F ABACEAE 231

FABACEAE
by Gerardo A. Aymard C., Nidia L. Cuello A., Paul E. Berry,
Velva E. Rudd, Richard S. Cowan, Paul R. Fantz,
Richard H. Maxwell, Charles H. Stirton,
Hans-Helmut Poppendieck, Haroldo Cavalcante de Lima,
Renée H. Fortunato, Basil Stergios, Nereida Xena de Enrich,
David A. Neill, R. Toby Pennington, and Celia Gil

Herbs, sometimes twining, shrubs, trees, or woody vines, rarely cauliflorous; roots
commonly bearing root nodules with nitrogen-fixing bacteria. Leaves alternate,
rarely opposite, pinnately or less often palmately compound or trifoliolate, some-
times unifoliolate or simple, the petiole and individual leaflets commonly with a
swollen basal pulvinus; stipules present, sometimes modified into spines or prickles.
Inflorescences mostly racemes, spikes, panicles, or heads. Flowers bisexual, usually
strongly irregular, mostly papilionaceous. Sepals mostly 5, generally connate into a
lobed, often bilabiate tube; petals usually 5, dissimilar [rarely 0–4 or 6, rarely 5 and
similar (Etaballia)], the uppermost (adaxial; in resupinate flowers the lowermost,
abaxial) petal the standard (sometimes called the vexillum and other parts associ-
ated with it are referred to by the adjective “vexillar”), borne externally to the other
petals, usually the largest and ± enclosing the others in bud, 2 lateral petals, the
wings, usually similar to each other and distinct, sometimes connivent basally, 2
lower petals innermost, similar to each other, and mostly connate to form a keel en-
closing the androecium and gynoecium. Stamens mostly 10, seldom 5–9 or 11, rarely
numerous, 9 of the filaments usually connate into an open sheath around the pistil,
the 10th, adaxial one, usually partly separated from the rest, or filaments rarely
free; anthers 4-sporangiate, 2-thecal, mostly opening longitudinally. Gynoecium a
single carpel or rarely 2 or several separate carpels, each with a terminal style and
stigma; ovules 1–many on a marginal placenta. Fruit commonly dry and dehiscent
along both sutures, but sometimes a follicle, or indehiscent and then sometimes
winged or breaking transversely into 1-seeded segments (loment), sometimes dru-
paceous, samaroid, or nut-like. Seeds with a short funiculus and usually a hard seed
coat; embryo with thickened cotyledons.
Cosmopolitan, ca. 400 genera and 10,000 species, 67 genera and 368 species in
the flora area.
This treatment follows the Cronquist system (A. Cronquist. An Integrated Sys-
tem of Classification of Flowering Plants. Columbia University Press. 1991), which
recognizes three different families among the legumes.

Key to the Genera of Fabaceae

by Paul E. Berry and Gerardo A. Aymard C.

1. Herbs, vines, small shrubs, or lianas ........................................................ 2

1. Large shrubs or trees ............................................................................... 38
2(1). Leaves digitately 2- or 4-foliolate; flowers enclosed by 2 bracts ... 67. Zornia
2. Leaves 1-foliolate, 3(4 or 5)-foliolate, or pinnately compound; flowers not
enclosed by 2 bracts, sometimes surrounded by several bracts ........... 3
232 F ABACEAE

3(2). Inflorescences spicate ................................................................................ 4

3. Inflorescences racemes, pseudoracemes, or solitary flowers .................... 5
4(3). Flowers surrounded by a series of floral bracts; stamens monadelphous;
fruit a loment ..................................................................... 58. Stylosanthes
4. Flowers not surrounded by a series of floral bracts; stamens diadelphous;
fruit not a loment .................................................................. 34. Indigofera
5(3). Leaves pinnately compound ...................................................................... 6
5. Leaves 1-foliolate, 3(4 or 5)-foliolate ....................................................... 18
6(5). Leaves even-pinnate .................................................................................. 7
6. Leaves odd-pinnate .................................................................................... 8
7(6). Vines; standard obovate; stamens monadelphous; seeds bicolored, orange
and black ......................................................................................... 1. Abrus
7. Herbs to small shrubs or trees; standard nearly round; stamens diadel-
phous; seeds brown ................................................................. 55. Sesbania
8(6). Mostly prostrate herbs; standard scarious, persistent; stamens in 2 lat-
eral bundles ..................................................................... 56. Soemmeringia
8. Vines, herbs, or small shrubs; standard usually deciduous; stamens not in
2 lateral bundles .................................................................................... 9
9(8). Herbs to small shrubs .............................................................................. 10
9. Woody or herbaceous vines ...................................................................... 13
10(9). Leaflets alternate or subopposite; fruit a loment ................ 3. Aeschynomene
10. Leaflets opposite; fruit not a loment ....................................................... 11
11(10). Pubescence of minutely hooked (view at 20–30×) trichomes; flowers resu-
pinate; ovary stipitate ................................................................ 17. Clitoria
11. Pubescence absent, or if present, never with minutely hooked trichomes;
flowers not resupinate; ovary sessile .................................................. 12
12(11). Pubescence of malpighiaceous (T-shaped) trichomes; anthers with an api-
cal projection ......................................................................... 34. Indigofera
12. Pubescence not of malpighiaceous trichomes; anthers without an apical
projection ................................................................................ 61. Tephrosia
13(9). Woody vines .............................................................................................. 14
13. Herbaceous vines ..................................................................................... 16
14(13). Pubescence of minutely hooked trichomes .................................... 17. Clitoria
14. Plants glabrous or pubescent, but never with minutely hooked trichomes
.............................................................................................................. 15
15(14). Fruits with distinct submarginal wings; pseudoracemes with 6–12-flow-
ered lateral short shoots .............................................................. 22. Derris
15. Fruits diverse, but never with distinct submarginal wings; pseudoracemes
either 6–12-flowered on lateral short shoots or with paired flowers on a
short common peduncle or flowers arising directly from the rachis and
appearing verticillate, paired, or single ......................... 36. Lonchocarpus
16(13). Flowers red; calyx long-tubular; style densely barbate, 1–1.5 cm long
................................................................................................... 8. Barbieria
16. Flowers yellow, blue, white, or purple; calyx cupular; style to 8 mm long
.............................................................................................................. 17
17(16). Stipules striate; calyx without tubercular-based trichomes; standard or-
bicular with a dorsal spur; stamens pseudodiadelphous; fruit mostly
linear, 2-valved, dehiscent .................................................. 14. Centrosema
 F ABACEAE 233

17. Stipules not striate; calyx with tubercular-based trichomes; standard obo-
vate to suborbicular without a dorsal spur; stamens monadelphous;
fruit a loment ......................................................................15. Chaetocalyx
18(5). Erect or prostrate herbs to small shrubs ................................................ 19
18. Vines or lianas .......................................................................................... 25
19(18). Fruit a loment .......................................................................................... 20
19. Fruit not a loment .................................................................................... 22
20(19). Leaves 4-foliolate; calyx glandular-punctate; stamens monadelphous
................................................................................................... 52. Poiretia
20. Leaves 1- or 3-foliolate; calyx not glandular-punctate; stamens diadelp-
hous ...................................................................................................... 21
21(20). Stipules scarious; calyx deeply divided, lobes unequal, the upper 2 gluma-
ceous; fruit a loment, terete to compressed, submoniliform, puberulent
............................................................................................... 6. Alysicarpus
21. Stipules not scarious; calyx subentire to bifid, the lower lobe 3-toothed
with the central tooth longer than the laterals, or the calyx almost
equally lobed; fruit a loment, not submoniliform, glabrous or with
hooked trichomes ................................................................ 23. Desmodium
22(19). Stamens monadelphous; anthers dimorphic; fruit inflated, not septate
............................................................................................... 19. Crotalaria
22. Stamens diadelphous; anthers uniform to subuniform; fruit not inflated,
septate .................................................................................................. 23
23(22). Lower surface of leaflets not gland-dotted; stipules setaceous; petioles not
canaliculate; flowers reddish purple or pink; anthers with an apical
projection ............................................................................... 34. Indigofera
23. Lower surface of leaflets gland-dotted; stipules not setaceous; petioles ca-
naliculate; flowers yellow; anthers without an apical projection....... 24
24(23). Shrubs; corolla yellow, lined with red; calyx not gland-dotted; standard
ovate; seeds 4–6 ........................................................................ 10. Cajanus
24. Perennial herbs or subshrubs; corolla yellow, sometimes with the stan-
dard veins reddish purple; calyx glandular; standard obovate; seeds
2 ............................................................................................... 27. Eriosema
25(18). Pubescence of minutely hooked trichomes (view at 20–30×) ....... 17. Clitoria
25. Plants glabrous or pubescent, if pubescent, never with minutely hooked
trichomes .............................................................................................. 26
26(25). Inflorescences nodose, 2–8 flowers at each node ..................................... 27
26. Inflorescences not nodose ........................................................................ 29
27(26). Calyx appearing bilabiate; fruits frequently with extra longitudinal ribs
............................................................................................... 12. Canavalia
27. Calyx 4- or 5-lobed; fruits without extra ribs .......................................... 28
28(27). Flowers reddish, the standard straight; stamens diadelphous; fruits with
long, down-curved beaks .................................................... 20. Cymbosema
28. Flowers shades of purple, occasionally white or lilac, the standards
mostly reflexed; stamens pseudomonadelphous; fruits short-beaked or
beak absent ................................................................................ 24. Dioclea
29(26). Leaflets and calyx gland-dotted ............................................. 54. Rhynchosia
29. Leaflets and calyx not gland-dotted ........................................................ 30
30(29). Calyx with irritating trichomes; anthers barbate; glandular disk present;
234 F ABACEAE

fruit densely covered with stiff irritating trichomes ............... 40. Mucuna
30. Calyx without irritating trichomes; anthers glabrous; glandular disk ab-
sent; fruit without stiff irritating trichomes ....................................... 31
31(30). Stamens monadelphous or pseudomonadelphous .................................. 32
31. Stamens diadelphous ............................................................................... 33
32(31). Twining herbs; inflorescences not nodose, bracteoles subtending the ca-
lyx; calyx 5-lobed; stamens pseudomonadelphous; fruit with a hooked
tip ........................................................................................... 62. Teramnus
32. Climbing or erect subshrubs; inflorescences nodose; bracteoles present
but not subtending the flowers; calyx 4-lobed; stamens monadelphous;
fruit without hooked tip ........................................................... 31. Galactia
33(31). Style glabrous ........................................................................................... 34
33. Style pubescent ........................................................................................ 35
34(33). Leaflets somewhat rhombic; calyx 5-lobed; standard with a pair of in-
flexed lateral auricles at base; fruit transversely grooved; anthers
8 or 9 ................................................................................ 11. Calopogonium
34. Leaflets linear or lanceolate; calyx 4-lobed; standard without a pair of
inflexed lateral auricles at base; fruit not transversely grooved;
anthers 10 ................................................................................ 31. Galactia
35(33). Roots tuberous; fruit articulate ............................................. 46. Pachyrhizus
35. Roots not tuberous; fruit not articulate .................................................. 36
36(35). Pubescence of uncinate trichomes; style coiled through 1.5–2.5 revolu-
tions; bracteoles persistent in fruit ....................................... 47. Phaseolus
36. Uncinate trichomes absent; style not coiled, or, if coiled, then through
3–5 revolutions; bracteoles caducous .................................................. 37
37(36). Inflorescence nodes not glandular; upper calyx teeth free; corollas pur-
plish upon drying, with elongated wing petals; thickened part of style
bent sharply ..................................................................... 38. Macroptilium
37. Inflorescence nodes glandular; upper calyx 2-lipped; corollas yellowish or
white, sometimes with some purple or violet upon drying, with wing
petals equal to the others and with erect, sigmoid-shaped, or coiled keel
petals; thickened part of style gradually bent or coiled .............. 65. Vigna
38(1). Leaves simple or unifoliolate ................................................................... 39
38. Leaves compound ..................................................................................... 45
39(38). Leaf margin serrate; fruits indehiscent .................................................. 40
39. Leaf margin entire; fruits dehiscent or indehiscent ............................... 41
40(39). Fruit globose to ovoid or broadly turbinate; seeds 1 or 2 .......... 35. Lecointea
40. Fruit ellipsoid, laterally compressed, stipitate, apiculate; mesocarp fleshy;
seeds 1–few ............................................................................. 66. Zollernia
41(39). Inflorescences axillary spikes; petals linear .............................. 29. Etaballia
41. Inflorescences racemes or panicles; petals not linear ............................. 42
42(41). Calyx entire and completely enclosing the flower in bud, opening irregu-
larly or in distinct lobes; flowers not papilionaceous, petals 0–6; sta-
mens numerous (> 15) ......................................................................... 43
42. Calyx lobes apparent in bud (entire and enclosing the bud in Alexa);
flowers papilionaceous or not, petals generally 5; stamens generally
< 15 ....................................................................................................... 44
43(42). Petals 3–6; fruit a single-seeded drupe ............................................ 4. Aldina
43. Petals 0 or 1; fruit a 1- to multiseeded legume .......................... 59. Swartzia
 F ABACEAE 235

44(42). Anthers not versatile; fruits 1-seeded ....................................... 21. Dalbergia

44. Anthers versatile; fruits 2- or 3-seeded .................................. 51. Poecilanthe
45(38). Leaves opposite ........................................................................................ 46
45. Leaves alternate ....................................................................................... 47
46(45). Leaf rachis terete; inflorescence axillary; calyx tubular, not pellucid-punc-
tate; petals yellow ............................................................ 49. Platymiscium
46. Leaf rachis usually winged; inflorescence terminal; calyx pellucid-punc-
tate, bilabiate; petals violet ....................................................... 60. Taralea
47(45). Stamens free or nearly so ........................................................................ 48
47. Stamens connate to a considerable extent .............................................. 60
48(47). Calyx entire and completely enclosing the flower in bud, opening irregu-
larly or in distinct lobes; flowers not papilionaceous, petals 0–6; sta-
mens numerous (> 15) ......................................................................... 49
48. Calyx lobes apparent in bud (but entire and enclosing the bud in Alexa);
flowers papilionaceous or not, petals generally 5; stamens generally
fewer ..................................................................................................... 50
49(48). Petals 3–6; fruit a single-seeded drupe, less often 2–4-seeded ....... 4. Aldina
49. Petals 0 or 1; fruit a multi-seeded legume ................................. 59. Swartzia
50(48). Flowers ± regular, the adaxial petal a little wider than the others; fruit
compressed, indehiscent, round to oblong, transversely rugulose
................................................................................................. 2. Acosmium
50. Flowers zygomorphic or subactinomorphic (Myrocarpus); fruits various,
but often either winged or dehiscent .................................................. 51
51(50). Fruit 1- or 2-seeded; leaves pellucid-punctate ........................................ 52
51. Fruit 1- or more-seeded; leaves mostly not pellucid-punctate ............... 54
52(50). Fruit narrowly elliptic or fusiform, the seed(s) flat and situated in the
middle of the winged fruit .................................................. 42. Myrocarpus
52. Fruit with a bulbous apical seed chamber and a large basal wing ........ 53
53(52). Leaflets 14–25; anthers shorter than the filaments; plants native in the
flora area ......................................................................... 43. Myrospermum
53. Leaflets 5–15; anthers longer than the filaments; plants mostly cultivated
in the flora area .................................................................... 44. Myroxylon
54(51). Calyx large, enclosing the bud; ovary long-stipitate; inflorescence a termi-
nal, woody raceme .......................................................................... 5. Alexa
54. Calyx various, not completely enclosing the bud; ovary sessile or short-
stipitate; inflorescence axillary or terminal, generally not woody .... 55
55(54). Calyx 2-lobed, the upper lobe enlarged and covering the petals; petals
pink; fruit flattened, 1-seeded, tapering at both ends, elastically dehis-
cent ...................................................................................... 39. Monopteryx
55. Calyx without a large lobe covering the petals in bud; petals mostly blue
or purple, less often white with darker nectar spot; fruit not as above
.............................................................................................................. 56
56(55). Fruits indehiscent .................................................................................... 57
56. Fruits dehiscent ....................................................................................... 58
57(56). Fruits linear-oblong to elliptic, membranous, upper suture shortly winged
................................................................................................. 9. Bowdichia
57. Fruits oblong, tapering at both ends, apex acute, not winged ...................
.............................................................................................. 57. Spirotropis
58(56). Stigma lateral; seeds reddish, hard, shiny .................................. 45. Ormosia
236 F ABACEAE

58. Stigma terminal; seeds not as above ....................................................... 59

59(58). Standard without lateral appendages; fruit with 1 or 2 seeds ...................
.............................................................................................. 25. Diplotropis
59. Standard with 2 lateral appendages; fruit with more than 2 seeds ..........
.......................................................................................... 16. Clathrotropis
60(47). Leaves 3-foliolate ..................................................................................... 61
60. Leaves pinnately compound with more than 3 leaflets .......................... 62
61(60). Flowers large, > 15 mm long; fruit dehiscent; seeds hard, red ..................
................................................................................................ 28. Erythrina
61. Flowers small, < 15 mm long; fruit indehiscent, drupaceous; seeds soft,
not red ........................................................................................... 7. Andira
62(60). Lower surface of leaflets gland-dotted; flowers yellow ................ 10. Cajanus
62. Lower surface of leaflets not gland-dotted; flowers red, orange, rose, vio-
let, or white .......................................................................................... 63
63(62). Fruits drupaceous .................................................................................... 64
63. Fruits nondrupaceous, dry at maturity, dehiscent or indehiscent ......... 65
64(63). Leaf rachis terete; calyx not pellucid-punctate, truncate or with short
obsolete teeth or shortly 5-dentate; petals pink, violet, or white .....
....................................................................................................... 7. Andira
64. Leaf rachis flattened or winged; calyx pellucid-punctate, bilabiate, upper
lip with 2 large lobes and lower lip very short, entire, or 3-dentate; pet-
als violet to whitish mauve ...................................................... 26. Dipteryx
65(63). Leaflets opposite on the rachis ................................................................ 66
65. Leaflets alternate or subopposite on the rachis ...................................... 73
66(65). Fruit with basal seed and apical wing, the basal part spinose with a stiff
lateral spur; leaves with numerous reddish orange resinous dots; pet-
als yellowish ..................................................................... 13. Centrolobium
66. Fruit without a basal spinose portion and lateral spur; leaves without
resin dots; petal color various .............................................................. 67
67(66). Fruit flat, ovate-elliptic, with broad lateral wings; calyx spathaceous; pet-
als yellow to orange; anthers opening by 2 apical pores ...... 30. Fissicalyx
67. Fruit not as above; calyx not spathaceous; petal color various; anthers
longitudinally dehiscent ...................................................................... 68
68(67). Fruit longitudinally 4-winged ....................................................... 48. Piscidia
68. Fruit not 4-winged ................................................................................... 69
69(68). Fruits long and narrow, either cylindrical and constricted between the
seeds or somewhat flattened and dehiscent ....................................... 70
69. Fruits thin and flattened, rounded to oblong, mostly indehiscent ......... 71
70(69). Fruit cylindrical, indehiscent, constricted between the seeds ... 41. Muellera
70. Fruit dehiscent, flattened, often wider toward the apex, very little con-
stricted between the seeds, the 2 valves hard and often coiling in dehis-
cence ....................................................................................... 32. Gliricidia
71(69). Fruit 1-seeded, elastically dehiscent, 2-valved, flat; calyx pellucid-punc-
tate, bilabiate ............................................................................. 60. Taralea
71. Fruit 1- or more-seeded, not elastically dehiscent; calyx not pellucid-punc-
tate ....................................................................................................... 72
72(71). Fruit with 2 subparallel prominent veins near the margin; seeds 1(2); in-
florescence a terminal panicle; vexillar stamen connate to staminal
tube ................................................................................ 33. Hymenolobium
 Abrus 237

72. Fruit without subparallel marginal veins; seeds 1–12; inflorescence a lat-
eral or terminal raceme or panicle; vexillar stamen free at base
......................................................................................... 36. Lonchocarpus
73(65). Fruits long and narrow, many-seeded, without appendages; leaves even-
pinnate .................................................................................... 18. Coursetia
73. Fruits rounded to oblong, but not long and narrow, 1–few-seeded, some-
times with appendages; leaves odd-pinnate or less often even-pinnate
.............................................................................................................. 74
74(73). Fruit with a central seed and surrounding wing (see also Dalbergia)
............................................................................................. 53. Pterocarpus
74. Fruit not as above .................................................................................... 75
75(74). Leaves even-pinnate, the rachis often flattened and ± winged; calyx pel-
lucid-punctate, bilabiate; fruit with dehiscent twisting woody valves
.................................................................................................... 60. Taralea
75. Leaves odd-pinnate, the rachis terete; calyx not pellucid-punctate or bila-
biate; fruit generally indehiscent, often 1-winged with an apical or
basal seed ............................................................................................. 76
76(75). Fruit with an apical seed and basal wing ............................ 50. Platypodium
76. Fruit various, but not as above ................................................................ 77
77(76). Flowers in pyramidal, usually terminal panicles; fruits with a basal seed
and distal wing or turgid and spongy-fibrous ..................................... 78
77. Flowers in subcorymbose, usually axillary panicles; fruits winged by flat-
tened valves, seeds either basal or central ......................................... 79
78(77). Fruit with 2 small, basal, lateral wings ................................ 64. Vataireopsis
78. Fruit without basal wings, apically winged or wingless with spongy-fi-
brous mesocarp ......................................................................... 63. Vatairea
79(77). Seeds basally positioned in the fruit; anthers longitudinally dehiscent;
stipules sometimes spinescent ......................................... 37. Machaerium
79. Seed(s) medially positioned in the fruit; anthers with short, transverse
slits; stipules never spinescent .............................................. 21. Dalbergia

1. ABRUS L., Syst. Nat. ed. 12, 2: 472. 1767.

by Nidia L. Cuello A.

Mostly twining vines, seldom shrubs. Leaves alternate, even-pinnate; rachis

grooved; leaflets 11–27, oblong to ovate, mucronate, sparingly to densely pubescent
with ascending trichomes, opposite along the rachis; stipels minute, appressed to
the rachis; stipules subulate, persistent. Inflorescences axillary or lateral
pseudoracemes, terminating leafy or leafless branches, with 1–several sessile flow-
ers at a node; bracts and bracteoles often caducous. Flowers white, reddish brown,
lavender, or pinkish. Calyx campanulate, lobed or short-toothed, sparingly short-
pubescent; standard obovate, apically notched and with a short claw; wing petals
upcurved, shorter than keel petals. Stamens 9, not exserted, monadelphous, the
staminal tube adnate to the standard. Style curved, glabrate. Fruit oblong to linear,
somewhat oblique, hardly stipitate, much-compressed or not, the valves softly pu-
bescent, often becoming indurated, septate, the short, slender beak downturned.
Seeds subglobose or slightly compressed-ovoid, arillate, sometimes with an annulus
around the hilum, the hilum depressed.
238 F ABACEAE

Fig. 213. Abrus precatorius subsp. africanus

Fig. 214. Abrus pulchellus subsp. tenuiflorus

 Acosmium 239

Paleotropics, now widely introduced and circumtropical; 17 species, 2 in Ven-

ezuela, both in the flora area.
The brightly colored orange-red and black seeds are highly toxic, but are some-
times used as an adornment or in artwork by Panare Amerindians.

Key to the Species of Abrus

1. Leaflets oblong-ovate, 3–8 mm wide; flowers in pedunculate racemes;

fruits usually thick and woody, the surface muriculate, densely ap-
pressed-pubescent, ca. 2 times as long as broad; seeds subglobose,
shiny scarlet with black ........................................................ A. precatorius
1. Leaflets oblong-obovate, 7–13 mm wide; flowers sessile in small clusters;
fruits mostly not woody, the surface smooth and sparsely pubescent, ca.
3 times as long as broad; seeds compressed, shiny brown .... A. pulchellus

Abrus precatorius L., Syst. Nat. ed. 12, 2:
472. 1767.
Paleotropics, introduced to the Neo-
tropics; 2 subspecies, 1 in Venezuela.
A. precatorius subsp. africanus Verdc.,
Kew Bull. 24: 241. 1970. —Peonía.
Abrus minor Desv., Ann. Sci. Nat. (Paris)
9: 418. 1826.
Woody vine over shrubs. Disturbed or cul-
tivated areas, near sea level to 50 m; Delta
Amacuro (Santa Catalina), Bolívar (Caicara,
Ciudad Bolívar). Anzoátegui, Aragua, Bari-
nas, Falcón, Guárico, Miranda, Monagas,
Nueva Esparta, Trujillo; U.S.A. (southern
Florida), Mexico, Guatemala, Belize, Hondu-
ras, Costa Rica, Panama, West Indies, Co-
lombia, Guyana, Peru, Brazil (Bahia), Africa,
Asia, Oceania. ŠFig. 213.
Abrus pulchellus Wall. ex Thwaites,
Enum. Pl. Zeyl. 91. 1864 [1859].
Paleotropics, introduced to Neotropics; 5
subspecies, 1 in Venezuela.
A. pulchellus subsp. tenuiflorus (Benth.)
Verdc., Kew Bull. 24: 250. 1970. —Abrus
tenuiflorus Benth. in Mart., Fl. Bras.
15(1): 216. 1859. —Arepillo, Bejuco
arepillo.
Vine over shrubs. Forest borders, 100–
1000 m; Bolívar (near Santa Elena de
Uairén), Amazonas (near confluence of Río
Negro and Brazo Casiquiare). Táchira; Bra-
zil (Amazonas, Goiás, Mato Grosso), Africa,
Sri Lanka. ŠFig. 214.

2. ACOSMIUM Schott. in Spreng., Syst. Veg. 4(cur. post.): 406. 1827.

by Charles H. Stirton and Gerardo A. Aymard C.
Large shrubs or trees. Stipules small, caducous. Leaves 3–31-foliolate, odd-pin-
nate, alternate. Leaflets membranaceous or coriaceous, often discolorous, opposite
on the rachis. Inflorescences paniculate, terminal, rarely axillary, racemes short to
very elongate. Flowers white, cream, or yellow, scarcely zygomorphic, pedicellate,
bracteate and bracteolate, with axillary clusters of stalked glands. Calyx turbinate-
campanulate, upper pair of teeth much larger and broader than lateral and keel
teeth; petals free, erect-ascending, uppermost petal outermost, differently shaped
than rest. Stamens free, 10 (rarely 5), subequal, usually longer than petals; anthers
uniform, ovate. Ovary sessile or shortly stipitate, 2–4-ovulate, glabrous or shaggy;
style filiform; stigma small or truncate, terminal. Legume indehiscent, compressed
and flat or turgid, 1–4-seeded. Seeds ovate or orbicular, compressed.
Central America, Colombia, Venezuela, Guyana, Suriname, Brazil, Bolivia,
Paraguay, Argentina, Madagascar; 17 species, 3 in Venezuela, 2 of these in the flora
area.
240 F ABACEAE

Key to the Species of Acosmium

1. Leaves 5–9-foliolate; leaflets ovate to broadly lanceolate, subcoriaceous,

not prominently reticulate on the upper surface; fruit sessile or
subsessile, turgid, woody; ovary glabrous ..................................... A. nitens
1. Leaves 5- or 7-foliolate; leaflets ovate to ovate-elliptic, chartaceous to sub-
coriaceous, reticulate on the upper surface; fruit distinctly stipitate,
plano-compressed, papery, winged; ovary pubescent ..................... A. sp. A

Acosmium nitens (Vogel) Yakovlev, Notes Acosmium sp. A. —Congrio, Inchapemen

Roy. Bot. Gard. Edinburgh 29: 353. 1969.
—Leptolobium nitens Vogel, Linnaea 11:
394. 1837. —Sweetia nitens (Vogel)
Benth., J. Linn. Soc., Bot. 8: 262. 1865.
—Chimaco, Congrio, Congrio rebalsero.
Small tree to 10 m tall; leaves 5–9-
foliolate; flowers cream. Semideciduous to
evergreen lowland forests along rivers and
adjacent slopes, savannas, riparian shrub-
land along black-water rivers, 50–200 m;
northern Bolívar, widespread in Amazonas.
Apure, Guárico; Colombia, Guyana, Suri-
name, Brazil (Amazonas, Para, Mato Grosso,
Rondônia), Bolivia. ŠFig. 215.
(Panare), Karamate yo.
Slender, spreading tree to 20 m tall,
branched from low down on the trunk; leaves
5- or 7-foliolate; flowers white; fruit winged.
Savannas, savanna/forest ecotone, semide-
ciduous forests, granitic outcrops, 50–300 m;
Bolívar (Ciudad Piar, Corozal, east of
Miamo, Morichal El Caballo, San Félix to
Upata road, San Pedro de los Bocas, islands
in Lago Guri), Amazonas (10 km south of
Puerto Rico). Guyana, Brazil (Roraima).

Fig. 215. Acosmium nitens

 Aeschynomene 241

3. AESCHYNOMENE L., Sp. Pl. 713. 1753.

by Velva E. Rudd
Herbaceous annuals, shrubs, or small trees. Leaves alternate, pinnate, 5–
many-foliolate; stipules usually persistent, peltate, appendiculate below the point of
attachment or attached at the base and not appendiculate; stipels lacking; leaflets
alternate or subopposite, usually small, entire to serrate-denticulate. Inflorescences
axillary or terminal, simple or compound racemes, sometimes fasciculate; bracts
mostly similar to and intergrading with the stipules; bracteoles paired at the base of
the calyx. Flowers ca. 5–25 mm long. Calyx bilabiate or campanulate with 5
subequal lobes; petals 5, white, yellow, orange, or purplish, glabrous or pubescent.
Stamens 10, monadelphous or diadelphous (5 + 5); anthers uniform, elliptic,
dorsifixed, or, sometimes nearly basifixed. Ovary (1)2–18-ovulate, sessile or short-
stipitate; style glabrous with base usually persisting as a cuspidate apex to the fruit;
stigma terminal, minutely capitate or penicillate. Fruit a loment, (1)2–18-articulate,
commonly with the upper (adaxial) margin essentially entire, the lower margin
crenate, sometimes both margins crenate or both subentire. Seeds reniform, smooth,
lustrous, the hilum orbicular.
Mainly tropical or subtropical America, Africa, Asia, and Australia, with a few
species in warm-temperate areas; 150 species, 15 in Venezuela, 10 of these in the
flora area.

Key to the Species of Aeschynomene

1. Stipules attached at the base, not peltate; calyx campanulate with 5 sub-
equal lobes; standard pubescent on the outer face ............................... 2
1. Stipules peltate, appendiculate below the point of attachment; calyx bi-
labiate with the vexillar lip bifid, the keel lobe trifid; petals glabrous ... 4
2(1). Stems suffrutescent, erect; inflorescence terminal, paniculate, or some
axillary and racemose; fruit moniliform, 4–6-articulate; stipe 4–5 mm
long, glabrous ......................................................................... A. paniculata
2. Stems herbaceous, prostrate or decumbent; inflorescences axillary, race-
mose; fruit with upper margin straight, the lower margin crenate with
constriction; stipe < 4 mm long, usually hispid .................................... 3
3(2). Leaves (5–)8–15(–20)-foliolate; leaflets obovate to obovate-oblong, (2–)
3–8 mm wide; fruit 3(4)-articulate; articles 2.5–5 × 2–4 mm; stipe
3–4 mm long ............................................................................ A. brasiliana
3. Leaves (11–)15–30-foliolate; leaflets oblong-elliptic, 1–2 mm wide; fruit
2-articulate; articles 2–2.5 × 2–2.5 mm; stipe 1–2 mm long ....... A. histrix
4(1). Leaflets asymmetrical, appearing to be 2–several-costate ....... A. americana
4. Leaflets asymmetrical or not, 1-costate .................................................... 5
5(4). Costa of leaflet excentric; fruit hispid and tuberculate ........... A. fluminensis
5. Costa of leaflets essentially central; fruit glabrous or pubescent, the sur-
face variously raised but not tuberculate ............................................. 6
6(5). Plants in general, especially in fruits, blackening on drying; calyx lips en-
tire or subentire ..................................................................................... 7
6. Plants, including fruits, turning brownish to straw-colored or remaining
green on drying; calyx lips clearly lobed or toothed ............................. 8
242 F ABACEAE

7(6). Fruit submoniliform at maturity, stipe and basal article separated by a

suture; stipe 8–15 mm long; flowers 7–12 mm long ................ A. pratensis
7. Fruit with one margin entire, the other crenate, stipe and basal article
continuous, not separated by a suture; stipe 4–8 mm long; flowers 4–9
mm long ..................................................................................... A. sensitiva
8(6). Flowers (8–)10–15 mm long, standard reflexed; fruit 4–6 mm wide; stipe
3–6(–10) mm long; leaflets entire or with a slight tendency toward mar-
ginal trichomes ............................................................................... A. rudis
8. Flowers (4–)5–12 mm long; standard essentially erect; fruit 2.5–4 mm
wide; stipe 3–15 mm long; leaflets serrulate-ciliate to entire .............. 9
9(8). Stipe of fruit 3–4(–6) mm long; flowers (4–)5–7(–9) mm long; calyx 4–5 mm
long ................................................................................................ A. evenia
9. Stipe of fruit (5–)10–15 mm long; flowers 8–12 mm long; calyx 6–9 mm
long ................................................................................................ A. scabra

Aeschynomene americana L., Sp. Pl. 715.
1753. —Cujicillo.
Mexico, Central America, West Indies,
South America, introduced in Africa and
Asia; 2 varieties, 1 in Venezuela.
A. americana var. americana
Herb, somewhat weedy, usually erect, to
ca. 2.5 m tall. Wet or moist roadsides,
ditches, fields, forest edges, near sea level to
500 m; Delta Amacuro (Caño Manamo near
Tucupita), Bolívar (Canaima). Northern and
western Venezuela; Mexico, Central Amer-
ica, West Indies, widespread in South
America, introduced in Africa and Asia.
Aeschynomene brasiliana (Poir.) DC.,
Prodr. 2: 322. 1825. —Hedysarum
brasilianum Poir. in Lam., Encycl. 6:
448. 1804 [1805].
Aeschynomene guarica Pittier, Bol. Técn.
Minist. Agric. 5: 41. 1844.
Prostrate or decumbent herb. Wet road-
sides, savannas, scrub. Mexico, Central
America, Cuba, Colombia, Venezuela,
Trinidad, Suriname, French Guiana, Ecua-
dor, Peru, Brazil, Bolivia; 3 varieties, all in
Venezuela, 2 of these in the flora area.
Key to the Varieties of A. brasiliana
1. Flowers (4–)5–8 mm long; segments of fruit
2.5–3 × 2–3 mm ......... var. brasiliana
1. Flowers ca. 10 mm long; segments of fruit
4–5 × 3–4 mm .......... var. carichanica
elsewhere in Venezuela; other distribution
as in species.
A. brasiliana var. carichanica Rudd, J.
Wash. Acad. Sci. 49: 48. 1959.
200–400 m; Bolívar (between Caicara and
Maniapure, Cerro Carichana along Río
Orinoco, Cerro El Médano 23 km southwest
of Caicara). Endemic. ŠFig. 220.
Aeschynomene evenia C. Wright in
Sauvalle, Anales Acad. Ci. Méd. Habana
5: 334. 1868.
West Indies, western Colombia, Venezu-
ela, Trinidad, eastern Brazil; 2 varieties,
both in Venezuela, 1 of these in the flora
area.
A. evenia var. serrulata Rudd, Contr. U.S.
Natl. Herb. 32: 61. 1955. —Dormidera,
Gusana.
Erect herb, often suffrutescent, to ca. 50
cm tall; leaflets serrulate-ciliate. Moist or
wet places, near sea level to 100 m; Delta
Amacuro (Capure), Bolívar (Ciudad Bolívar),
Amazonas (Maroa, San Carlos de Río Negro).
Aragua, Barinas, Guárico, Monagas; West
Indies, western Colombia, Trinidad, eastern
Brazil. ŠFig. 216.
Aeschynomene fluminensis Vell., Fl.
Flumin. 310. 1825 [1829].
Cuba, Hispaniola, Dominican Republic,
Colombia, Venezuela, Brazil, Bolivia, Para-
guay; 2 varieties, 1 in Venezuela.

A. brasiliana var. brasiliana A. fluminensis var. fluminensis

50–200 m; northeastern Bolívar, Ama- Suffrutescent, shrubby herb; stems erect
zonas (Culebra, Isla Ratón). Widespread to ca. 4 m tall. In water, wet savannas, river

banks, beaches, 50–300 m; eastern Bolívar
(between Caicara and Río Cuchivero, be-
tween Tumeremo and El Dorado). Cuba,
Dominican Republic, Brazil, Bolivia, Para-
guay.
Aeschynomene histrix Poir. in Lam.,
Encycl. suppl. 4: 77. 1816.
Mexico, Guatemala, Honduras, El Salva-
dor, Nicaragua, Colombia, Venezuela, Guy-
ana, Suriname, French Guiana, Peru, Brazil,
Bolivia, Paraguay, Argentina, Uruguay; 5 va-
rieties, 3 in Venezuela, 1 of these in the flora
area.
A. histrix var. histrix
Herb, commonly prostrate, sometimes
erect; stems to ca. 1 m long. Disturbed areas,
savannas, forest edges, 50–300 m; Delta
Amacuro (between Piacoa and Castillos de
Guayana), northern Bolívar, Amazonas
(around and north of Puerto Ayacucho).
Anzoátegui, Guárico; Guatemala, Honduras,
El Salvador, Nicaragua, Colombia, Guyana,
Suriname, French Guiana, Peru, Brazil,
Paraguay. ŠFig. 217.
Some specimens trend toward var. incana
(Vogel) Benth., and some have been previ-
ously cited as var. incana, but in the strict
sense, var. incana occurs in southern South
America, particularly in Argentina and Uru-
guay.
Fig. 216. Aeschynomene evenia var. serrulata
Aeschynomene 243
Aeschynomene paniculata Willd. ex
Vogel, Linnaea 12: 95. 1838.
Suffrutescent, erect herb to ca. 2.5 m tall.
Rocky slopes, savannas, open woods, road-
sides, 100–500 m; northern Bolívar, Ama-
zonas (Puerto Ayacucho). Anzoátegui, Ara-
gua, Trujillo; widespread in Mexico, Central
America, Colombia, Guyana, Suriname, Bra-
zil, Bolivia.
Aeschynomene pratensis Small, Bull.
New York Bot. Gard. 3: 423. 1905.
Nicaragua, Costa Rica, Panama, Cuba,
Hispaniola, Colombia, northern Brazil, Bo-
livia; 2 varieties, 1 in Venezuela.
A. pratensis var. caribaea Rudd, Contr.
U.S. Natl. Herb. 32: 47. 1955.
Erect herb, sometimes suffrutescent;
flowers 7–8 mm long; calyx 4–5 mm long;
fruit with articles 5–6 × 4–5 mm; stipe 8–10
mm long. Marshy areas, river banks, 100–
800 m; Bolívar (La Paragua, Santa Elena de
Uairén). Anzoátegui, Cojedes, Trujillo; Nica-
ragua, Costa Rica, Panama, Cuba, His-
paniola, Colombia, Guyana, northern Brazil,
Bolivia. ŠFig. 219.
Aeschynomene rudis Benth., Pl. Hartw.
116. 1839 [1843].
Erect herb to ca. 2 m tall. Moist or wet
places, river banks or in water, ca. 50 m;
Fig. 217. Aeschynomene
histrix var. histrix
244 F ABACEAE

Fig. 218. Aeschynomene scabra

Fig. 220. Aeschynomene brasiliana

Fig. 219. Aeschynomene pratensis var. caribaea var. carichanica
 Aldina 245

Bolívar (Ciudad Bolívar). Guárico; widely
distributed from southern U.S.A., Mexico,
Central America, Cuba, Colombia, Brazil,
Bolivia, to Argentina.
Aeschynomene scabra G. Don, Gen. Hist.
2: 284. 1832.
Herb, sometimes suffrutescent, to ca. 3 m
tall. Swamps, humid savannas, 50–300 m;
Bolívar (Ciudad Bolívar, Río Orinoco be-
tween Río Horeda and Cerro Carichana).
Mexico, Central America, Ecuador, Peru,
Brazil, Bolivia, Paraguay. ŠFig. 218.
The two collections from the flora area
(Wurdack & Monachino 39868, F, NY, US;
Wurdack & Monachino 39941, NY, US, VEN)
were previously referred to Aeschynomene
rostrata Benth., a species known only from
Bahia, Brazil. The leaflets on specimens from
the flora area have more serrulate-ciliate mar-
gins than most collections, and thus A. scabra
seems a better determination.
Aeschynomene sensitiva Sw., Prodr. 107.
1788.
Southern Mexico, Central America, West
Indies, Colombia, Ecuador, Peru, Brazil, Bo-
livia, Paraguay, Argentina; 3 varieties, 1 in
Venezuela.
A. sensitiva var. sensitiva
Erect, suffrutescent herb 1–2(–4) m tall.
In water or on wet river banks and mud flats,
near sea level to 100 m; Delta Amacuro
(Caño Araguao, Caño Güiniquina, Sacupana,
between Tucupita and Los Güires), Bolívar
(Ciudad Bolívar, south of Tumeremo).
Aragua, Carabobo, Distrito Federal, Falcón,
Miranda, Táchira; other distribution as in
species.

4. ALDINA Endl., Gen. Pl. 1322. 1840, nom. cons.

Allania Benth., J. Bot. (Hooker) 2: 91. 1840.
by Basil Stergios and Richard S. Cowan
Trees. Leaves odd-pinnate or the upper sometimes 1-foliolate, exstipulate; leaf-
lets opposite to subopposite, blades mostly coriaceous to rigid with rounded to cor-
date base. Inflorescences terminal or axillary to leaves nearest end of the branch,
racemose or paniculate-racemose; bracts and bracteoles minute; pedicels short, 2-
bracteolate apically. Calyx entire in bud, opening about halfway in 2–5 recurved or
reflexed segments, tube turbinate; petals (3)4–6, elliptic to oblanceolate, obovate or
oblong, usually concave to cucullate, white to yellowish, glabrous. Stamens numer-
ous, glabrous; filaments borne on margin of short hypanthial cup adnate to calyx
base; anthers large, uniform, narrowly oblong to linear, opening lengthwise by slits.
Gynophore arising from the bottom of hypanthium cup, swollen-articulate at junc-
tion with the incurved, somewhat oblong ovary; ovary usually 1, but potentially 2 or
3; style subulate, straight or usually apically curved; stigma punctiform. Fruit to 10
cm long, dehiscent, thick, woody, ovoid, and 1-seeded or else oblong-fusiform and 2–
4-seeded.
Colombia, Venezuela, Guyana, northwestern Brazil; 22 species, 18 in Venezu-
ela, all in the flora area.
The genus is currently under revision by the senior author. The treatment here
of the published taxa is largely an extract from an earlier, preliminary treatment.

Key to the Species of Aldina

1. Leaflets completely glabrous on all surfaces; racemes simple or multiple

................................................................................................................ 2
1. Leaflets pubescent on lower surface, sometimes minutely so; inflorescence
an elongated, open, multiple raceme or terminal panicle of racemes ..... 8
2(1). Flowers showy, calyx tube 2.5–3 cm, petals 3–4 × 2 cm; ovaries 2 or 3 on
the stipe; fruits elongated, sausage-shaped to thickly fusiform and no-
246 FABACEAE

ticeably constricted between the 2–4 seeds; leaflets thickly coriaceous,

almost cartilaginous, noticeably cordate at the base ..................... A. sp. A
2. Flowers half again as large or smaller than above; ovary solitary on stipe;
fruits ovoid to oblong, 1- or 2-seeded; leaflets chartaceous to ± coria-
ceous, but not cordate at the base ......................................................... 3
3(2). Leaflets pallid and waxy on lower surface, broadly lanceolate to acumi-
nate, 10–11 × 4–4.5 cm; calyx and unopened buds very sparsely white-
appressed-puberulent near the base or usually glabrous; calyx tube
truncate, indentedly barrel-shaped at the base; pedicels 11–12 mm
long; bracteoles 3–3.5 mm below base of calyx tube ....................... A. sp. B
3. Leaflets shiny on lower surface, without a waxy coating, oblong to ovate-
elliptic, obtuse-mucronate, (8–)9–14 × 4.5–6 cm; calyx and unopened
buds uniformly and usually densely pubescent; calyx tube truncate,
claviform at the base; pedicels 1–3 mm long; bracteoles 0–1 mm below
base of calyx tube ................................................................................... 4
4(3). Leaflets noticeably tessellate with dark red lines on lower surface; ma-
ture buds 15–17 × 8 mm; racemes simple and axillary or occasionally
paired .......................................................................................... A. latifolia
4. Leaflets not tessellate, smooth and homogeneous on lower surface; ma-
ture buds 9–11 × 5–7 mm; racemes multiple-branched or in small
panicles ................................................................................................... 5
5(4). Leaflets glaucous or with an ash-gray-green bloom on lower surface;
ovary mostly glabrous .................................................................... A. berryi
5. Leaflets entirely shiny on both surfaces, not at all discolorous; ovary pu-
bescent .................................................................................................... 6
6(5). Leaves 1- or 3-foliolate ............................................................ A. heterophylla
6. Leaves 5- or 7-foliolate ............................................................................... 7
7(6). Leaflets coriaceous, 7–8 × 5–6 cm, elliptic, mucronate with no extended
acumen; inflorescence and flower buds ferruginous-velutinous, with a
golden brown lustre ......................................................................... A. sp. C
7. Leaflets chartaceous, 11–13 × 4.5–5.5 cm, lanceolate to obtuse, acumi-
nate, the acumen 1–1.5 cm long; inflorescence and flower buds tan-
brown, appressed-pubescent .......................................................... A. sp. G
8(1). Ovary glabrous or nearly so ....................................................................... 9
8. Ovary either densely and uniformly pubescent, or densely golden-brown-
puberulent on lower, dorsal portion only, and either ventral suture or
both sutures glabrous .......................................................................... 10
9(8). Pistil as long as or longer than the stamen filaments at anthesis; leaflets
pruinulose-lepidote on lower surface, with semierect or reclining tri-
chomes; petals 5, 18–20 × 6–6.5 mm .............................................. A. sp. D
9. Pistil noticeably shorter than the stamen filaments at anthesis; leaflets
appressed-puberulent, subglaucous with a uniform bloom on lower sur-
face; petals 4, 20–25 × ca. 10 mm ....................................... A. macrophylla
10(8). Leaflets rich erect-brown-villous on lower surface; racemes and flower
buds notably dark, golden-brown-velutinous; calyx 2-lobed at anthesis
........................................................................................... A. kunhardtiana
10. Leaflets appressed-puberulent, either pale lilac or with a pruinose, waxy
bloom on lower surface; racemes and flower buds tan-brown, ap-
pressed-puberulent; calyx 3–5-lobed at anthesis ................................ 11
11(10). Stipe ≥ 10 mm long, 2 or more times as long as the ovary at anthesis .. 12
 Aldina 247

11. Stipe 2–2.5 mm long, never longer than the ovary at anthesis .............. 13
12(11). Leaflets 10–12 × 6–7.5 cm, obtuse-mucronulate to rounded, pale lilac on
lower surface with a pruinulose, waxy bloom; calyx tube cupular, 7–8 ×
9 mm wide at anthesis; petals 15 mm long and strongly cucullate
.......................................................................................................... A. sp. E
12. Leaflets 16–17 × 7–8 cm, obtuse-acuminate, monocolorous and nearly
shiny, waxy bloom not prominent on lower surface; calyx tube funnel-
shaped, 10 × 8–9 mm at anthesis; petals 20–25 mm long and somewhat
flattened ...................................................................................... A. latifolia
13(11). Petals 3 or 4 .............................................................................................. 14
13. Petals 5 or 6 .............................................................................................. 15
14(13). Leaflet venation stongly elevated, conspicuous on both surfaces; petals
2–2.5 × 1–1.8 cm; leaflets oblong to oblong-ovate, 7.5–18 × 4.5–9 cm,
scantily appressed-puberulent but shiny and the same color on the
lower surface ............................................................................ A. reticulata
14. Leaflet venation inconspicuous, usually obscure on upper surface; petals
1–2 cm long; leaflets elliptic, obtuse to rounded at the apex, 8–9 × 5–6 cm,
dark wine-colored on the lower surface .......................................... A. sp. F
15(13). Leaflets acute to acuminate apically, upper and lower surfaces markedly
discolorous ............................................................................................ 16
15. Leaflets obtuse to rounded-obtuse, upper and lower surfaces not mark-
edly discolorous .................................................................................... 17
16(15). Leaflets tessellate and appressed-puberulent on lower surface; petals
5, 2–2.3 cm long .......................................................................... A. discolor
16. Leaflets not tessellate, golden-pubescent on lower surface; petals 6, ca.
1.8 cm long ...................................................................................... A. aurea
17(15). Leaf axis and petiole appressed-puberulent, petiolules 5–7 mm long; pet-
als 6 ............................................................................................. A. elliptica
17. Leaf axis and petiole glabrous when mature, petiolules 10–25 mm long;
petals 5 ................................................................................................. 18
18(17). Leaflets ovate to orbicular, 7–15 × 6–14 cm, lateral veins conspicuous,
7–13 per side; racemes paired, 7–19-flowered ....................... A. paulberryi
18. Leaflets elliptical, 8.5–10 × 3.5–4.5 cm, lateral veins inconspicuous, 22–25
per side; racemes paniculate, many-flowered, elongate ....... A. petiolulata

Aldina aurea R.S. Cowan, Mem. New York
Bot. Gard. 10(1): 145. 1958.
Tree 3–4 m tall; leaflets discolorous,
pallid on lower surface, apically acute-
acuminate; ovary densely pubescent; pet-
als 6, ca. 1.8 cm long. Montane forests,
500–1300 m; Amazonas (Cerro Yutajé).
Endemic.
Aldina berryi R.S. Cowan & Steyerm., Ann.
Missouri Bot. Gard. 71: 312. 1984.
Tree ca. 20 m tall; leaflets completely gla-
brous, shiny on both surfaces, ± coriaceous;
petals 5–7, white; ovary glabrous. Montane
and gallery forests, 800–1000 m; Bolívar
(near Cerro Guaiquinima, Río Chiguao). En-
demic.
Aldina discolor Spruce ex Benth. in Mart.,
Fl. Bras. 15(2): 12. 1870. —Cojón de
verraco.
Tree 18–25 m tall; leaflets completely gla-
brous, discolorous, tessellate, appressed-pu-
berulent on lower surface; inflorescence in
multiple, axillary racemes; petals 5, 2–2.3
cm long, yellowish. Nonflooded evergreen
lowland forests, 100–200 m; Amazonas
(Yavita). Brazil (Amazonas).
Aldina elliptica R.S. Cowan, Mem. New
York Bot. Gard. 10(1): 146. 1958.
Tree 10–25 m tall; leaf stalk including
petiole appressed-puberulent into maturity,
petiolules 5–7 mm; leaflets appressed-pubes-
248 FABACEAE
cent on lower surface, obtuse to rounded-ob-
tuse, not discolorous; petals 6, ca. 1.5 cm
long; ovary uniformly pubescent, not ex-
serted at anthesis. Tepui slope forests, ca.
1800 m; Amazonas (Cerro Yutajé). Endemic.
Aldina heterophylla Spruce ex Benth. in
Mart., Fl. Bras. 15(2): 13. 1870.
Small tree (in the flora area) ca. 2 m tall;
leaves 5-foliolate; leaflets glabrous and
shiny, ovate-elliptic, obtuse to fairly mucr-
onate, 8–10 × 4–5 cm; ovary golden-seri-
ceous; fruit 1–4 cm long, irregularly shaped.
White-sand shrub savannas, ca. 500 m;
Amazonas (12 km west-southwest of Cerro
Autana). Brazil.
Aldina kunhardtiana R.S. Cowan, Mem.
New York Bot. Gard. 8: 106. 1958.
Small (10 m tall) to giant (35 m tall) tree;
leaflets “lauraceous” with pronounced obtuse
tip, the lower surface with distinctive rich,
brown-villous pubescence; inflorescence an
elongate, open, terminal panicle of racemes,
dark, golden-brown-velutinous; fruits glo-
bose, to 8 cm diameter. Evergreen lowland
forests, forest-savanna ecotones, 100–200 m;
Amazonas (Río Casiquiare, from Río Sipapo
south to San Fernando de Atabapo, upper
Río Temi, San Carlos de Río Negro to Solano,
Yavita to Maroa road). Brazil. ŠFig. 222.
Aldina latifolia Spruce ex Benth. in Mart.,
Fl. Bras. 15(2): 12. 1870.
Tree 5–17 m tall. Southern Venezuela, ad-
jacent Brazil; 3 varieties, all in the flora
area.
Key to the Varieties of A. latifolia
1. Leaflets appressed-puberulent on the
lower surface ............... var. pubescens
1. Leaflets glabrous on both surfaces ........ 2
2. Leaves 1- or 3-foliolate ..............................
........................... var. auyantepuiensis
2. Leaves mostly 3- or 5-foliolate ..................
........................................ var. latifolia
A. latifolia var. auyantepuiensis Pittier ex
H.S. Irwin, Acta Bot. Venez. 2: fig. 13.
1967. —Automoyek (Arekuna).
Tree 5–8 m tall. Dwarf montane forests,
700–1100 m; Bolívar (Guayaraca, east-north-
east of Icabarú, Kamarata, Río Carapo along
Cerro Guaiquinima). Endemic. ŠFig. 221.
This is perhaps the most weakly defined
variety, for the leaves at the apex of the
branchlets often have a reduced number of
leaflets.
A. latifolia var. latifolia. —Cojón de
verraco.
Tree 5–15 m tall. Lowland, usually ripar-
ian forests, white-sand savannas, 100–200
m; Amazonas (Maroa, Pimichín, Río Autana,
Río Guayapo). Endemic.
A. latifolia var. pubescens R.S. Cowan,
Mem. New York Bot. Gard. 8: 107. 1953.
Bushy tree ca. 8 m tall. White-sand sa-
vannas at edges of riparian forests, 100–200
m; Amazonas (Caño Cupueni near San
Fernando de Atabapo, near Yavita). Brazil
(Amazonas).
Aldina macrophylla Spruce ex Benth. in
Mart., Fl. Bras. 15(2): 13. 1870.
Tree to 17 m tall; leaflets appressed-pu-
bescent on lower surface; ovary glabrous or
nearly so. Venezuela; 2 varieties, both in the
flora area. Endemic.
Key to the Varieties of A. macrophylla
1. Stipe 6–10 mm with ovary noticeably ex-
serted in mature flower; calyx tube cu-
pulate, 6–8 × (6–)7–9 mm; petals 18–25
mm long; bracteoles inserted 1.5–2 mm
below base of calyx tube; stipe com-
pletely and densely appressed-puberu-
lent .......................... var. macrophylla
1. Stipe ≤ 2 mm with ovary not exserted in
mature flower; calyx tube 3–4 × 5–6 mm;
petals 8–10 × 4–5 mm; bracteoles in-
serted at base of calyx tube; stipe pubes-
cence restricted to a central circular
band ....................... var. yapacanensis
A. macrophylla var. macrophylla
Tree to 17 m tall. Riparian forests, white-
sand savannas, 100–200 m; Amazonas (Caño
Chimoni in Río Siapa basin, Caño San
Miguel, Río Yatúa). Endemic. ŠFig. 224.
A. macrophylla var. yapacanensis (R.S.
Cowan) Stergios, comb. & stat. nov.
—Aldina yapacanensis R.S. Cowan,
Mem. New York Bot. Gard. 8: 108. 1953.
Tree 8–10 m tall; flowers brownish to
white; ovary generally not exserted in ma-

ture flowers; calyx funnel-shaped, 3–4 × 5–6
mm; bracteoles inserted at base of calyx
tube; pubescence on stipe restricted to a
band in the middle. Black-water riparian
and swamp forests, 100–200 m; Amazonas
(Caño Yapacana, Río Emoni in lower Río
Siapa basin). Endemic.
This variety was reduced to synonomy by
Cowan (Mem. New York Bot. Gard. 10(1):
146–147. 1958), because of relatively high
variability in leaflet size and shape, and in
variation in petal and stipe size as flowers
opened and matured. However, with further
scrutiny and an additional, copiously flow-
ered collection, the ovary was found to con-
sistently not be exserted. Also, as shown by
the type collection, the bracteoles are in-
serted 1.5–2 mm below the base of the calyx
tube in var. macrophylla, but in var.
yapacanensis, they are at the base. In var.
yapacanensis, the pubescence on the stipe is
restricted to a band in the middle. Although
these characters may seem weak, they fur-
nish more reliable criteria than those distin-
guishing some varieties of Aldina latifolia.
Aldina paulberryi Aymard, Novon 8: 330.
1998.
Tree ca. 15 m tall; leaves 1- or 3-foliolate; leaf-
lets ovate-orbicular, the apex rounded-obtuse, 7–
15 × 6–14 cm, not discolorous, lateral veins 7–13
per side; racemes 2–many, 7–19-flowered. Ever-
green lowland forests, 100–200 m; Amazonas
(near Macuruco on upper Río Orinoco). Endemic.
ŠFig. 225.
This species can be distinguished from its
nearest apparent relative, Aldina petio-
lulata, by orbicular, not elliptical leaflets,
more prominent but less numerous lateral
veins, and paniculate, fewer flowered,
shorter racemes.
Aldina petiolulata R.S. Cowan, Mem. New
York Bot. Gard. 10(4): 70, fig. 59. 1961.
Tree to 40 m tall; leaflets appressed-pu-
berulent on lower surface, elliptical, with
22–25 lateral veins; inflorescence a many-
flowered panicle of elongate racemes. Mon-
tane forests, 700–800 m; Amazonas (Sierra
de la Neblina). Endemic.
Aldina reticulata R.S. Cowan, Mem. New
York Bot. Gard. 8: 107. 1953.
Small tree; lower surface of leaflets and
ovary appressed-puberulent; venation
Aldina 249
strongly elevated and conspicuous on both
surfaces; stipe ≤ ovary length; petals 4.
Lower montane forests, lowland swampy for-
ests, 100–800 m; Amazonas (headwaters of
Caño Grande in Cerro Sipapo, Maroa area,
Río Temi). Endemic. ŠFig. 223.
Aldina sp. A
Shrubby tree 2–6(–8) m tall; leaflets gla-
brous, cartilaginous-coriaceous, cordate at
the base, 7–8 × 4–5 cm; flowers showy, petals
3–4 × 2 cm; ovaries 2 or 3 on a single stipe;
fruits thickly fusiform, 2–4-seeded. Open,
low-rocky, shrubby slopes, 400–1000 m;
Bolívar (Cerro Kampe, middle Río Paragua
basin, Serranía Senkopirén). Endemic.
This spectacular species is probably the
most distinct one in the genus, easily recog-
nized by very large flowers with a long-ex-
serted, thick stipe bearing 2 or 3 ovaries.
Aldina sp. B
Tree 8–10 m tall; leaflets 10–11 × 4–4.5
cm; calyx and unopened buds usually gla-
brous, but sometimes sparsely white-ap-
pressed-puberulent near the base; pedicels
11–12 mm long; bracteoles borne 3–3.5 mm
below the base of the calyx tube. Humid,
lower montane forests, ca. 500 m; Bolívar
(Quebrada El Trueno 90 km south of La
Paragua and at the base of Cerro Guai-
quinima). Endemic.
This species is readily distinguished by the
glabrous, broadly lanceolate-acuminate leaf-
lets that have an off-white, pallid-waxy coat-
ing on the lower surface. It is probably closest
to Aldina latifolia, which also has simple, axil-
lary racemes and densely appressed-tomen-
tose stipe and ovary. Mature flowers of both
species have a noticeably exserted pistil.
Aldina sp. C
Shrubby tree 2–5 m tall, unique by the com-
bination of glabrous leaflets and branched,
multiple racemes; leaflets ovate-elliptic, mu-
cronate, coriaceous; inflorescence appressed-
ferruginous-tomentose; ovary densely puberu-
lent-tomentose. White-sand shrub savannas,
forest-savanna ecotones, 100–1300 m; Ama-
zonas (Cerro Autana, Cerro Parú).
Aldina sp. C is closely related to A. lati-
folia, A. berryi, and A. heterophylla, all of
which have shiny, glabrous leaflets without
the waxy coating on the lower surface. It can
be distinguished from A. latifolia by the non-
250 FABACEAE
tessellate leaflets and multiple-branched ra-
cemes; from A. berryi by the entirely shiny
leaflets and pubescent ovary; and from A.
heterophylla by the 5–7 leaflets and lustrous,
ferruginous pubescence.
Aldina sp. D
Tree ca. 10 m tall; leaflets glabrous with a
glaucous, somewhat ashy bloom on the lower
surface, ovate-elliptic, obtuse-rounded at the
apex, 9–11 × 6–6.5 cm; ovary glabrous. Semi-
black-water, riparian swamp forests, 100–
200 m; Amazonas (Río Emoni in the lower
Río Siapa basin). Endemic.
This species most resembles Aldina berryi
in its 4 petals and glabrous ovary, but differs
in the more notably pallid-discolorous, ob-
tuse to rounded leaflets, in the longer, ex-
serted pistil, and in the longer pedicellate
but smaller flowers.
Aldina sp. E
Tree 4–6 m tall, occasionally 10–30 m tall.
Riparian or swamp forests, 100–200 m;
Amazonas (Caño Momoni in the Río Casi-
quiare basin, Caño Yagua, Macuruco, Río Ne-
gro). Endemic.
This species is similar to Aldina macro-
phylla var. pubescens in the 3–5-lobed calyx
and the long stipe, but differs in having
smaller, obtuse-mucronate to rounded, dis-
colorous leaflets.
Fig. 221. Aldina latifolia var. auyantepuiensis
Aldina sp. F
Tree 3–6 m tall. Swamp forests, flooded
riparian alluvial plains, 50–200 m; Ama-
zonas (Laguna Pasiba, Río Negro). Endemic.
Aldina sp. F and and A. reticulata are the
only two species that have 3 or 4 petals in-
stead of the usual 5 or 6. Aldina sp. F differs
from A. reticulata in the obscure venation on
the upper leaflet surface, more obtuse, ovate-
elliptic leaflets, smaller flowers, and a dark,
wine-colored lower leaflet surface.
Aldina sp. G
Medium-sized tree; leaves 5- or 7-
foliolate; leaflets completely glabrous on
both surfaces and shiny without any waxy
coating on lower surfaces and concolorous,
chartaceous; petiolules 4–6 mm long; inflo-
rescence and flower buds tan-brown, ap-
pressed-puberulent. Savannas, 800–1000 m;
Amazonas (Cerro Sipapo). Colombia
(Vaupés), Brazil (Amazonas: Manaus) .
This species is most closely related to
Aldina sp. C, both with 5- or 7-foliolate
leaves, entirely shiny, discolorous leaflets,
and pubescent ovary. It can be distinguished
from Aldina sp. C by its chartaceous rather
than coriaceous leaflets, which are obtuse
and acuminate rather than elliptic and mu-
cronate, the inflorescence and flower buds
are brown-tan appressed-puberulent rather
than loosely ferruginous-velutinous.
 Aldina 251

Fig. 222. Aldina kunhardtiana

252 FABACEAE

Fig. 223. Aldina reticulata

Fig. 224. Aldina macrophylla var. macrophylla

 Aldina 253

Fig. 225. Aldina paulberryi

254 F ABACEAE

5. ALEXA Moq. in A. DC., Prodr. 13(2): 168. 1849.

Alexandra R.H. Schomb., London J. Bot. 4: 12. 1845, non Alexandra Bunge
1843.
by Charles H. Stirton and Gerardo A. Aymard C.

Trees 10–36 m tall, rarely cauliflorous. Stipules minute, rapidly caducous.

Leaves 5–17-foliolate, odd-pinnate, alternate. Leaflets membranaceous or coria-
ceous. Inflorescences terminal, racemes short to very elongate, becoming quite
woody with age. Flowers white, green, cream, yellow, orange, or crimson, strongly
zygomorphic, pedicellate, bracteate (the bracts sometimes swollen and glandular);
bracteoles minute and caducous; pearl glands absent. Calyx turbinate-campanulate
to campanulate, incurved or erect, densely pubescent, teeth 3–5, minute. Petals free,
erect or ascending; standard outermost, elliptic to obovate, base truncate, apex
emarginate, densely pubescent or glabrous; wing and keel petals scarcely differenti-
ated, mostly falcate. Stamens 10–12, equal, free, staminodes sometimes present;
anthers dorsifixed, versatile, longitudinally dehiscent, elongate. Ovary stipitate, 4–
9-ovulate, glabrous or pubescent; style filiform; stigma inconspicuous, terminal. Le-
gume dehiscent, woody, apex acute, calyx woody and persistent in fruit, densely red-
dish brown, glabrescent, 1–7-seeded. Seeds swollen, ovate or elliptic, glabrous.
Venezuela, Guyana, Suriname, French Guiana, Brazil; 10 species, 6 in Venezu-
ela, all in the flora area.

Key to the Species of Alexa

1. Leaves membranaceous, with well-developed drip tips; flowers tubular,

but petals reflexing very quickly; flower bracts swollen and glandular;
corolla cream to greenish white ............................................................ 2
1. Leaflets subcoriaceous, with drip tips absent or scarcely developed; flow-
ers typically papilionoid without petals reflexing; flower bracts not
swollen and glandular; corolla pale yellow to crimson and orange ..... 3
2(1). Inflorescence dense; stipe of ovary glabrous; calyx as wide as long; style
finely pubescent, to 2 cm long, uncinate at the apex ................. A. cowanii
2. Inflorescence lax; stipe of ovary pubescent; calyx 2–3 times longer than
broad; style glabrous, 6–8 cm long, curved at the middle ......................
........................................................................................ A. canaracunensis
3(1). Leaflets densely yellow- to brown-puberulous and opaque on the upper
surface .................................................................................................... 4
3. Leaflets glabrous or glabrescent and shiny on the upper surface ............ 5
4(3) Leaflets 7 or 9, densely pubescent on the lower surface only; pedicels
2–3.5 cm long ..................................................................... A. bauhiniiflora
4. Leaflets 11 or 13(15, 17), pubescent on both surfaces; pedicels 0.4–1.5 cm
long .............................................................................................. A. confusa
5(3). Inflorescences in terminal racemes 10–21 cm long; corolla densely pubes-
cent, pale yellow; petals membranous ................................ A. herminiana
5. Inflorescences borne on trunk, to 7 cm long; corolla glabrous, crimson to
orange; petals subcoriaceous ............................................... A. imperatricis
 Alexa 255

Fig. 226. Alexa cowanii

256 F ABACEAE

Fig. 227. Alexa herminiana

 Alexa 257

Fig. 228. Alexa imperatricis

258 F ABACEAE

Alexa bauhiniiflora Ducke, Bull. Mus.
Hist. Nat. (Paris), sér. 2: 732. 1932.
Tree 5–10 m tall. Along rivers, 100–500
m; Amazonas (Río Mawarinuma, Río Pa-
duari, headwaters of Río Siapa, Sierra de la
Neblina). Brazil (Amazonas, Pará).
guaza, Río Tabaro, Río Tonoro, Santa María
de Erebato, Serranía Pia-Zoi), Amazonas
(Ocamo, upper Río Cunucunuma, Río Pada-
mo). Guyana, Brazil (Amazonas, Roraima).
Some branches become hollowed out and
are occupied by stinging ants.

Alexa canaracunensis Pittier, Bol. Soc.
Venez. Ci. Nat. 7: 308. 1942. —Cai-
careño, Kadaka (Arekuna), Kayapa,
Tinajito, Tunadi (Yekwana).
Small tree 3–12 m tall, rarely taller;
leaves firmly membranaceous; corolla green-
ish cream. Evergreen lowland to montane
forests, 100–1500 m; Bolívar (Amaruay-
tepui, Cerro Paují, El Dorado, Las Claritas,
Macizo del Chimantá [Chimantá-tepui], Río
Canaracuni, Río Caura, Río Cuyuní, Río
Icabarú, Río Padauri, upper Río Paragua,
Río Tírica), Amazonas (Sierra Parima). Bra-
zil (Roraima).
Young stems subtending the inflores-
cences become hollowed out and occupied by
ants, which feed on nectar provided by swol-
len pear-shaped flower bracts.
Alexa confusa Pittier, Bol. Soc. Venez. Ci.
Nat. 8: 262. 1943. —Caicareño monta-
ñero, Dequira-arepillo (Piaroa), Guamo
peludo, Kure-yek (Arekuna), Leche de
cochino, Pitopitosiji (Yanomami), Tune-
ene (Yekwana).
Alexa superba R.S. Cowan, Mem. New
York Bot. Gard. 8: 117. 1953.
Alexa imperatricis auct. non R.H. Schomb.
1845, nec Baill. 1870.
Tree 10–30 m tall; leaves subcoriaceous,
the upper surface dark green, the lower sur-
face yellowish green and densely pubescent;
corolla cream-colored on inner faces, buff
brown outside. Evergreen lowland to lower
montane forests, occasionally along savanna
borders, 50–1300 m; Bolívar (Icabarú, Río
Caura, Río Karún, Río Pacairao, Río Par-
Alexa cowanii Yakovlev, Bot. Zurn. (Mos-
cow & Leningrad) 62(3): 436. 1977.
—Tinajito.
Tree to 20 m tall; leaves membranaceous,
glabrous; corolla white to pale greenish white.
Evergreen lowland to lower montane forests,
100–500 m; Bolívar (near El Dorado, along the
Guyana border, Kilómetro 88, Río Uiri-yuk
near La Escalera). Guyana (upper reaches of
the Kamarang River watershed). ŠFig. 226.
The branches often become hollowed out
and are occupied by ants.
Alexa herminiana N. Ramírez, Ernstia 37:
41. 1986.
Tree 10–20 m tall; leaves coriaceous, the
upper surface rich green, the lower surface
paler; corolla pale yellow. Evergreen lowland
to lower montane forests, 50–500 m; Ama-
zonas (Caño Piedra on Cerro Sipapo massif,
Raudal Rabipelado near Samariapo, Río
Cataniapo). Endemic. ŠFig. 227.
Alexa imperatricis (R.H. Schomb.) Baill.,
Hist. Pl. 2: 362. 1870. —Alexandra
imperatricis R.H. Schomb., London J.
Bot. 4: 12. 1845, non Baker 1888, nec
Pittier 1942. —Coreco, Leche de cochino.
Tree 30–40 m tall, cauliflorous; leaves co-
riaceous, slightly glossy, the upper surface
darker green; corolla orange, one petal and
stamens crimson to orange. Evergreen low-
land forests, 50–300 m; Delta Amacuro (east-
northeast of El Palmar, Río Toro, Serranía de
Imataca), Bolívar (El Dorado, Kilómetro 88,
Río Cuyuní, Río Toro). Adjacent Guyana.
ŠFig. 228.

6. ALYSICARPUS Desv., J. Bot. Agric. 1: 120. 1813.

by Nidia L. Cuello A.
Herbs, suberect to prostrate; stems ascending to procumbent, terete, striate.
Leaves alternate, 1(–3)-foliolate; leaflets linear-lanceolate to ovate, reticulate-
venose; stipules scarious, acuminate, free or connate; bistipellate. Inflorescence ter-
minal or rarely axillary, racemose; bracts scarious, deciduous; pedicels paired, short.
Flowers small. Calyx deeply 5-lobed, the lobes unequal, the upper 2 connate nearly
to the apex, glumaceous; corolla reddish purple, rarely orange; standard obovate to
 Andira 259

orbicular, clawed; wing petals obliquely oblong; keel petals slightly incurved, obtuse.
Stamens 10, diadelphous, the upper connate only basally; anthers uniform. Ovary
sessile or shortly stipitate; ovules numerous; style filiform, incurved at the apex;
stigma broadly capitate, terminal. Fruit a loment, terete to compressed, sub-
moniliform, indehiscent. Seeds suborbicular or globose, smooth, lustrous, estro-
phiolate.
Old World tropics; 25 species, 1 introduced into tropical America as forage, 1 in
Venezuela.

Alysicarpus vaginalis (L.) DC., Prodr. 2:

353. 1825. —Hedysarum vaginale L., Sp.
Pl. 746. 1753.
Annual herb, suberect to prostrate. In
Trachypogon savannas on low hills and dis-
turbed areas, 200–500 m; Delta Amacuro
(Tucupita), north-central and eastern Bolí-
var (Altiplanicie de Nuria, 16 km north of El
Manteco, El Pao, Tumeremo). Aragua, Ba-
rinas, Guárico, Miranda, Portuguesa; U.S.A.
(Florida), Mexico, Honduras, Panama, West
Indies, Colombia, Guyana, Suriname, French
Guiana, Ecuador, India, Asia, Oceania, Ma-
dagascar. ŠFig. 229.

Fig. 229. Alysicarpus vaginalis

7. ANDIRA Juss., Gen. Pl. 363. 1789, nom. cons.

by Gerardo A. Aymard C., Nidia L. Cuello A., and R. Toby Pennington
Trees or shrubs. Leaves alternate, odd-pinnate, rarely trifoliolate or unifoli-
olate; leaflets opposite or rarely alternate; stipels setaceous, persistent or absent.
Inflorescences terminal and axillary panicles, often crowded and subsessile. Flowers
rose, violet, or white; bracts and bracteoles small, caducous. Calyx truncate or
shortly 5-dentate; petals long-clawed; standard suborbicular, exappendiculate; wing
petals straight, oblong, free; keel petals similar, overlapping at the back. Stamens
10, diadelphous; vexillar stamen free; anthers ovate, small, versatile; ovary stipi-
tate, ovules (1)2–8; style short, incurved; stigma small, terminal. Fruit drupaceous,
ovoid, obovoid, or globose, indehiscent, shortly stipitate with fleshy mesocarp and
hard, woody endocarp. Seeds 1(–3), large, filling the entire cavity, exalbuminate.
Mexico, Central America, West Indies, Colombia, Venezuela, Guyana, French
Guiana, Suriname, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Africa (1
species); ca. 30 species, 5 in Venezuela, all in the flora area.
260 F ABACEAE

Key to the Species of Andira

1. Leaves all trifoliolate ................................................................. A. trifoliolata

1. Leaves with 5–15 leaflets (occasionally some leaves trifoliolate) ............ 2
2(1). Leaves with 5–7 leaflets (occasionally trifoliolate) ................................... 3
2. Leaves with 9–15 leaflets (never trifoliolate) ............................................ 4
3(2). Leaflets minutely pubescent on lower surface, the apex rounded to emar-
ginate ................................................................................... A. tervequinata
3. Leaflets glabrous on lower surface, the apex cuspidate ..................... A. sp. A
4(2). Branches and branchlets glabrous; leaflets glabrous, the apex acuminate;
stipules 5–10 mm long ................................................................ A. inermis
4. Branches and branchlets brown-tomentose; leaflets minutely pubescent
on lower surface, the apex obtuse or retuse; stipules ca. 3 mm long
............................................................................................ A. surinamensis

Andira inermis (W. Wright) DC., Prod. 2:
475. 1825. —Geoffroea inermis W.
Wright, London Med. J. 8: 256. 1787.
—Pilón.
Mexico, Central America, Colombia, Ven-
ezuela, Guyana, Suriname, French Guiana,
Ecuador, Peru, Brazil, Bolivia, Argentina; 2
subspecies, 1 in Venezuela.
A. inermis subsp. inermis
Tree to 35 m tall; stipules linear; petals
pink or purplish, the standard with a central
white spot. Deciduous to evergreen lowland
forests, 50–100 m; Delta Amacuro (Río
Cuyubini), Bolívar (southwest of Caicara,
Maniapure). Barinas, Distrito Federal, Fal-
cón, Miranda, Portuguesa, Yaracuy, Zulia;
Mexico, Central America, Antilles, Colombia,
Guyana, Suriname, French Guiana, Ecua-
dor, Peru, Brazil, Bolivia, Argentina, culti-
vated elsewhere in the tropics. ŠFig. 230.

Fig. 230. Andira inermis subsp. inermis

 Andira 261

Fig. 231. Andira surinamensis

Fig. 232. Andira trifoliolata

262 F ABACEAE

Andira surinamensis (Bondt) Splitg. ex with sandy savannas, 300–500 m; Bolívar

Amshoff, Meded. Bot. Mus. Herb. Rijks
Univ. Utrecht 52: 60. 1939. —Geoffroea
surinamensis Bondt, Cort. Geoffr. Suri-
nam. 13, figs. 1–8. 1788. —Arisoru
(Warao), Canelito negro, Palo blanco,
Pilón, Pilón rebalsero, Sobó (Yekwana),
Winka, Wonka (Panare).
Geoffroea retusa Poir. in Lam., Encycl. 8:
182. 1808. —Andira retusa (Poir.) DC.,
Prod. 2: 475. 1825.
Tree to 40 m tall; petals pink to violet, the
standard with a central white spot. Decidu-
ous to evergreen and riparian forests, some-
times on granitic outcrops, 50–400 m; Delta
Amacuro (Caño Arature, east-northeast of El
Palmar), northern Bolívar, Amazonas (Caño
Yutajé, Capibara, La Esmeralda, Puerto Aya-
cucho, Río Casiquiare, Río Cataniapo, Sama-
riapo). Anzoátegui, Apure, Barinas, Táchira;
Colombia, Trinidad, Guyana, Suriname,
French Guiana, Ecuador, Peru, Brazil. ŠFig.
231.
The fruits are used by the Warao Amer-
indians to treat skin diseases.
Andira tervequinata R.T. Penn., Aymard
& Cuello, Novon 7: 72. 1997.
Tree to 20 m tall. Lower montane forests
(Río Aparamán at base of Amaruay-tepui,
banks of Río Trueno at base of Cerro
Guaiquinima). Endemic.
Andira trifoliolata Ducke, Arq. Inst. Biol.
Veg. 4(1): 22. 1938. —Congrio.
Tree to 20(–30) m tall; petals white or dull
flesh-colored. Evergreen lowland forests,
edges of white-sand savannas, riparian for-
ests, 100–200 m; Amazonas (Caño San
Miguel, lower Río Baría, Río Casiquiare, San
Carlos de Río Negro, Santa Bárbara, Solano).
Brazil (Amazonas). ŠFig. 232.
A collection currently included here, Gen-
try and Tillett 10889 (FHO, MO), may repre-
sent a new species. It is a shrub, while all
other records of Andira trifoliolata are trees.
Moreover, the leaflets have sparse appressed
trichomes on their lower surfaces, whereas
all other specimens of A. trifoliolata are gla-
brous.
Andira sp. A. —Congrio.
Small tree ca. 5 m tall; petals white.
Shrubby white-sand savannas, 100–200 m;
Amazonas (Caño San Miguel, Caño Ucata
southeast of Síquita, lower Río Ventuari).
Brazil (Amazonas, Rondônia).

8. BARBIERIA DC., Prodr. 2: 234. 1825.

by Paul R. Fantz
Woody (not in flora area) or herbaceous vine, scandent or trailing; branches
striate-terete, densely hirsute, trichomes reddish brown becoming whitish with age;
pubescence of minutely hooked trichomes common (view at 20–30×). Leaves alter-
nate, odd-pinnate, petiolate, stipulate, stipellate; leaflets (9–)13–21(–25), opposite,
oblong to elliptic-oblong, entire, broadly acute to obtuse, mucronate, appressed-pi-
lose on lower surface; stipules persistent, striate, lanceolate, subulate-acuminate;
stipels persistent, 1-veined, linear-subulate; petiolules quadrate. Inflorescence axil-
lary or terminal, pseudoracemose, few-flowered, long-pedunculate; pedicels paired
at nodes; bracts striate, lance-subulate; bracteoles paired at calyx base, lanceolate,
subulate-acuminate. Flowers resupinate, showy, papilionaceous, red. Calyx narrow-
tubular, persistent in fruit, 5-lobed, the lobes deltoid to rapidly subulate above; stan-
dard complicate, long-clawed, spurless, oblong-oblanceolate; wing petals oblong,
long-clawed, much shorter than the long-clawed, elliptic-oblong keel petals which
nearly equals length of standard. Stamens diadelphous, often persistent in fruit.
Pistil enclosed within staminal sheath; ovary sessile; style linear, densely barbate.
Legume sessile, linear, straight, without costa, enclosed at base by persistent calyx,
valves puberulent-hirsute, strongly transversly impressed between the seeds, spi-
rally twisting upon dehiscence 0.5–1.5 turns. Seeds dark brown to black, smooth,
transverse-oblong.
 Bowdichia 263

Fig. 233. Barbieria pinnata

Widespread in the Neotropics, but apparently rare locally since few collections
exist in herbaria as compared to its range; 1 species.

Barbieria pinnata (Pers.) Baill., Hist. Pl. 2:
263. 1870. —Galactia pinnata Pers.,
Syn. Pl. 2: 302. 1807. —Clitoria pinnata
(Pers.) R.H. Sm. & G.P. Lewis, Kew Bull.
46: 320. 1991.
Clitoria polyphylla Poir. in Lam., Encycl.
suppl. 2: 300. 1811. —Barbieria poly-
phylla (Poir.) DC., Mém. Légum. 242, t.
39. 1825.
Moist soils in open areas of forests and forest
edges, 500–1000 m; Bolívar (Río Icabarú, Santa
Elena de Uairén). Barinas, Sucre; Southern
Mexico, Central America, West Indies, Colom-
bia, Peru, Brazil, Bolivia. ŠFig. 233.

9. BOWDICHIA H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 376. 1823.

by Charles H. Stirton
Large shrubs or small trees to 10 m tall. Leaves 9–21-foliolate, odd-pinnate, al-
ternate, deciduous; stipules 3–7 mm long; leaflets chartaceous or coriaceous, alter-
nate to opposite, stipellate. Inflorescences axillary or terminal racemes. Flowers
blue, bisexual, zygomorphic, bracts and bracteoles small, triangular, fringed with
pearl glands. Calyx turbinate with subequal lobes, sinus of vexillar lobes broad,
teeth fused for half their length; petals 5, unequal, free, glabrous, clawed; standard
blue with yellow or rose nectar patch, very broadly ovate, emarginate; wing petals
2–3 times larger than the keel petals, asymmetrically obovate; keel petals narrowly
cultrate (knife-blade-like). Stamens 10, free, filaments subequal with thickened
bases; anthers subequal, often staminodal, oblong, dorsifixed. Ovary stipitate, 5–10-
ovulate, densely or sparsely pubescent; style curved, shorter than ovary; stigma
large, terminal, capitate. Fruits stipitate, compressed, indehiscent, linear-oblong to
264 F ABACEAE

Fig. 234. Bowdichia virgilioides

elliptic, membranous, upper suture shortly winged. Seeds oblong, compressed,

asymmetric, with subterminal hilum.
Colombia, Venezuela, Suriname, Guyana, Brazil, Bolivia; 2 species, 1 in Ven-
ezuela.

Bowdichia virgilioides H.B.K., Nov. Gen.
Sp. (quarto ed.) 6: 376. 1823. —Alcor-
noque, Alcornoque sabanero, Cornoco.
Tree 6–10 m tall; corolla bluish violet.
Trachypogon-Curatella-Byrsonima savanna-
dry forest ecotone, granitic outcrops, 50–500
m; Bolívar (Caicara, Ciudad Bolívar, Puerto
Ordaz, Represa Guri, Río Cuchivero), Ama-
zonas (Cacuri, Puerto Ayacucho, lower Río
Ocamo, Río Ventuari). Anzoátegui, Apure,
Aragua, Barinas, Cojedes, Guárico, Mérida,
Monagas, Portuguesa, Sucre, Trujillo, Zulia;
Colombia, Guyana, Brazil, Bolivia. ŠFig.
234.
This species is quite variable throughout
its range.

10. CAJANUS DC., Cat. Pl. Horti Monsp. 85. 1813, nom. cons. —Cajan Adans.,
Fam. Pl. 2: 326. 1763.
by Nidia L. Cuello A.
Shrubs or subshrubs. Leaves alternate, pinnately or sometimes digitately 3-
foliolate; leaflets with vesicular glands below, membranous, puberulent, oblan-
ceolate to lanceolate, basally cuneate, apically acute; stipules triangular-lanceolate,
caducous; stipels present or absent. Inflorescence an axillary or terminal raceme,
pedunculate or almost sessile; flowers 6–12 per rachis; bracts small, caducous. Calyx
campanulate, 5-lobed, the upper lobe longer, lobes acute or acuminate; corolla yellow
 Cajanus 265

or lined with red or standard dorsally reddish, to 3 cm long; standard ovate, auricu-
late; wing petals obliquely-obovate; keel petals auriculate, rounded-oblique, obtuse.
Stamens 10, diadelphous, the vexillar stamen free; anthers uniform. Ovary sessile;
ovules 4–6; style slender, glabrous; stigma capitate. Fruit linear-oblong, 2-valved
with oblique constrictions between the seeds, yellowish green with brown mottling,
puberulent. Seeds reniform to suborbicular, the hilum oblong, strophiolate.
Paleotropics (2 species sometimes escaping from cultivation in Neotropics); 32
species, 1 in Venezuela.

Cajanus cajan (L.) Millsp., Publ. Field

Columbian Mus., Bot. Ser. 2: 53. 1900.
—Cytisus cajan L., Sp. Pl. 739. 1753.
—Quinchoncho.
Shrub 1–3 m tall; widely cultivated for
food and forage, sometimes escaped on dis-
turbed ground, 50–1000 m; Delta Amacuro
(Santa Catalina), Bolívar (Gran Sabana, Isla
Anacoco, Serranía de Imataca). Aragua, Co-
jedes, Distrito Federal, Guárico, Mérida,
Miranda, Monagas, Portuguesa, Táchira,
Yaracuy, Zulia; Mexico, Central America, Co-
lombia, Guyana, Suriname, French Guiana,
Ecuador, Peru, Brazil (Goiás, Mato Grosso,
Minas Gerais, São Paulo), Bolivia, wide-
spread in Africa and Asia. ŠFig. 235.

Fig. 235. Cajanus cajan

266 F ABACEAE

11. CALOPOGONIUM Desv., Ann. Sci. Nat. (Paris) 9: 423. 1826.

Stenolobium Benth., Ann. Wiener Mus. 2: 125. 1837.
by Richard H. Maxwell
Small twining to large climbing vines. Leaves alternate, pinnately 3-foliolate;
stipules acute or lanceolate; leaflets mostly ovate, entire. Inflorescence axillary,
erect and pseudoracemose or fasciculate; bracts and bracteoles mostly linear, ciliate,
mostly caducous. Flowers medium-sized to small, numerous. Calyx tube 5-lobed, the
upper lobes united, the apex entire or emarginate; corolla blue or violet; standard
reflexed, basally biauriculate; wing and keel petals adherent. Vexillar stamen free;
anthers mostly 8 or 9. Ovary sessile, many-ovulate; style filiform or subulate, usu-
ally incurved; stigma terminal, capitate, glabrous. Fruit a legume, linear to linear-
oblong, several- to many-seeded, compressed, septate, dehiscent; exocarp trans-
versely furrowed between seeds. Seeds reniform to cuboid, the hilum small, oval.
Neotropics; 6–8 species, 3 in Venezuela, 2 of these in the flora area.

Key to the Species of Calopogonium

1. Calyx and fruit with appressed or spreading trichomes to ca. 1 mm long;

calyx lobes acute, shorter than or about equal to the tube length; fruit
ca. 8 mm wide; terminal leaflets broadly ovate to rhomboidal, occasion-
ally lobed ................................................................................. C. coeruleum
1. Calyx and fruit with erect or spreading trichomes to ca. 2 mm long or longer;
calyx lobes linear, greatly exceeding the tube length; fruit ca. 4 mm
wide; terminal leaflets ovate to elliptic or lanceolate ......... C. mucunoides

Fig. 236. Calopogonium coeruleum Fig. 237. Calopogonium mucunoides

 Canavalia 267

Calopogonium coeruleum (Benth.) C. lagartilla, Peine del diablo.

Wright in Sauvalle, Anales Acad. Ci. Méd.
Habana 5: 337. 1869. —Stenolobium
coeruleum Benth., Ann. Wiener Mus.
Naturgesch. 2: 125. 1838. —Caraotillo.
Vine, may become long and high-climbing.
Roadsides, fields, disturbed areas, 50–400 m;
Delta Amacuro (Río Toro), northern Bolívar,
Amazonas (Río Ocamo). Anzoátegui, Apure,
Barinas, Carabobo, Guárico, Miranda, Mona-
gas, Portuguesa, Sucre, Zulia; Mexico, Central
America, West Indies, Guyana, Suriname,
French Guiana, throughout Brazil and Colom-
bia, extending south through Ecuador and Peru
into Bolivia and tropical Argentina. ŠFig. 236.
Calopogonium mucunoides Desv., Ann.
Sci. Nat. (Paris) 9: 423. 1826. —Frijol de
Vine; the long trichomes, especially on the
fruit, may be yellow or a darker rusty color.
Disturbed, semi-open, humid sites, near sea
level to 300 m; Delta Amacuro (Caño
Angosturito, Caño Güiniquina), Bolívar
(middle Río Orinoco), Amazonas (San Carlos
de Río Negro). Guárico, Monagas, Zulia, and
probably throughout rest of Venezuela; Cen-
tral America, West Indies, Colombia, Guy-
ana, Suriname, French Guiana, Ecuador,
Peru, Brazil (Amapá, the drier northeastern
states south to Bahía, Pará, Amazonas,
Rondônia). ŠFig. 237.
This has been widely used as a cover
crop and apparently escapes as a weed
throughout central Brazil and in the Old
World tropics.

12. CANAVALIA DC., Prodr. 2: 403. 1825, nom. cons.

Canavali Adans., Fam. Pl. 2: 325, 531. 1763.
Wenderothia Schltdl., Linnaea 12: 330. 1838.
by Gerardo A. Aymard C. and Richard H. Maxwell
Climbing, twining, or trailing vines. Leaves alternate, 3-foliolate; stipules
small, lanceolate or oblong, deciduous or caducous; leaflets mostly ovate to subor-
bicular, entire, unlobed. Inflorescence axillary, pseudoracemose, mostly pendent, tu-
berculate; each tubercle 2–6-flowered, bracteate. Flowers mostly resupinate, showy,
bracteolate. Calyx 5-lobed, appearing bilabiate, the upper lobes largest, entire or
emarginate, the lower and lateral lobes crowded, minute; corolla whitish, pink, rose,
or shades of violet or purple; standard reflexed; wing petals free; keel petals wider
than wings, upcurved, frequently beaked, united distally. Stamens 10, monadelp-
hous or pseudomonadelphous. Pistil mostly curved; ovary pubescent, sessile or
short-stipitate; stigma capitate. Fruit a legume, oblong, compressed, dehiscent, the
upper sutures with 2 parallel ribs, frequently each valve with 1 or 2 extra ribs,
several- to many-seeded. Seeds ellipsoid, mostly compressed, the hilum oblong or
linear.
Pantropics; ca. 60 species, 11 in Venezuela, 5 of these in the flora area.

Key to the Species of Canavalia

1. Common trailer on seacoast beaches or twining vines on backbeach veg-

etation; leaflets somewhat orbicular, the apices frequently emarginate;
seed hilum ca. 6 mm long ............................................................... C. rosea
1. Inland species; leaflets ovate, elliptic, lanceolate, or combinations, not or-
bicular, the apices rounded, acute, to acuminate; seed hilum 6–22 mm
long ......................................................................................................... 2
2(1). Fruits without an extra rib; pubescence of leaflets long-silky, very dense
on lower surface of blade ..................................................... C. sericophylla
2. Fruits with an extra rib; leaflets glabrous or pubescence sparse to moder-
ately dense on lower surface of blade .................................................... 3
268 F ABACEAE

3(2). Upper calyx lobe gradually concave behind the apex; flowers to ca. 4 cm
long; seeds ca. 3 mm thick .................................................... C. grandiflora
3. Upper calyx lobe depressed behind the apex; flowers 2–3 cm long; seeds
9–12 mm thick ....................................................................................... 4
4(3). Leaflets broadly ovate, chartaceous; standard ca. 2.5 cm long; seed hilum
8–12 mm long, shorter than the seed; fruit to ca. 2.8 cm wide ..............
............................................................................................... C. brasiliensis
4. Leaflets lanceolate-elliptic, coriaceous; standard to ca. 3 cm long; seed hi-
lum 15–18 mm long, nearly as long as the seed; fruit to ca. 3.5 cm wide
..................................................................................................... C. dictyota

Canavalia brasiliensis Mart. ex Benth.,

Fig. 238.
Canavalia Ann. Wiener Mus. Naturgesch. 2: 135.
brasiliensis 1838.
Canavalia fendleri Piper, Contr. U.S.
Natl. Herb. 20: 570. 1925, pro parte ma-
jor, fide Sauer, Brittonia 16: 144. 1964.
Trailing vine in the open, or twining over
shrubs. Mud flats, lakeside lowlands, into
upper savannas, disturbed areas, 50–200 m;
Bolívar (near Caicara, Ciudad Bolívar).
Widespread elsewhere in Venezuela; U.S.A.
(Florida), Mexico, Central America, West
Indies, widespread in South America al-
though apparently rare in Amazon basin.
ŠFig. 238.
Canavalia brasiliensis is planted occa-
sionally as a cover crop and is sometimes a
weedy invader, usually in lowlands.
 Centrolobium 269

Canavalia dictyota Piper, Contr. U.S. Natl.
Herb. 20: 574. 1925.
Vine trailing, twining and climbing on un-
dershrubs, or occasionally high-climbing.
Open ground, thickets, rocky outcrops, near
sea level to 100 m; Delta Amacuro (Caño
Angosturita, Caño Güiniquina, Tucupita),
Bolívar (lower Río Caura, Río Pargueni),
Amazonas (Puerto Ayacucho). Distrito Fed-
eral, Lara, Nueva Esparta, Sucre, Zulia;
Panama, West Indies, Colombia (Magda-
lena), Guyana, Suriname, French Guiana,
Ecuador, Peru, Brazil (Bahia).
Canavalia grandiflora Benth., Ann.
Wiener Mus. Naturgesch. 2: 135. 1837.
Liana or trailing or climbing vine. Humid
forests, but extending from forest edges into
open areas, including rocky outcrops, 50–100
m; Bolívar (Anacoco, Caicara, Represa Guri,
lower Río Caroní, Río Caura, Río Cuyuní,
Santa María de Erebato, Tumeremo), Ama-
zonas (Puerto Ayacucho). French Guiana,
Peru, Brazil (Goiás, Mato Grosso, Minas
Gerais, Pará, Rondônia).
Canavalia rosea (Sw.) DC., Prodr. 2: 404.
1825. —Dolichos roseus Sw., Prodr. 105.
1788.
Dolichos maritimus Aubl., Hist. Pl.
Guiane 765. 1775.
Canavalia maritima Thouars, J. Bot.
Agric. 1: 80. 1813.
Woody prostrate vine on beach, or twin-
ing, climbing over vegetation on the back
beach, from the drift line to ca. 100 m; Delta
Amacuro (Caño Güiniquina). Anzoátegui,
Aragua, Distrito Federal, Falcón, Sucre,
Zulia; Pantropics.
The name Canavalia maritima (Aubl.)
Thouars has frequently been used, but Thouars’s
epithet is here considered to be a new name
rather than a combination from Aublet’s epi-
thet. Thus, C. rosea is the correct name.
Canavalia sericophylla Ducke, Arq. Inst.
Biol. Veg. 4: 23. 1938.
Vine. Evergreen lowland forests, 100–200
m; Amazonas (Río Mawarinuma at base of
Sierra de la Neblina). Amazonian Peru, Bra-
zil (Amazonas).

13. CENTROLOBIUM Mart. ex Benth., Comm. Legum. Gen. 31. 1837.

by Velva E. Rudd

Trees to ca. 30 m tall, commonly buttressed at the base; bark grayish, smooth
or fissured; sap red. Leaves deciduous, alternate, odd-pinnate, 7–21-foliolate; leaf-
lets oblong, ovate, or elliptic, the lower surface with numerous reddish orange resin-
ous dots; stipules deltoid to broadly orbicular or rhomboid, often caducous; stipels
lacking. Bracts and bracteoles stipule-like but smaller. Flowers ca. 1–2 cm long in
terminal panicles. Calyx turbinate-campanulate with 4 subequal lobes, the vexillar
lobe emarginate or bipartite; petals yellowish, sometimes with red or violet mark-
ings. Stamens 10, monadelphous; anthers uniform, dorsifixed. Ovaries sessile or
short-stalked, 1–3(4)-ovulate; style filiform, usually persistent as a stout spine;
stigma small, terminal. Fruit samaroid, indehiscent with a large 1–3(4)-seeded
basal body and a large terminal, sterile wing, the body echinate with spines to 4 cm
long, the wings spatulate or cultriform, the persistent stylar spine terminal. Seeds
oblong-subreniform.
Panama, Colombia, Venezuela, Trinidad, Guyana, Ecuador, Peru, Brazil, Bo-
livia; ca. 6 species, 1 in Venezuela.

Centrolobium paraense Tul., Arch. Mus.
Hist. Nat. 4: 87. 1844. —Cartán.
Tree to 30 m tall. Evergreen lowland for-
ests, savannas. Panama, Colombia, Venezu-
ela, Trinidad (cultivated?), Guyana, Brazil; 2
varieties, both in the flora area.
The wood of Centrolobium paraense is
fine-grained and has beautiful shades of red-
dish orange. It is used for furniture and cabi-
net work as well as for general construction.
270 F ABACEAE

Fig. 239. Centrolobium paraense var. orinocense

 Centrosema 271

Key to the Varieties of C. paraense (Benth.) Pittier, Bol. Mus. Comerc.

1. Leaflets rounded to subcordate at the
base, subglabrous on the upper surface
or often with some pubescence along the
midvein, glabrous on lower surface at
maturity; flowers 12–15 mm long
.................................... var. orinocense
1. Leaflets subcordate at the base, some-
times revolute, tomentose on upper sur-
face, subglabrous on lower surface ex-
cept for tomentum on major veins; flow-
ers 15–17 mm long ....... var. paraense
C. paraense var. orinocense Benth. in
Mart., Fl. Bras. 15(1): 266. 1862,
“orenocense.” —Centrolobium orinocense
Venez. 1: 19. 1927 [1926]. —Balaústre,
Cartán.
Centrolobium patinense Pittier, J. Wash.
Acad. Sci. 5: 570. 1915.
100–300 m; Bolívar (Ciudad Bolívar,
Represa Guri area, La Prisión, middle Río
Caura, San Mateo, Serranía de Guayapo in
lower Caura basin). Anzoátegui, Aragua,
Guárico, Zulia; Panama, Colombia. ŠFig. 239.
C. paraense var. paraense. —Cartán.
100–300 m; Bolívar (El Manteco, La
Paragua, Represa Guri, Río Asa, mouth of
Río Paragua, Upata). Anzoátegui, Guárico,
Sucre, Zulia; Trinidad (cultivated?), Guyana,
Brazil (Roraima).

14. CENTROSEMA (DC.) Benth., Comm. Legum. Gen. 53. 1837, nom. cons. —Cli-
toria sect. Centrosema DC., Prodr. 2: 234. 1825.
Bradburya Raf., Fl. Ludonear 104. 1817.
by Paul R. Fantz
Scandent herbs, trailing to high-climbing, suffrutescent at base from perennial
xylopodium; pubescence of minutely hooked trichomes common (view at 20–30×);
aerial stems wiry to lianaceous, twining, angular-terete. Leaves alternate, odd-pin-
nate or occasionally digitate, 3-, 5-, or 7-foliolate, entire, stipulate, stipellate, peti-
olate; stipules ovate to deltoid, striate, persistent; stipels linear to linear-deltoid,
striate, persistent; petiolules subquadrangular. Inflorescences axillary, usually soli-
tary, pseudoracemose, few to several-flowered, crowded apically, often with one
flower open at a time; peduncle elongate, rachis straight to weakly flexuose, nodose;
pedicels paired at nodes; bracts crowded, in 3 series, striated, smaller than
bracteoles, outer pair largest; bracteoles paired at calyx base, striated, enfolding
buds, appressed to calyx and often obscuring calyx tube and upper and lateral teeth,
deciduous in fruit. Flowers resupinate, papilionaceous, showy, lilac to purple or
white with purplish veins. Calyx short-campanulate, usually persisting in early
fruiting stages before deteriorating, upper 2 lobes connate, lower ventral tooth elon-
gated, often conspicuous, extending from between bracteoles; standard complicate,
orbicular, emarginate, spurred or gibbous above claw, pubescent on outer surface;
wing petals falcate-obovate, auriculate above claw; keel petals incurved, clawed,
subequal to scarcely shorter than wings, ventral margin broad U-shaped. Stamens
pseudodiadelphous, vexillar stamen fused basally, filaments dilated apically. Pistil
enclosed in staminal sheath; ovary sessile, linear, appressed-pubescent; style
strongly incurved, broadly U-shaped, ± dilated apically, persistent as a beak in fruit;
stigma marginal at style apex, barbellate basally. Legume sessile, linear to occasion-
ally broadened, compressed and flat, beaked; valves costate, longitudinal veins near
each margin, occasionally winged, sutures slightly thickened, subseptate within and
between the seeds; dehiscence by spiral twisting of valves. Seeds transverse-oblong,
thick, numerous to occasionally few per pod.
272 F ABACEAE

New World tropics and subtropics, a single species extending into temperate
southern U.S.A., a few species introduced into Old World tropics; 34 species, 12 in
Venezuela, 10 of these in the flora area.

Key to the Species of Centrosema

1. Leaves digitate, rachis absent; leaflets 3, 5, or 7 ......................... C. venosum

1. Leaves pinnate, rachis 0.2–4 cm long; leaflets 3 ....................................... 2
2(1). Fruit 8–50 mm wide; flowers 4–6 cm long; petiolules 5–8 mm long; stipels
5–8 mm long; leaflets 6–12 cm wide; inflorescences 6–15 cm long ...... 3
2. Fruit 4–7 mm wide; flowers 1.5–4 cm long; petiolules 1–5 mm long; stipels
2–5 mm long; leaflets 0.5–6(–7) cm wide; inflorescences 1–8(–11) cm
long ......................................................................................................... 4
3(2). Fruit 8–13 mm wide, beak 10–15 mm long; calyx tube 4–5 mm long, teeth
obscure or ventral tooth to 2 mm long; bracteoles 8–16 × 6–9 mm;
pedicels 5–10 mm long; stipules 4–9 mm long .......................... C. plumieri
3. Fruit 35–50 mm wide, beak 25–40 mm long; calyx tube 6–9 mm long,
teeth 2–3 mm long with ventral tooth 4–5 mm long; bracteoles 15–20 ×
9–13 mm; pedicels 10–20 mm long; stipules 10–16 mm long .................
................................................................................................ C. triquetrum
4(2). Bracteoles 14–23 × 6–10 mm; bracts 6–10 mm wide; legume beak 18–27 mm
long ......................................................................................................... 5
4. Bracteoles 4–15 × 2–7 mm; bracts 1.5–5 mm wide; legume beak 3–16 mm
long ......................................................................................................... 6
5(4). Calyx tube 5–6 mm wide, ventral tooth 2.5–4 mm long, upper and lateral
teeth 1–2 mm long; fruit 55–90 × 4–4.5 mm, beak 16–27 mm long;
pedicels 5–11 mm long ........................................................ C. brasilianum
5. Calyx tube 6–10 mm wide, ventral tooth 5–8 mm long, upper and lateral
teeth 2–3 mm long; fruit 100–180 × 7–8 mm, beak 25–52 mm long;
pedicels 10–15 mm long ................................................. C. tetragonolobum
6(4). Calyx tube 2–3 mm long, 3–5 mm wide; fruit 3–4 mm wide, 5–10 mm long;
leaflets linear, 0.3–1 cm wide ................................................................ 7
6. Calyx tube 3–6 mm long, 5–10 mm wide; fruit 4–7 mm wide, (8–)10–20 mm
long; leaflets ovate to elliptic to rhombic to oblong, 1–7 cm wide ........ 8
7(6). Flowers 1–1.5 cm long; bracteoles 4–7 × 2–3 mm; calyx teeth subequal,
upper and lateral teeth 3–4 mm long, ventral tooth 4–6 mm long;
fruits 5–8 cm long, beak 3–7 mm long; leaflets with main lateral veins
14–19; rachis 0.5–1.2 cm long; stipules 5–6 mm long ........... C. pascuorum
7. Flowers 2–4 cm long; bracteoles 9–13 × 3–5 mm; calyx teeth unequal in
length, upper and lateral teeth 0.4–1 mm long, ventral tooth 5–8 mm
long; fruits 7–10 mm long, beak 10–15 mm long; leaflets with main lat-
eral veins 20–40; rachis nearly lacking, 0.1–0.3 mm long; stipules 3–4 mm
long .................................................................................... C. angustifolium
8(6). Bracteoles 10–15 mm long; bracts 8–10 mm long; calyx tube 8–10 mm
wide, 4–6 mm long, ventral tooth 9–15 mm long; leaf rachis 1.5–3.5 cm
long, leaflets commonly large, 6–15 cm long; robust vine or lianaceous
........................................................................................... C. macrocarpum
8. Bracteoles 6–10 mm long; bracts 3–7 mm long; calyx tube 5–7 mm wide,
 Centrosema 273

3–4 mm long, ventral tooth 5–10 mm long; leaf rachis 0.4–1.5(–2) cm

long, leaflets commonly smaller, 2–7(–10) cm long; slender vine ........ 9
9(8). Upper and lateral calyx teeth subequal to calyx tube, 2–4 mm long, ven-
tral tooth 5–8 mm long; fruit 6–7 mm wide, beak 8–10 mm long, valve
dehiscence 1–1.5 turns; bracts 6–7 mm long; pedicels 6–8 mm long in
flower, 8–13 mm long in fruit; petioles 3–4 mm long .................... C. molle
9. Upper and lateral calyx teeth longer than calyx tube, 6–8 mm long, ven-
tral tooth 8–10 mm long; fruit 4–5 mm wide, beak 10–15 mm long,
valve dehiscence 1.5–5 turns; bracts 3–4 mm long; pedicels 2–5 mm in
flower, 4–8 mm long in fruit; petioles 2–3 mm long .......... C. virginianum

Centrosema angustifolium (H.B.K.) Guyana, Suriname, French Guiana, Brazil,

Benth., Comm. Legum. Gen. 54. 1837,
non (H.B.K.) Benth. in Mart., Fl. Bras.
15(1): 129. 1859. —Clitoria angustifolia
H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 417.
1823 [1824]. —Bradburya angustifolium
(H.B.K.) Kuntze, Revis. Gen. Pl. 1: 164.
1891.
Perennial herbaceous vine with wiry stem
and linear leaves. Savannas, edges of semi-
deciduous forests, 200–800 m; Bolívar
(Altiplanicie de Nuria east of Miamo, north-
east of El Manteco, Represa Guri, Serra do
Sol on Brazilian border), Amazonas (Caño
Picure in Río Ventuari basin). Anzoátegui,
Apure, Guárico, Monagas; Central America,
Colombia, Guyana, Suriname, Brazil, culti-
vated in Peru.
Centrosema brasilianum (L.) Benth.,
Comm. Legum. Gen. 54. 1837. —Clitoria
brasiliensis L., Sp. Pl. 753. 1753.
—Bradburya brasilianum (L.) Kuntze,
Revis. Gen. Pl. 1: 164. 1891.
Clitoria angustifolia auct. non H.B.K.
1823 [1824]: sensu Centrosema angus-
tifolium (H.B.K.) Benth., Comm. Legum.
Gen. 54. 1837; Centrosema angusti-
folium (H.B.K.) Benth. in Mart., Fl.
Bras. 15(1): 129. 1859; Centrosema
brasilianum var. angustifolium (H.B.K.)
Amshoff, Meded. Bot. Mus. Herb. Rijks
Univ. Utrecht 52: 62. 1939.
Perennial herbaceous vine with narrow to
broad leaflets. Savannas, roadsides, forest
edges, 50–500 m; Delta Amacuro (near Los
Castillos de Guayana, Sacupana), Bolívar
(widespread across northern part of state,
Río Hacha), Amazonas (Puerto Ayacucho to
Samariapo). Apure, Distrito Federal, Mona-
gas, Nueva Esparta, Sucre, Zulia; Colombia,
Bolivia. ŠFig. 245.
Centrosema macrocarpum Benth., Ann.
Mag. Nat. Hist. ser. 1, 3: 436. 1839.
—Bradburya macrocarpa (Benth.)
Kuntze, Revis. Gen. Pl. 1: 164. 1891.
—Centrosema macrocarpon auct., ortho-
graphic error.
Liana or robust vine. Open areas and sec-
ondary regrowth of forests, sandy soils, 50–
300 m; Bolívar (Altiplanicie de Nuria, Río
Cuchivero, Río Orinoco), Amazonas (Isla
Ratón, Río Cataniapo). Anzoátegui, Aragua,
Cojedes, Zulia; Central America, Colombia,
Guyana, Ecuador, Peru, Brazil, Bolivia. ŠFig.
240.
Centrosema molle Mart. ex Benth., Comm.
Leg. Gen. 55. 1837; Ann. Wein. Mus.
Naturgesch. 2: 119. 1839.
Centrosema pubescens auct. non Benth.
1837: sensu Benth. in Mart., Fl. Bras.
15: 131. 1839; Barbosa-Fevereio, Rodri-
guésia 42: 184. 1977.
Perennial herbaceous vine. Open areas of
evergreen lowland forests, river banks, sa-
vannas, sandy areas, near sea level to 300 m;
Delta Amacuro (Río Orinoco), Bolívar
(Ciudad Bolívar, La Paragua, Río Cuyuní,
Río Pargueni off middle Orinoco), Amazonas
(Puerto Ayacucho to Samariapo, Santa
Bárbara del Orinoco). Widespread elsewhere
in Venezuela north of the Orinoco; Mexico,
Central America, West Indies, Colombia,
Suriname, French Guiana, Ecuador, Peru,
Brazil, Bolivia, Argentina, introduced into
tropical Africa and Asia.
This is the most commonly collected spe-
cies of Centrosema and has often been con-
fused with and misidentified as C. pubescens
(see Fantz, Sida 17: 321–332. 1996).
274 F ABACEAE

Fig. 240. Centrosema macrocarpum Fig. 241. Centrosema pascuorum

Fig. 242. Centrosema plumieri

 Centrosema 275

Fig. 243. Centrosema triquetrum

Fig. 244. Centrosema

venosum

Fig. 245. Centrosema brasilianum

276 F ABACEAE

Centrosema pascuorum Mart. ex Benth.,
Comm. Legum. Gen. 561. 1837. —Brad-
burya pascuora (Mart. ex Benth.)
Kuntze, Revis. Gen. Pl. 1: 164. 1891.
Suffrutescent herb with narrow leaves,
trailing to weakly climbing. Rocky area and
roadsides, ca. 800 m; Bolívar (near Santa
Elena de Uairén). Anzoátegui, Cojedes,
Guárico, Miranda, Monagas, Zulia; Mexico,
Central America, Ecuador, Brazil. ŠFig. 241.
Centrosema plumieri (Turpin ex Pers.)
Benth., Comm. Legum. Gen. 54. 1837.
—Clitoria plumieri Turpin ex Pers., Syn.
Pl. 2: 303. 1807. —Bradburya plumieri
(Turpin ex Pers.) Kuntze, Revis. Gen. Pl.
1: 164. 1891.
Liana or robust vine. Evergreen lowland
forests, disturbed forests, 50–200 m; Delta
Amacuro (Sacupana), Bolívar (La Unión in
middle Río Caura), Amazonas (Capibara,
Mavaca, Río Mawarinuma). Apure, Guárico,
Monagas, Portuguesa; Mexico, Central
America, West Indies, Colombia, Guyana,
Ecuador, Peru, Brazil, Bolivia, introduced
into tropical Africa. ŠFig. 242.
Centrosema tetragonolobum Schultze-
Kraft & R.J. Williams, Caldasia 16: 133.
1990.
Perennial herbaceous vine with narrow to
broad leaflets. Borders of gallery forests in
savanna zone between 4°–6° latitude on both
sides of the Río Orinoco, 50–200 m; Bolívar
(18 km southeast of Río Parguaza on the
road to Puerto Ayacucho). Amazonas (Sama-
riapo to north of Puerto Ayacucho). Colombia
(Vichada).
Centrosema triquetrum (Hoffmanns. ex
Benth.) Benth. in Benth. & Hook. f.,
Gen. Pl. 1(2): 528. 1865. —Platysema tri-
quetra Hoffmanns. ex Benth., Ann.
Wiener Mus. Naturgesch. 2: 122. 1839.
—Bradburya triquetra (Hoffmanns. ex
Benth.) Kuntze, Revis. Gen. Pl. 1: 164.
1891.
Liana or robust vine. Evergreen lowland
forests, riparian bamboo thickets, 50–200 m;
Delta Amacuro (Río Cuyubini), Amazonas
(Río Mawarinuma). Táchira; Colombia, Guy-
ana, Ecuador, Peru, Brazil (Amazonas, Goi-
ás, Maranhão, Pará). ŠFig. 243.
Centrosema venosum Mart. ex Benth. in
Mart., Fl. Bras. 15(1): 133. 1859.
—Bradburya venosum (Mart. ex Benth.)
Kuntze, Revis. Gen. Pl. 1: 164. 1891.
Perennial herb, stems creeping or trail-
ing, weakly twining. Open grassy savan-
nas, 50–100 m; Bolívar (between Caicara
and Puerto Ayacucho, Río Parguaza),
Amazonas (Canaripó, from north of Puerto
Ayacucho to Samariapo, Rincones, Río
Ventuari). Anzoátegui; Colombia, Brazil
(Bahia, Goias, Minas Gerais, Pará). ŠFig.
244.
Centrosema virginianum (L.) Benth.,
Ann. Wiener Mus. Naturgesch. 2:120.
1837. —Clitoria virginiana L., Sp. Pl.
753. 1753. —Bradburya virginianum
(L.) Kuntze, Revis. Gen. Pl. 1: 164. 1891.
Perennial herbaceous vine. Semide-
ciduous woods and savannas, 100–500 m;
Bolívar (Altiplanicie de Nuria, Upata to
Guasipati), Amazonas (near Puerto Aya-
cucho). Aragua, Distrito Federal, Miranda;
U.S.A., Mexico, Central America, West
Indies, South America to Argentina, intro-
duced into India, Reunión.

15. CHAETOCALYX DC., Prodr. 2: 243. 1825.

by Velva E. Rudd
Perennial vines, herbaceous or suffrutescent. Stems slender, terete, longitudi-
nally striate. Leaves alternate, odd-pinnate, 5–17-foliolate; leaflets oblong, elliptic,
ovate, obovate, or suborbicular; stipules deltoid or deltoid-ovate to lanceolate, at-
tached at the base; stipels absent. Bracts intergrading with the stipules, often cadu-
cous; bracteoles lacking. Flowers ca. 1.2–3 cm long, solitary or in axillary, racemose,
paniculate, or fasciculate inflorescences. Calyx campanulate with 5 subequal lobes
or teeth, base symmetrical or asymmetrically gibbous, often setose with glandular
trichomes; petals cream-white to yellow, sometimes marked with red or violet, pu-
bescent or glabrous. Stamens 10, monadelphous; anthers uniform, elliptic,
 Chaetocalyx 277

Fig. 246. Chaetocalyx scandens var. pubescens

dorsifixed. Ovaries sessile or stipitate, 6–16-ovulate, glabrous or pubescent; stigma

terminal, minute, capitate. Fruit a loment, linear to subelliptic, laterally com-
pressed or subterete, longitudinally striate or reticulate-striate, to 16-articulate, the
segments approximately isometric to several times longer than wide. Seeds reni-
form-rod-shaped, sublustrous, brownish, the hilum elliptic, subterminal.
Mexico, Central America, West Indies, Colombia, Venezuela, Trinidad, Suri-
name, Ecuador, Peru, Brazil, Bolivia, Paraguay, Argentina, Uruguay; 14 or 15 spe-
cies, 1 in Venezuela.

Chaetocalyx scandens (L.) Urb., Symb. —Chaetocalyx pubescens DC., Prodr. 2:

Antill. 2: 292. 1900. —Coronilla scan-
dens L., Sp. Pl. 743. 1753.
Herbaceous vine; leaflets densely pubes-
cent to subglabrous on one or both surfaces;
calyx and fruit puberulent. Stream banks,
open woods, savannas, rocky hillsides.
Southern Mexico, Dominican Republic, Les-
ser Antilles, northern Colombia, Venezuela,
eastern Brazil; 2 varieties, both in Venezu-
ela, 1 of these in the flora area.
The other variety, Chaetocalyx scandens
var. scandens, is glabrous or subglabrous and
is found in Venezuela along the Coastal Cor-
dillera in Aragua, Carabobo, and Distrito
Federal.
C. scandens var. pubescens (DC.) Rudd,
Contr. U.S. Natl. Herb. 32: 236. 1958.
243. 1825.
Chaetocalyx vestita Standl., Field Mus.
Nat. Hist., Bot. Ser. 8: 14. 1930.
Chaetocalyx fissa Pittier, Bol. Soc. Venez.
Ci. Nat. 6: 189. 1940.
Chaetocalyx magniflora Pittier, Bol. Soc.
Venez. Ci. Nat. 6: 186. 1940.
Chaetocalyx nigrescens Pittier, Bol. Soc.
Venez. Ci. Nat. 6: 188. 1940.
Chaetocalyx paucifolia Pittier, Bol. Soc.
Venez. Ci. Nat. 6: 185. 1940.
Chaetocalyx perglandulosa Pittier, Bol.
Soc. Venez. Ci. Nat. 6: 187. 1940.
Woody, rocky slopes, savannas, 300–700
m; Bolívar (Altiplanicie de Nuria). Venezu-
elan Coastal Cordillera; southern Mexico,
Dominican Republic, Lesser Antilles, north-
ern Colombia, eastern Brazil. ŠFig. 246.
278 F ABACEAE

16. CLATHROTROPIS Harms in Dalla Torre & Harms, Gen. Siphon. 221. 1901.
by Charles H. Stirton and Gerardo A. Aymard C.
Tall trees. Leaves 5- or 7(9)-foliolate, odd-pinnate, alternate; leaflets char-
taceous or coriaceous, opposite, the upper surface dark green, the lower surface
paler; stipules small, caducous. Inflorescences terminal, racemose-paniculate. Flow-
ers white with purple nectar patch, fragrant, bisexual, zygomorphic, pedicellate,
bracteate and bracteolate. Calyx unequally 5-toothed, upper pair fused for 1/4–1/2
their length; petals clawed, ± auriculate; standard orbicular to obovate, glabrous;
wing petals ± same length as keel petals; sculpturing present, lamellate. Stamens
10, free or nearly so; filaments subequal; anthers uniform, oblong, versatile. Ovary
subsessile or subsessile, 4- or 5-ovulate, hairy; style filiform; stigma small, terminal.
Fruit dehiscent, compressed to flattened, sutures often thickened and dilated,
valves woody or coriaceous, glabrous or densely golden-pubescent, elliptic, oblong,
trapezoid or semireniform. Seeds few, large, compressed; testa fragile.
Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil; 6 spe-
cies, 5 in Venezuela, all in the flora area.

Key to the Species of Clathrotropis

1. Lower surface of leaves minutely pubescent; fruits densely velutinous,

with thickened and dilated sutures ...................................................... 2
1. Lower surface of leaves glabrous; fruits glabrous, without prominent su-
tures ....................................................................................................... 3
2(1). Lower surface of leaves glaucous; fruits crispate, copper-colored, shortly
ridged along upper sutures ................................................ C. glaucophylla
2. Lower surface of leaves green; fruits densely velutinous, rusty brown,
strongly ridged along upper sutures ................................... C. macrocarpa
3(1). Leaflets ovate-oblong to sublanceolate, apex attenuate, the secondary and
tertiary venation scarcely visible on both surfaces; flowers borne in
dense short panicles; fruits < 3.5 cm long ..................................... C. nitida
3. Leaflets broadly elliptic to oblong-elliptic, apex rounded to short-acumi-
nate, the secondary and tertiary venation prominently reticulate on
both surfaces; flowers borne in large lax panicles; fruits > 3.5 cm long
................................................................................................................ 4
4(3). Young branches, inflorescences, and buds finely silvery to yellowish
white-pubescent ................................................................. C. brachypetala
4. Young branches, inflorescences, and buds densely chocolate brown pubes-
cent ............................................................................................. C. brunnea

Clathrotropis brachypetala (Tul.) Klein.,
Recueil Trav. Bot. Néerl. 22: 398. 1925.
—Diplotropis brachypetala Tul., Arch.
Mus. Hist. Nat. 4: 111. 1844. —Caica-
reño, Mahomo montañero, Montanallari
(Warao).
Tree to 30 m tall; leaves 5- or 7-foliolate,
broadly elliptic to oblong-elliptic, glabrous,
nitid, to ca. 25 cm long; fruits dark, glabrous,
flattened; seeds to 5 cm long and 1.5 cm
thick. Evergreen lowland and riparian for-
ests, near sea level to 300 m; Delta Amacuro
(Caño Joba-Suburu west of Caño Guayo and
east of Caño Sacupana, Río Cuyubini),
Bolívar (Jabillal on Río Caura, Río Ariza, Río
Cuyuní between Isla Anacoco and Acarabisi),
Amazonas (23 km northeast of Puerto
Ayacucho near Cachama). St. Vincent, Trini-
dad, Guyana, Suriname. ŠFig. 247.
 Clathrotropis 279

Fig. 247. Clathrotropis brachypetala

280 F ABACEAE

Clathrotropis brunnea Amshoff, Acta Bot.
Neerl. 17: 103. 1968. —Kajadi (Ye-
kwana), Ma’nour’uruwe’dasu (Piaroa).
Clathrotropis colombiana Yakovlev,
Nauch. Dokl. Near Sch. Biol. Nauki 1:
58. 1971.
Tree to 40 m tall; leaves (3)5(7)-foliolate,
oblong to obovate-oblong, pubescent on vena-
tion, otherwise glabrescent, coriaceous or
chartaceous; inflorescences and buds densely
chocolate brownish-pubescent; fruits dark
blue. Evergreen lowland forests, 100–300 m;
Bolívar (Río Tabaro off Río Nichare),
Amazonas (Caño Iguana, Caño Mosquito off
Caño Marieta). Colombia, Guyana, Suri-
name.
White phloem tissue of Clathrotropis
brunnea is rubbed on insect stings to relieve
pain.
Clathrotropis glaucophylla R.S. Cowan,
Bol. Soc. Venez. Ci. Nat. 15: 99. 1954.
—Cabari, Guapocoji (Yanomami), Kajadi
(Yekwana).
Tree 30–40 m tall; leaves 3- or 5-foliolate,
elliptic, obovate to oblong-obovate, the lower
surface minutely puberulent and glaucous;
fruits crispate, light copper-colored, shortly
ridged along upper sutures. Evergreen low-
land to lower montane forests, 100–600 m;
Bolívar (Amaruay-tepui, Aparamán-tepui,
Auyán-tepui, Río Cácaro, upper Río Caura,
upper Río Erebato, Río Icabarú), Amazonas
(Cerro Huachamacari, Maroa, Río Coromoto,
Río Cuao, Río Ocamo, Río Padamo, near San
Carlos de Río Negro, Yavita). Guyana.
Clathrotropis macrocarpa Ducke, Trop.
Woods 31: 16. 1932. —Barbasco, Cabarí.
Tree to 40 m tall; leaves 5- or 7-foliolate,
obovate, coriaceous; fruits velutinous,
strongly ridged along upper sutures. Ever-
green lowland to montane, nonflooded for-
ests, 100–900 m; Bolívar (widespread), Ama-
zonas (slopes of Cerro Huachamacari, Río
Cataniapo, Río Cuao, Río Mawarinuma, San
Carlos de Río Negro, Sierra de la Neblina,
Solano road). Colombia, Guyana, Brazil
(Amazonas, Roraima).
Clathrotropis nitida (Benth.) Harms, Bot.
Jahrb. Syst 33: 27. 1903. —Diplotropis
nitida Benth., J. Bot. (Hooker) 2: 71.
1841. —Chimako, Guacapú.
Tree to 20 m tall; leaves 5- or 7-foliolate,
ovate-oblong to sublanceolate, glabrous, coria-
ceous, nitid; fruits small, oval-oblong, gla-
brous. Riparian forests, 50–400 m; Amazonas
(Río Baría, Río Casiquiare between Culi-
macare and junction with Río Negro, Río
Guianía, Río Pacimoni, Río Siapa, Río Vena-
mo, Río Yatúa). Colombia, Brazil (Amazonas).

17. CLITORIA L., Sp. Pl. 753. 1753.

Ternatea Tourn. ex Mill., Gard. Dict. Abr. ed. 4, 3: Ternatea. 1754.
Neurocarpum Desv., J. Bot. Agric. 1: 119. 1813.
by Paul R. Fantz
Trees, shrubs, lianas, or subshrubs to perennial herbs or vines; pubescence of
minutely hooked trichomes common (view at 20–30×). Leaves alternate, odd-pin-
nate, 3-foliolate, petiolate to subsessile or 1-foliolate, sessile; stipules and stipels
persistent, striate, rarely caducous; petioles and rachis longitudinally striated,
sometimes canaliculate; petiolules subquadrate, rugose. Inflorescences axillary, ter-
minal or cauliferous, bearing chasmogamous or infrequently cleistogamous flowers,
1–many-flowered; peduncles usually solitary or several-fascicled; pedicels paired at
nodes; bracts striate, in 3 series below pedicels; bracteoles paired at calyx base, per-
sistent, striate, usually appressed to calyx, rarely borne below on pedicels.
Chasmogamous flowers resupinate, showy, papilionaceous, pink, blue to violet, or
white fading to pale yellow. Calyx funnel-shaped, persistent in fruit, 5-lobed, upper
2 lobes subconnate, lowermost narrower, often longer. Standard complicate, emar-
ginate, short-clawed, spurless, veins dark-colored, converging in throat, abaxial sur-
face pubescent. Wings long-clawed, extending beyond the falcate-incurved, short-
clawed keel. Stamens 10, diadelphous, often persistent in fruit. Pistil enclosed
 Clitoria 281

within staminal sheath; ovary stipitate; style geniculate, bearded longitudinally

apically. Cleistogamous flowers uncommon, inconspicuous unless with fruit; petals
lacking or remnants hidden in calyx; calyx funnel-shaped, small, persistent in fruit,
bracteolate, 5-lobed; pistil similar to that of chasmogamous flowers, but style bent
abruptly back toward base, in contact with anthers. Legume stipitate, linear,
straight to subfalcate, thickened at sutures, valves flat or convex, weak to strongly
depressed between seeds, ecostate or costate, beaked, spirally twisting upon dehis-
cence. Seeds dark brown to black, smooth or viscid.
Tropical to subtropical areas, mostly Neotropics, a few species in temperate ar-
eas; 60 species, 15 in Venezuela, 12 of these in the flora area.
The other three species in Venezuela are Clitoria fairchildiana Howard, a cul-
tivated Brazilian endemic that has been collected in Miranda state and in Ciudad
Bolívar; C. glaberrima Pittier, from Panama, rare elsewhere in Central America, in
Carabobo and Miranda states in Venezuela; and C. ternatea L., a cultivated and
naturalized African introduction found in Carabobo, Cojedes, Mérida, Sucre, and
Zulia states.

Key to the Species of Clitoria

1. Calyx 10-veined; fruits turgid, costate with medial longitudinal vein, 3.5–
6(–7) × 0.6–1.1 cm; stipe 7–12(–14) cm long, enclosed within calyx;
cleistogamy present (calyx 1–1.2 cm long); petiolules 2–5 mm long;
seeds viscid; subshrubs to perennial herbs, rarely vines ..................... 2
1. Calyx many-veined or striate; fruits flat, ecostate, (6–)8–24 × 1–2 cm;
stipe 15–37 mm long, exerted beyond calyx; cleistogamy absent; peti-
olules 4–10 mm long; seeds smooth; trees, shrubs, or lianas ............... 5
2(1). Leaves petiolate, petiole 2–8 cm long; inflorescence 5–20 cm long; stem
pubescence rufous; flowers white fading dull yellow; vines ....... C. falcata
2. Leaves subsessile to sessile, petiole to 1.5 cm long; inflorescence 0.5–7 cm
long; stem pubescence whitish; flowers pale bluish, lilac, or lavender;
subshrubs or perennial herbs, aerial stems erect ................................. 3
3(2). Leaves sessile, 1-foliolate; leaflets 3–7 mm wide ................... C. simplicifolia
3. Leaves subsessile, 3-foliolate; leaflets 1–3 mm wide ................................ 4
4(3). Calyx tube 16–22 mm long, 9–13 mm broad at throat, lobes 9–15 mm long;
bracteoles 8–14 mm long; flowers 5.5–7.5 cm long; petioles to 10 mm
long; lower surface of leaflets glabrate ................................. C. guianensis
4. Calyx tube 11–15 mm long, 6–9 mm broad at throat, lobes 6–9 mm long;
bracteoles 6–9 mm long; flowers 4–5.5 cm long; petioles 2–4 mm long;
lower surface of leaflets pubescent ......................................... C. laurifolia
5(1). Flowers 2.5–4 cm long, purple or white with purple marginally; calyx tube
4–7 mm wide at throat, lobes 2–4 mm long; valves of fruit depressed
between seeds (fruit unknown in C. steyermarkii); trees or shrubs .... 6
5. Flowers 4–8 cm long, lilac-purple to pinkish mauve; calyx tube 8–13 mm
wide at throat, lobes 4–13 mm long; valves of fruit not depressed be-
tween seeds; lianas or virgate shrubs ................................................... 8
6(5). Inflorescence to 0.5 cm long; flowers white with purple marginally; brac-
teoles linear, 3–5 × 0.3–0.5 mm; calyx pubescence uncinate; petiolules
4–6 mm long; leaflets 5–11 cm long ................................... C. steyermarkii
6. Inflorescence 1–3.5 cm long; flowers dull purple; bracteoles ovate, 1.5–3 ×
282 F ABACEAE

1–2.5 mm; calyx pubescence densely strigose; petiolules 8–10 mm long;

leaflets 10–20(–30) cm long ................................................................... 7
7(6). Calyx tube 7–11 mm long, lobes 2–3 mm long; bracteoles 1.5–3 mm wide;
leaflets broadly ovate to orbicular, 8–18 cm wide; trees .......... C. dendrina
7. Calyx tube 13–15 mm long, lobes 3–4 mm long; bracteoles 1–1.5 mm wide;
leaflets lanceolate-elliptic, 6–8 cm wide; shrubs .................... C. canescens
8(5). Lower surface of leaflets waxy, margins revolute; petiolules 2–4 mm long;
fruits 9–15 mm wide; fruit dehiscence twisting 1–1.3 turns; virgate
shrubs ......................................................................................... C. coriacea
8. Lower surface of leaflets not waxy, margins not revolute; petiolules
4–8 mm long; fruits 15–26 mm wide; fruit dehiscence twisting
0.2–0.5 turns; lianas .............................................................................. 9
9(8). Inflorescence 5–22 cm long; flowers 4–6 cm long, bluish violet; bracteoles
10–15 mm long; stipe 14–20 mm long; lower surface of leaflets rufous-
tomentose .............................................................................. C. arborescens
9. Inflorescence to 5 cm long; flowers 6–8 cm long, pink-rose to pinkish
mauve; bracteoles 2–10 mm long; stipe 24–37 mm long; lower surface of
leaflets glabrate ................................................................................... 10
10(9). Calyx tube 11–16 mm long, pubescence predominately uncinate; bracte-
oles 7–10 mm long; inflorescence to 0.5 cm long; staminal tube 3–4 cm
long; vexillar claw 6–9 mm long; stipels 1–3 mm long ................ C. sagotii
10. Calyx tube 16–24 mm long, pubescence strigose; bracteoles 2–7 mm long;
inflorescense 0.5–5 cm long; staminal tube 4–5 cm long; vexillar claw
14–19 mm long; stipels 3–7 mm long .................................................. 11
11(10). Style 16–25 mm long, subequal to ovary length; fruit pubescence rufo-
strigose; leaflets with 9–13(–15) pairs of lateral veins; inflorescences
1–5 cm long .............................................................................. C. javitensis
11. Style 24–32 mm long, ca. twice the ovary length; fruit pubescence unci-
nate; leaflets with 7–9 pairs of lateral veins; inflorescences 0.5–1 cm
long ........................................................................................ C. cavalcantei

Clitoria arborescens W.T. Aiton, Hortus
Kew ed. 2, 4: 302. 1814. —Ternatea
arborescens (W.T. Aiton) Kuntze, Revis.
Gen. Pl. 1: 210. 1891.
Clitoria poitaei DC., Prodr. 2: 234. 1825.
Clitoria amoena Miq., Stirp. Surinam. Se-
lect. 24. 1850 [1851].
Liana, occasionally scandent shrub. Open
forest areas, along forest margins of river
banks, 50–500 m; Delta Amacuro (Río Ama-
curo, Río Cuyubini, Río Orinoco, Serranía de
Imataca), Bolívar (widespread in northern
part of state, base of Auyán-tepui). Miranda,
Monagas, Sucre, Zulia; Colombia, Guyana,
Suriname, French Guiana.
Clitoria arborescens is not in Central
America as has been reported in the litera-
ture. These erroneous reports are based
upon misidentified specimens, usually Cli-
toria javitensis or Clitoria glaberrima.
This species has been cultivated for its
climbing habit and showy floral display. The
seeds are used as a fish poison in Guyana.
Clitoria canescens Pittier ex Fantz, Sida 8:
305. 1980.
Shrub. Sandy places along rivers, ca. 100
m; Amazonas (Río Guainía at Caño San
Miguel). Endemic.
Clitoria cavalcantei Fantz, Sida 9: 201.
1982.
Small shrub with a climbing apex or li-
ana. Riparian forests and clearings, 500–600
m; Bolívar (Río Curutu in upper Río Para-
gua). Brazil (Amazonas, Pará).
Clitoria coriacea Schery, Fieldiana, Bot.
28: 260. 1952.
Clitoria cerifera R.S. Cowan, Mem. New
York Bot. Gard. 9: 349. 1957.

Virgate shrub 1.5–2.5 m tall. White-sand
savannas and shrublands, 50–200 m; scat-
tered in western Amazonas. Endemic.
Clitoria coriacea will likely be found in
adjacent Colombia (Guianía).
Clitoria dendrina Pittier, Contr. U.S. Natl.
Herb. 20: 126. 1918. —Generala, Oreja
de tigre, Peonío de cerro.
Tree 2–11(–25) m tall. Open areas of semi-
deciduous forests, along forest borders, sa-
vannas, 50–400 m; Bolívar (Caño Asisa off
Río Parú, La Prisión, middle Río Orinoco, Río
Parguaza, east of Túriba), Amazonas (Caño
Asisa, Puerto Ayacucho). Aragua, Barinas,
Carabobo, Cojedes, Lara, Mérida, Portu-
guesa; Colombia. ŠFig. 248.
Clitoria falcata Lam., Encycl. 2: 51. 1786.
Clitoria rubiginosa Juss. ex Pers., Syn. Pl.
2: 303. 1807.
Clitoria glycinoides DC., Prodr. 2: 234.
1825.
Herbaceous vine. Mexico, Central Amer-
ica, West Indies, Colombia, Venezuela, Guy-
ana, Suriname, French Guiana, Ecuador,
Peru, Brazil, Bolivia, Paraguay, introduced
and naturalized in Africa and Asia; 4 variet-
ies, 1 in Venezuela.
Variety aurantiaca (Benth.) Fantz and
var. latifolia (Rizzini) Fantz are endemic to
Brazil; var. glabrescens (Verdc.) Fantz occurs
in Africa and is introduced into Guadeloupe
and Tobago.
C. falcata var. falcata
Savannas, forest borders, 50–1000 m;
Bolívar (Gran Sabana), Amazonas (between
Caicara and Puerto Ayacucho, near Yavita).
Widespread throughout most of Venezuela
north of the Orinoco; other distribution
neotropically as in species, and also intro-
duced into Africa (Gold Coast, Liberia, Nige-
ria) and Asia (Java, Taiwan). ŠFig. 253.
Clitoria guianensis (Aubl.) Benth., J. Proc.
Linn. Soc., Bot. 2: 40. 1858. —Crotalaria
guianensis Aubl., Hist. Pl. Guiane 761, t.
305. 1775. —Clitoria guyanensis Benth.
in Mart., Fl. Bras. 15(1): 121. 1862.
Subshrub to perennial, suffrutescent
herb 10–60 cm tall. Southern Mexico, Cen-
tral America, western Cuba, Colombia,
Venezuela, Brazil, Bolivia; 3 varieties, 1 in
Venezuela.
Clitoria 283
Clitoria guianensis is not in eastern Cuba
and elsewhere in the West Indies as has been
reported in the literature. These reports are
usually misidentified Clitoria laurifolia. The
varieties chapadensis (Malme) Fantz and
macrocleistogama Fantz are localized en-
demics in Brazil (Mato Grosso).
C. guianensis var. guianensis. —Generala.
Savannas, 200–800 m; Bolívar (wide-
spread), Amazonas (Río Ventuari basin).
Widespread in northern Venezuela; other
distribution as in species. ŠFig. 251.
Clitoria javitensis (H.B.K.) Benth., J. Proc.
Linn. Soc., Bot. 2: 42. 1858. —Neuro-
carpum javitense H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 409. 1823 [1824], emend.
Fantz, Sida 9: 162. 1981. —Ternatea
javitensis (H.B.K.) Kuntze, Revis. Gen.
Pl. 1: 210. 1891.
Liana. Tropical forests, usually associated
with sandy soil, 50–700 m; Costa Rica,
Panama, Colombia, Venezuela, Peru, Brazil;
4 varieties, 3 in Venezuela, 2 of these in the
flora area.
Variety portobellensis (Beurl.) Fantz is
distributed in Central America, Colombia,
and Venezuela (Carabobo, Miranda, Yara-
cuy). Variety longiloba Fantz is endemic to
Peru (Loreto).
Key to the Varieties of C. javitensis
1. Inflorescence 4–18 cm long, many-flow-
ered; petiole commonly 10–25 cm long,
petiolule 6–9 mm long; stipe 19–24 mm
long; leaflets ovate, to 28 cm long, 15 cm
wide ........................... var. grandifolia
1. Inflorescence 0.5–6 cm long, few-flowered;
petiole commonly 4–10 cm long, peti-
olule 4–6 cm long; stipe 25–37 mm long;
leaflets oblong-elliptic, to 18 cm long,
7(–9) cm wide ............... var. javitensis
C. javitensis var. grandifolia (Ducke)
Fantz, Sida 9: 170. 1981. —Clitoria
grandifolia Ducke, Arch. Jard. Bot. Rio
de Janeiro 5: 141. 1930.
100–500 m; Amazonas (Cerro Sipapo, Río
Cunucunuma). Colombia, Peru, Brazil.
C. javitensis var. javitensis
50–700 m; Bolívar (Río Caroní, Río Caura,
Río Paragua, Río Parguaza), Amazonas (wide-
284 F ABACEAE
spread). Amazonian Colombia, Brazil (Ama-
zonas, Rondônia). ŠFig. 250.
The typical form and f. bracteosubtenda
Fantz are known from the flora area. The
typical form has bracteoles 2–3 mm long,
subopposite to alternate on the pedicel, in-
serted 2.5–5 mm below the base of the calyx,
the apex below the calyx base or barely
reaching it. Forma bracteosubtenda has
bracteoles 3–5 mm long, subopposite, sub-
tending calyx base, inserted 1–2 mm below
the base of the calyx, the apex reaching the
swollen calyx base (represented internally by
the disk).
Clitoria laurifolia Poir. in Lam., Encycl.
supp. 2: 301. 1811.
Neurocarpum cajanifolium C. Presl.,
Symb. Bot. 1: 17, t. 9. 1830. —Clitoria
cajanifolia (C. Presl.) Benth. in Mart.,
Fl. Bras. 15(1): 121. 1862.
Clitoria parvifolia Pittier, Bol. Técn.
Minist. Agric. 5: 149. 1944, nom. nud.
Subshrub. Sandy soil in savannas and
along riverbanks, near sea level to 1300 m;
Delta Amacuro (Isla Las Cidras near Misión
San Francisco de Guayo), Bolívar (El Pal-
mar, Gran Sabana, Puerto Ordaz), Amazonas
(Río Negro). Anzoátegui, Apure, Zulia; West
Indies, Colombia, Guyana, Suriname,
French Guiana, Brazil, introduced and es-
tablished in eastern Africa and southeastern
Asia.
Clitoria laurifolia is not known from Cen-
tral America as has been reported in the lit-
erature. That material is usually mis-
identified Clitoria guianensis.
Three forms are found in the flora area:
(1) f. parvifolia with leaflets oblong-linear,
2.5–3.5 × 0.7–1.3 cm, petiole 1–1.5 mm long,
and rachis 7–13 mm long; (2) f. glabrior with
leaflets oblong to elliptic-oblong, 4–13 × 1.5–
3.5 cm, petioles 2–5 mm long, rachis 4–9 mm
long, and a legume with weakly raised costa,
extending 20–60% of the length of the valve,
or ecostate (often a mixture of ecostate and
costate legumes on same plant); and (3) f.
laurifolia with leaflets similar to f. glabrior,
but legume with prominently raised costa,
extending ca. 75–90% of the length of the
valve (otherwise all legumes similar).
Clitoria sagotii Fantz, Sida 8: 94. 1979.
Clitoria javitensis var. glabra Sagot, Ann.
Sci. Nat. Bot. sér. 6, 13: 299. 1882.
Neurocarpum javitense auct. non H.B.K.,
Nov. Gen. Sp. (quarto ed.) 6: 409. 1823
[1824]: sensu Clitoria javitensis (H.B.K.)
Benth., J. Proc. Linn. Soc., Bot. 2: 42.
1858.
Liana. Climber in forests or thickets, or
occasionally as trailing vine in open areas,
associated with sandy soil. Colombia, Ven-
ezuela, Guyana, Suriname, French Guiana,
Brazil; 3 varieties, 2 in Venezuela, both in
the flora area.
Variety sagotii is endemic to French
Guiana.
Key to the Varieties of C. sagotii
1. Petiole ± canaliculate adaxially; bracte-
oles 4–6 mm long, inserted 2–4 mm be-
low the calyx; petiolules 4–6 mm long;
main veins of leaflets 6–8 pairs ..............
................................. var. canaliculata
1. Petiole ± flattened adaxially; bracteoles 7–
10 mm long, inserted 1–2 mm below the
calyx; petiolules 6–9 mm long; main veins
of leaflets 8–10 pairs ....... var. sprucei
C. sagotii var. canaliculata Fantz, Sida 9:
351. 1982.
100–200 m; Bolívar (Río Cuyuní). Guy-
ana, Suriname. ŠFig. 249.
C. sagotii var. sprucei Fantz, Sida 9: 353.
1982.
100–200 m; Amazonas (Río Casiquiare,
Río Guainía, Río Negro, San Antonio on up-
per Río Orinoco). Colombia (Amazonas), Bra-
zil (Amazonas).
Clitoria simplicifolia (Kunth) Benth., J.
Proc. Linn. Soc., Bot. 2: 40. 1858.
—Neurocarpum simplicifolium Kunth,
Mimoses 213, t. 59. 1819 [1824]. —Ter-
natea simplicifolia (Kunth) Kuntze,
Revis. Gen. Pl. 1: 120. 1891.
Subshrub to perennial suffrutescent herb,
trailing to erect, 10–40 cm tall. Riparian for-
ests, disturbed areas, 50–200 m; Amazonas
(Isla Ratón, around Puerto Ayacucho, Río
Orinoco). Brazil (Goiás, Mato Grosso, Pará),
Paraguay. ŠFig. 252.
Clitoria steyermarkii Fantz, Ann. Mis-
souri Bot. Gard. 76: 1165. 1989.
Slender shrub to 2 m tall. Upland scrub,
ca. 700–800 m; Bolívar (Cerro Guaiquinima).
Endemic.
 Clitoria 285

Fig. 248. Clitoria dendrina

Fig. 249. Clitoria sagotii var. canaliculata

286 F ABACEAE

Fig. 250. Clitoria javitensis var. javitensis

Fig. 251. Clitoria guianensis var. guianensis

 Coursetia 287

Fig. 252. Clitoria simplicifolia Fig. 253. Clitoria falcata var. falcata

18. COURSETIA DC., Ann. Sci. Nat. (Paris) 4: 92. 1825.

by Nidia L. Cuello A.
Small trees or shrubs. Stems usually spindly, erect to scandent. Leaves pulvi-
nate, odd- or even-pinnate, or unifoliolate; petiolules shorter than the leaflets; leaf-
lets orbicular or narrowly to widely elliptic, mucronate, upper surfaces glabrate to
silky, lower surfaces pilose to silky or woolly, caducous; stipules small, subulate, per-
sistent; stipels present or absent. Inflorescences of pendent to erect, sessile or pe-
dunculate, axillary racemes, rarely panicles. Flowers ebracteolate; floral bracts
broadly lanceolate to linear-lanceolate, readily caducous or sometimes persistent.
Calyx campanulate, rounded or rarely attenuate at base, strigose to silky or tomen-
tose, or sometimes stipitate-glandular, the lobes ligulate, lanceolate to triangular,
acute to rounded; petals all yellowish brown, reddish brown, or purple; standard or-
bicular, emarginate, usually whitish to yellow and/or reddish, calluses and inflexed
auricles usually well developed; wing petals commonly protruding beyond the keel
at anthesis, parallel to the keel, whitish yellow to reddish; keels rostrate, the tip
acute. Ovary granuliferous to silky, villous, or woolly, rarely glabrous, sessile or
nearly so on the receptacle, 7–30-ovulate. Legumes glabrous to pubescent. Seeds
uniformly brownish, with darker purple mottling.
Neotropics (most diverse in Mexico and Central America); ca. 39 species, 4 in
Venezuela, 1 of these in the flora area.
288 F ABACEAE

Coursetia ferruginea (H.B.K.) Lavin in

Stirton, Adv. Legume Syst. 3: 63. 1987.
—Robinia ferruginea H.B.K., Nov. Gen.
Sp. (quarto ed.) 6: 395. 1823 [1824].
—Humboldtiella ferruginea (H.B.K.)
Harms, Repert. Spec. Nov. Regni Veg.
19: 12. 1923. —Majomo sabanero.
Fig. 254. Coursetia arborea Griseb., Fl. Brit. W. I. 2:
Coursetia
183. 1859.
ferruginea
Tree or shrub 2–3 m tall; flowers white to
pale lavender. Deciduous to evergreen low-
land forests or savannas, 100–400 m; Delta
Amacuro (Los Castillos), northern Bolívar,
Amazonas (Caño Asisa, Río Ocamo). Wide-
spread elsewhere in Venezuela; Panama, Co-
lombia, Trinidad, Guyana, Brazil (Roraima).
ŠFig. 254.
 Crotalaria 289

19. CROTALARIA L., Sp. Pl. 714. 1753; Gen. Pl. ed. 5. 320. 1754.
by Nidia L. Cuello A.
Herbs or shrubs, erect to prostrate. Stems pubescent or glabrous, smooth or
striate. Leaves ternately compound or unifoliolate by reduction; stipules varying
greatly in size and shape or lacking; leaflets entire, glabrous or pubescent. Inflores-
cences racemose, terminal or opposite the leaves; bracts setaceous to foliaceous;
bracteoles paired, similar in shape to the bracts, borne on the pedicel or the calyx.
Flowers sometimes showy, bisexual, 5-merous. Calyx green, bilabiate, the tube usu-
ally campanulate, glabrous or pubescent, the lobes usually longer to much longer
than the tube; corolla papilionaceous, usually yellow; standard orbicular or obovate,
the apex rounded or retuse, the base clawed, the adaxial surface and veins near the
base frequently becoming red-tinged; wing petals oblong, attached to a stalk con-
tinuous with the lower margin (cleaver-shaped), the blades usually puckered be-
tween some of veins; keel petals usually twisted at the tip, woolly-ciliate. Stamens
10, monadelphous, the staminal tube open on the upper side; anthers dimorphic
with basifixed, long anthers alternating with medifixed short anthers, the short an-
thers positioned above the elongate ones at anthesis. Ovary terminated by a genicu-
late style. Fruit inflated, subcylindric, sessile or short-stipitate, glabrous or pubes-
cent, the fruits of some species black at maturity. Seeds 7–46, oblique-cordiform;
testa smooth and mostly impermeable to water.
Tropics and subtropics, a few species in temperate areas; ca. 600 species, ca. 20
in Venezuela, 10 of these in the flora area.

Key to the Species of Crotalaria

1. Leaves 3-foliolate ....................................................................................... 2

1. Leaves 1-foliolate ....................................................................................... 4
2(1). Petioles equal to or longer than the terminal leaflet; bracts absent; fruits
villous to spreading-pubescent, trichomes often ca. 2 mm long .............
....................................................................................................... C. incana
2. Petioles shorter than the terminal leaflet; bracts present; fruit appressed-
pubescent to strigulose, trichomes usually < 0.5 mm long .................. 3
3(2). Leaflets narrow or linear; racemes elongate; bracts 1–3 mm long, almost
straight ................................................................................ C. maypurensis
3. Leaflets oblong to ovate-lanceolate; racemes short; bracts 5–10 mm long,
curled, silky-pubescent ................................................................ C. micans
4(1). Stipules usually absent, inconspicuous when present.............................. 5
4. Stipules present and conspicuous, or caducous ........................................ 7
5(4). Leaflets linear-lanceolate to lanceolate, 0.2–0.9 cm wide; stipules usually
absent, when present consisting of an inconspicuous, narrow (0.2–0.5 mm
wide) decurrent wing terminating at the base of the leaf without visible
lobes .......................................................................................... C. sagittalis
5. Leaflets obovate, obovate-elliptic, lanceolate, or oblanceolate, 0.5–5 cm
wide; stipules absent.............................................................................. 6
6(5). Leaflets obovate to obovate-elliptic, 1.5–5 cm wide; densely appressed-
pubescent on both sides; inflorescences 6–12 cm long; flowers 1–1.5 cm
long ................................................................................................. C. nitens
290 F ABACEAE

6. Leaflets lanceolate to oblanceolate, 0.5–1 cm wide; sparsely appressed pu-

bescent on both sides; inflorescences 1.5–5 cm long; flowers 0.5–0.8 cm
long ....................................................................................... C. aff. velutina
7(4). Stipules free; leaves oblanceolate .............................................................. 8
7. Stipules decurrent; leaves elliptic, obovate, lanceolate, or narrow-lan-
ceolate ..................................................................................................... 9
8(7). Stipules and bracts persistent, ovate, 5–15 mm long; calyx glabrous
................................................................................................ C. spectabilis
8. Stipules and bracts caducous, narrow, 2–3 mm long; calyx strigulose-
pubescent .......................................................................................C. retusa
9(7). Stipules decurrent, longer than the length of one internode (3–8 mm wide,
the width constant), this occurring throughout the length of the stem and
thus the entire stem appearing winged; stipule lobes usually not spread-
ing; inflorescences terminal or terminal and leaf-opposed .......... C. pilosa
9. Stipules decurrent, shorter than the length of one internode, usually only
along the uppermost internodes of the stem; stipule lobes spreading or
recurved; inflorescences not terminal, always leaf-opposed .............. 10
10(9). Leaflets obovate to ovate-elliptic, the lowermost usually obovate, the
uppermost ovate to ovate-elliptic; stipule lobes falcate or recurved
................................................................................................. C. stipularia
10. Leaflets all elliptic-lanceolate or linear-lanceolate; stipule lobes spread-
ing, sagittate, apiculate ........................................................... C. sagittalis

Crotalaria incana L., Sp. Pl. 716. 1753.
Herb or small shrub to 1.3 m tall; flowers
yellow. Roadsides, ca. 50 m; Delta Amacuro
(between San José and Clavellina, near
Tucupita), Bolívar (Caicara). Anzoátegui,
Apure, Aragua, Distrito Federal, Guárico,
Mérida, Miranda, Monagas, Nueva Esparta,
Portuguesa, Zulia; Mexico, Central America,
Colombia, Ecuador, Peru, Brazil (Bahia,
Pará), Bolivia, Argentina, Africa, Asia (Chi-
na, Philippines, Malaysia).
C. maypurensis var. depauperata (Mart.)
Windler & S.G. Skinner, Phytologia 50:
186. 1982. —Crotalaria depauperata
Mart., Fl. Bras. 15(1): 30. 1859.
—Generala.
Herb with ascending stem to 1 m tall;
flowers pale yellow. Swampy savannas, 50–
100 m; Bolívar (near Caicara, near Maripa,
near San Juan de Manapiare), Amazonas
(Río Ocamo, near San Juan de Manapiare).
Brazil (Minas Gerais). ŠFig. 255.

Crotalaria maypurensis H.B.K., Nov. Gen. C. maypurensis var. maypurensis

Sp. (quarto ed.) 6: 403. 1823 [1824].
—Generala.
Crotalaria lectophylla Benth., Ann. Nat.
Hist. 3: 430. 1839.
Crotalaria anagyroides var. pauciflora
Griseb., Cat. Pl. Cub. 69. 1866.
Neotropics; 2 varieties, both in the flora
area.
Key to the Varieties of C. maypurensis
1. Leaflets linear, the terminal leaflet usu-
ally 0.8–3.5 × 0.1–0.25 cm ......................
................................. var. depauperata
1. Leaflets oblanceolate, lanceolate, or ellip-
tic, the terminal leaflet usually 4.5–5.5 ×
0.7–1.5 cm .............. var. maypurensis
Shrub or suffrutescent herb; flowers yel-
low. Savannas, sandy open river banks, 50–
1000 m; Bolívar (widespread), Amazonas
(near Canaripó, Culebra, Puerto Ayacucho to
Samariapo, Santa Bárbara). Monagas, Zulia;
Mexico, Central America, Colombia, British
Guiana, Peru, Brazil, Paraguay, Argentina.
ŠFig. 256.
Crotalaria micans Link, Enum. Hort.
Berol. Alt. 2: 228. 1822.
Crotalaria anagyroides H.B.K., Nov. Gen.
Sp. (quarto ed.) 6: 404. 1823 [1824].
Suffrutescent herb or shrub to 3 m tall;
flowers yellow. Sandy savannas, 50–800 m;
Bolívar (Altiplanicie de Nuria, lower Río

Caroní, near Santa Elena), Amazonas (Is-
la Ratón, upper Río Orinoco). Barinas, Cara-
bobo, Distrito Federal, Lara, Mérida, Mi-
randa, Monagas, Portuguesa, Táchira, Sucre,
Trujillo, Zulia; Mexico, Central America, West
Indies, Colombia, Guyana, Suriname, Ecua-
dor, Peru, widespread in Brazil, Bolivia, Para-
guay, Argentina, Africa, Sri Lanka.
Crotalaria nitens H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 399. 1824.
Crotalaria bracteata Schltdl. & Cham.,
Linnaea 5: 575. 1830.
Crotalaria schiedeana Steud., Nomencl.
Bot. ed. 2, 1: 445. 1840.
Herb 0.5–2 m tall; flowers yellow. Dis-
turbed areas, 100–200 m; Amazonas (Ta-
yari along upper Río Orinoco). Barinas,
Mérida, Táchira; Mexico, Guatemala,
Honduras, Costa Rica, Colombia, Ecuador,
Peru, Bolivia.
Crotalaria pilosa Mill., Gard. Dict. ed. 8.
no. 2. 1768, non Crotalaria pilosa
Thunb. 1800, nec Crotalaria pilosa
Roxb. ex Mart. 1820. —Wareka
marakae-yó (Panare).
Crotalaria pterocaula Desv., J. Bot. Agric.
3: 76. 1814.
Crotalaria pilosa var. skutchii Senn,
Rhodora 41: 331. 1939.
Suffrutescent to 60 cm tall; flowers yellow.
Savannas, 100–400 m; Bolívar (Altiplanicie
de Nuria, near Ciudad Piar, near Corozal on
Río Maniapure, lower Río Caroní, near Santa
Elena de Uairén, Serranía de los Pijiguaos),
Amazonas (between Maguari and Río Paru-
cito, Río Negro). Barinas, Mérida, Portuguesa;
Central America, Antilles, Colombia, Guyana,
Suriname, French Guiana, Ecuador, Brazil,
Bolivia, Paraguay. ŠFig. 259.
The fruit of Crotalaria pilosa is used as a
rattle by the Panare Amerindians.
Crotalaria retusa L., Sp. Pl. 715. 1753.
—Anilillo, Maraca de cascabel.
Dolichos cuneifolius Forssk., Fl. Aegypt.-
Arab. 134. 1775. —Crotalaria cuneifolia
(Forssk.) Schrank, Syll. Pl. Nov. 2: 78.
1828.
Crotalaria retusifolia Stokes, Bot. Mat.
Med. 3: 516. 1812.
Herb or suffrutescent herb to 1 m tall;
flowers yellow with reddish brown on outside
of standard. Roadsides, sandy savannas, 50–
Crotalaria 291
400 m; Delta Amacuro (Pedernales, near
Tucupita), Bolívar (lower Río Caroní, middle
Río Caura, Tumeremo, near Upata). Anzo-
átegui, Apure, Barinas, Distrito Federal,
Falcón, Guárico, Miranda, Monagas, Portu-
guesa, Zulia; Central America, Colombia,
Guyana, Suriname, French Guiana, Ecua-
dor, Peru, Brazil, Africa, India, Sri Lanka,
China, Malesia, Philippines.
Crotalaria sagittalis L., Sp. Pl. 714. 1753.
Herb or suffrutescent herb 10–60 cm tall;
flowers yellow. Savannas, ca. 100 m;
Amazonas (La Esmeralda, near Puerto
Ayacucho). Apure, Barinas, Yaracuy; U.S.A.,
Mexico, Central America, Colombia, Ecua-
dor, Peru, Brazil (Goiás), Bolivia, Paraguay.
Crotalaria sagittalis is a polymorphic species
with the stipules varying in size and shape, 2–10
mm long or more, decurrent, sagittate, apiculate,
pubescent or glabrate, or lacking.
Crotalaria spectabilis Roth, Nov. Pl. Sp.
341. 1821.
Erect herb; flowers yellow. Disturbed ar-
eas, 50–200 m; Bolívar (vicinity of El Callao
and Tumeremo, lower Río Caroní). Barinas,
Monagas, Portuguesa; Panama, Colombia,
Trinidad, Peru, Brazil (Pará, São Paulo), na-
tive to India, naturalized in many tropical
(or occasionally temperate) localities in the
Old and New Worlds.
Crotalaria stipularia Desv., J. Bot. (Desvaux)
3: 76. 1814, spelling variant: Crotalaria
stipularis. —Ajonjolí sabanero.
Erect to creeping herb with silvery green
stems and leaves; flowers yellow. Savannas
with igneous outcrops, slopes, 50–300 m;
northern Bolívar. Anzoátegui, Aragua,
Distrito Federal, Guárico, Mérida, Miranda,
Monagas, Portuguesa, Sucre, Yaracuy; Co-
lombia, Guyana, Suriname, French Guiana,
Ecuador, Brazil, Bolivia, Paraguay, Argen-
tina. ŠFig. 257.
Crotalaria aff. velutina Benth., Ann. Nat.
Hist. 3: 429. 1839.
Erect herb to 60 cm long; flowers yellow.
Savannas, 50–300 m; Bolívar (road between
Caicara and Puerto Ayacucho). Apure; Bra-
zil. ŠFig. 258.
This taxon may represent a new species,
but more fruits and flowers are needed to de-
termine this with certainty.
292 F ABACEAE

Fig. 255. Crotalaria

maypurensis var.
depauperata

Fig. 256. Crotalaria maypurensis var. maypurensis

 Crotalaria 293

Fig. 258. Crotalaria

Fig. 257. Crotalaria stipularia aff. velutina Fig. 259. Crotalaria pilosa
294 F ABACEAE

20. CYMBOSEMA Benth., J. Bot. (Hooker) 2: 60. 1840.

by Richard H. Maxwell
Vines to 10 m long. Leaves alternate, 3-foliolate; stipules acute; leaflets broadly
lanceolate to oblong, entire. Inflorescence axillary, erect, pseudoracemose, un-
branched; bracts ovate, persistent. Flowers ca. 4 cm long, several on each node. Ca-
lyx 4-lobed; corolla reddish, drying purplish, petals with claws 4–7 mm long; stan-
dard obovate; wing petals oblanceolate, free; keel petals oblanceolate, fused distally.
Stamens 10, diadelphous, the vexillar one free. Ovary sessile, ca. 6-ovulate; stigma
and style glabrous. Fruit a legume, ca. 44 × 18 × 4 mm, oblong, usually ± falcate,
with a long, down-curved beak, glabrescent, ca. 4-seeded. Seeds ca. 10 × 6 × 4 mm,
hard, dark brown, the hilum nearly halfway encircling the seed.
Mexico, Costa Rica, Panama, Colombia, Venezuela, Amazonian Peru and Bra-
zil; 1 species.

Fig. 260. Cymbosema roseum

 Dalbergia 295

Cymbosema roseum Benth., J. Bot.
(Hooker) 2: 60. 1840. —Raudal Boba-
dilla.
Cymbosema apurense (H.B.K.) Pittier:
sensu Pittier, Bol. Soc. Venez. Ci. Nat. 7:
154. 1941, non Dioclea apurensis H.B.K.
1823 [1824].
Vine; flowers reddish (may change to
purple when dried). Mostly river banks and
forests, near sea level to 200 m; scattered in
Delta Amacuro, northern Bolívar, and Ama-
zonas. Apure, Miranda, Yaracuy; extra-Ven-
ezuelan distribution as in genus. ŠFig. 260.
The standard of the flowers is not re-
flexed, which suggests hummingbird pollina-
tion.

21. DALBERGIA L. f., Suppl. Pl. 52. 1781 [1782], nom. cons.
Amerimnon P. Browne, Civ. Nat. Hist. Jamaica 288. 1756.
Ecastaphyllum P. Browne, Civ. Nat. Hist. Jamaica 299. 1756.
by Gerardo A. Aymard C.
Trees or shrubs, sometimes with scandent branches, or woody lianas. Leaves
alternate, odd-pinnate, some species 1-foliolate; stipules ovate to subulate, small,
deciduous; stipels absent; leaflets usually alternate, 3–several. Inflorescences pan-
iculate, sometimes racemose, terminal or axillary. Flowers small, on short pedicels;
bracts and bracteoles deciduous. Calyx campanulate, the teeth 5, the 2 upper ones
broadest, the lowest one usually longest; petals subequal, clawed; standard gla-
brous, ovate, obovate, or orbicular; wing petals oblong; keel petals coherent along
the lower margin. Stamens 9 or 10, monadelphous or disposed in 2 fascicles of 5, dia-
delphous, or absent; anthers minute, basifixed, dehiscing by 2 horizontal slits.
Ovary stipitate, few-ovulate; style subulate, glabrous; stigma capitate or indetermi-
nate. Fruits stipitate, reniform to oblong or linear, occasionally constricted medially,
wingless or with the wing surrounding the seminiferous area, lightly reticulate, the
margin not thickened, indehiscent. Seeds 1 or rarely 2, reniform and compressed.
Pantropics; ca. 100 species, 16 in Venezuela, 15 of these in the flora area.

Key to the Species of Dalbergia

1. Leaves with 17–41 leaflets ......................................................................... 2

1. Leaves with 1–15 leaflets ........................................................................... 6
2(1). Leaves with 25–41 leaflets ........................................................... D. inundata
2. Leaves with 17–21 leaflets ......................................................................... 3
3(2). Leaflets 0.8–1 × 0.4–0.6 cm .......................................................... D. inundata
3. Leaflets 2–8 × 1–5 cm ................................................................................ 4
4(3). Leaflets oblong, the base rounded, apex obtuse; fruit with obvious reticu-
late venation ........................................................................... D. spruceana
4. Leaflets ovate, ovate-elliptic, or circular, the base acute, apex acuminate;
fruit without venation or with barely discernible reticulate venation
................................................................................................................ 5
5(4). Lower surface of leaflets appressed-pubescent; flowers dark violet; fruit
3–6 cm long, ovate, woody, not winged ........................................ D. foliosa
5. Lower surface of leaflets glabrescent or sparsely pilose; flowers purple
with white spots; fruit 7–10 cm long, oblong-elliptic, papery, winged
................................................................................................. D. spruceana
6(1). Leaves 1-foliolate ....................................................................................... 7
6. Leaves 2–15-foliolate ................................................................................. 9
296 F ABACEAE

7(6). Leaflets orbicular; pedicels 3.5–4 mm long ........................................ D. sp. A

7. Leaflets ovate or ovate-lanceolate; pedicels 1–2.5 mm long ..................... 8
8(7). Lower surface of leaflets not glaucous; calyx lobes unequal, the upper 2
fused; fruit oblong, glabrous .................................................. D. amazonica
8. Lower surface of leaflets glaucous; calyx lobes equal, the upper 2 free or
fused basally; fruit orbicular to ovate, sparsely pubescent ....................
.......................................................................................... D. ecastophyllum
9(6). Branches and branchlets glabrous or puberulent; lower surface of leaflets
glabrous or sparsely pilose .................................................................. 10
9. Branches and branchlets ferruginous-tomentose or yellowish-tomentose;
lower surface of leaflets pilose or densely pubescent ......................... 12
10(9). Leaflets orbicular or broadly elliptic; fruit orbicular ................. D. monetaria
10. Leaflets ovate, ovate-lanceolate, ovate-oblong, or obovate; fruit reniform
or oblong-elliptic .................................................................................. 11
11(10). Leaflets ovate, glabrous on upper surface, sparsely pubescent on lower
surface, the apex retuse; inflorescence not fasciculate and clustered in
the leaf axil, terminal or axillary; fruit oblong-elliptic, winged, mem-
branaceous .............................................................................. D. frutescens
11. Leaflets ovate-lanceolate, glabrous on both surfaces, the apex acuminate;
inflorescence ± fasciculate, clustered in the leaf axil; fruit reniform, co-
riaceous, not winged, not membranaceous ........................... D. hygrophila
12(9). Lower surface of leaflets appressed-pubescent or densely ferruginous-to-
mentose ................................................................................................ 13
12. Lower surface of leaflets pilose, or, if pubescent, pubescence not ferrugi-
nous-tomentose .................................................................................... 14
13(12). Lower surface of leaflets densely ferruginous-tomentose; calyx teeth
equal, 4–5 mm long; fruit reniform, suborbicular, or quite orbicular, not
winged, densely ferruginous-tomentose ..................................... D. riedelii
13. Lower surface of leaflets appressed-pubescent; calyx teeth unequal, 1.5–2
mm long; fruit oblong, winged, glabrous, or sparsely pubescent ...........
................................................................................................ D. intermedia
14(12). Leaflets widely ovate or orbicular ........................................................... 15
14. Leaflets oblong, obovate, ovate-elliptic, or obovate-oblanceolate ........... 16
15(14). Leaflets 3–6 cm wide, ovate, the upper surface shiny, the lower surface
appressed-pubescent ....................................................................... D. sp. B
15. Leaflets 7–12 cm wide, widely ovate or orbicular, chartaceous, the upper
surface dull, the lower surface completely appressed-pubescent ..........
......................................................................................................... D. sp. C
16(14). Lower surface of leaflets completely appressed-pubescent; calyx 1.2 mm
long; ovary densely pubescent ........................................................ D. sp. D
16. Lower surface of leaflets glabrous or sparsely appressed-pubescent; calyx
5–6 mm long; ovary glabrescent or pilose ........................................... 17
17(16). Leaflets elliptic-lanceolate; petals white; fruit reticulate ......... D. subcymosa
17. Leaflets ovate-elliptic, oblong, or obovate; petals lilac or purple; fruit not
reticulate ....................................................................................... D. foliosa
Dalbergia amazonica (Radlk. ex Köpff) Radlk. ex Köpff, Anat. Charakt. Dalberg.
Ducke, Leguminosas Amaz. Brasil. 122. 40. 1892. —Anicillo, Caraota rebalsera,
1939. —Ecastaphyllum amazonicum Kamatata (Yekwana).

Scandent shrub or small tree; flowers
white. Evergreen riparian lowland to lower
montane forests, 50–500 m; Bolívar (Río
Acanán, mouth of Río Botanamo, lower Río
Caroní, upper Río Caura, Río Cuyuní, Río
Erebato, mouth of Río Nichare, mouth of Río
Paragua), Amazonas (Puerto Ayacucho,
Raudal de Atures, lower Río Ventuari).
Apure; Colombia, Guyana, French Guiana,
Brazil (Amapá, Amazonas, Pará).
Dalbergia ecastophyllum (P. Browne ex
L.) Taub. in Engl. & Prantl, Nat.
Pflanzenfam. 3(3): 335. 1894. —Hedy-
sarum ecastaphyllum P. Browne ex L.,
Syst. Nat. ed. 10, 2: 1169. 1759. —Chi-
nay-yek (Arekuna), Bejuco saliva, Cara-
ota rebalsera.
Dalbergia brownei (Jacq.) Urb., Symb.
Ant. 4: 295. 1905. —Amerimnon brownei
Jacq., Enum. Syst. Pl. 27. 1760,
“brownii.”
Climbing shrub, liana, or small tree; pet-
als white. Evergreen lowland to montane for-
ests, flooded forests, 50–1300 m; Delta
Amacuro (Caño Güiniquina, near Peder-
nales), Bolívar (45 km north of Cerro
Impacto, Ciudad Bolívar, Puerto Ordaz, Río
Botanamo, Río Caura, Río Cuyuní, Río
Paragua, Río Tirica to Urimán, Sierra de
Maigualida), Amazonas (Isla Ratón, ca. 9 km
south of Puerto Ayacucho). Apure, Carabobo,
Guárico, Mérida, Miranda, Monagas, Sucre,
Táchira, Zulia; U.S.A. (southern Florida),
Central America, West Indies, Colombia,
Guyana, Suriname, French Guiana, Peru,
Brazil (Amazonas, Bahia, Distrito Federal,
Espirito Santo, Paraná, Rio de Janeiro),
West Africa.
Dalbergia foliosa (Benth.) A.M. Carvalho,
Brittonia 49: 100. 1997. —Ecasta-
phyllum foliosum Benth., J. Bot.
(Hooker) 2: 64. 1840. —Arepillo,
Jadewa-jadakwadiyo (Yekwana), Uay-
peu-roy-yek (Pemón).
Ecastaphyllum glaucum Desv., Ann. Sci.
Nat. (Paris) sér. 1, 9: 423. 1826. —Dal-
bergia glauca (Desv.) Amshoff, Meded.
Bot. Mus. Herb. Rijks Univ. Utrecht 52:
50. 1939, non Roxb. 1831, nec Kurz
1876.
Shrub with climbing branches; petals vio-
let. Evergreen lowland to lower montane for-
Dalbergia 297
ests, 100–500 m; Bolívar (El Foco near
Upata, Isla de Anacoco, La Escalera, Raudal
Cotua on Río Asa, Río Caroní, Río Nichare),
Amazonas (La Esmeralda, Río Casiquiare,
Río Pasimoni, Río Siapa, Río Sipapo).
Anzoátegui, Apure, Guárico; Colombia,
Guyana, Suriname, French Guiana, Peru,
Brazil (Amazonas, Bahia, Pará, Rio de
Janeiro), Bolivia, Paraguay, northern Argen-
tina. ŠFig. 261.
Dalbergia frutescens (Vell.) Britton, Bull.
Torrey Bot. Club. 16: 324. 1889. —Ptero-
carpus frutescens Vell., Fl. Flumin. 283.
1825 [1829]. —Sangrito.
Dalbergia variabilis Vogel, Linnaea 11:
196. 1837.
Neotropics; 2 varieties, 1 in Venezuela.
D. frutescens var. frutescens. —Bejuco de
estribo, Bejuco roblito, Sangrito.
Liana; flowers white. Deciduous to ever-
green lowland or lower montane forests, 50–
300 m; Bolívar (Puerto Ordaz, Represa Guri,
mouth of Río Paragua, Upata). Anzoátegui,
Apure, Falcón, Miranda, Guárico, Sucre,
Zulia; Peru, Brazil, Bolivia, Paraguay, north-
ern Argentina. ŠFig. 265.
Dalbergia hygrophila (Mart. ex Benth.)
Hoehne, Fl. Brasilica 25(3): 21. 1941.
—Ecastaphyllum hygrophilum Mart. ex
Benth., Comm. Legum. Gen. 29. 1837.
—Bejuco colorado.
Scandent shrub, small tree, or liana; flow-
ers white. Gallery forests, evergreen lowland
to lower montane forests, 50–500 m; Delta
Amacuro (Caño Araguabisi, Caño Güini-
quina), Bolívar (Río Acanán, lower Río
Caura), Amazonas (Caño Campana in Río
Atabapo basin, Caño Ucata southeast of
Síquita, Maroa, Río Baría, Río Cataniapo,
Río Mawarinuma, Río Orinoco). Anzoátegui,
Apure; Colombia, Peru, Brazil (Amazonas,
Pará). ŠFig. 264.
Dalbergia intermedia A.M. Carvalho,
Brittonia 49: 101. 1997. —Ecasta-
phyllum tomentosum Spruce ex Benth,
J. Linn. Soc. 4(supp.): 51. 1860.
—Dalbergia tomentosa (Spruce ex
Benth.) Ducke, Arch. Jard. Bot. Rio de
Janeiro 3: 144. 1982, non Vogel 1837,
nec (Benth.) Taub. 1891.
298 F ABACEAE
Liana; flowers white. Evergreen lowland
forests, 50–500 m; Bolívar (Río Zariapo on
upper Río Caura, Sierra de Maigualida),
Amazonas (Isla Ratón, Puerto Ayacucho).
Apure; Guyana, Brazil, Bolivia. ŠFig. 263.
Dalbergia inundata Spruce ex Benth., J.
Proc. Linn. Soc., Bot. 4(suppl.): 49. 1860.
—Drepanocarpus paludicola Standl.,
Trop. Woods 33: 12. 1933. —Uña de
gato.
Dalbergia aturensis Pittier, Bol. Soc.
Venez. Ci. Nat. 8: 260. 1943.
Shrub with climbing branches, small tree,
or shrub; flowers white and purple. Gallery
and evergreen lowland forests, flooded for-
ests, forests bordering white-water and
black-water rivers, 100–200 m; Amazonas
(widespread). Anzoátegui; Colombia, Guy-
ana, Peru, Brazil (Amazonas, Minas Gerais,
Pará). ŠFig. 262.
Dalbergia monetaria L. f., Suppl. Pl. 317.
1781 [1782]. —Eburu, Oteratana, Otero,
Realito, Tu-basabosa.
Climbing shrub, small tree, or liana; pet-
als white. Semideciduous to evergreen low-
land forests, lower montane forests, gallery
forests, flooded forests along black-water riv-
ers, 50–1000 m; Delta Amacuro (wide-
spread), Bolívar (Canaima, Cerro Guai-
quinima, Río Karaurín, upper Río Paragua,
Río Parupa, San Felíx), Amazonas (Caño San
Miguel, Maroa, Puerto Ayacucho, Río
Atabapo, Río Baria, Río Casiquiare, Río
Mawarinuma, upper Río Orinoco, Río Yudi).
Falcón, Miranda, Zulia; Central America,
West Indies, Colombia, Guyana, Suriname,
French Guiana, Ecuador, Peru, Amazonian
Brazil. ŠFig. 268.
This species has also sometimes been
called Dalbergia volubilis Urb. (Repert. Spec.
Nov. Regni Veg. 16: 136. 1919), but that
name was incorrectly based on Securidaca
volubulis L. (Species Plantarum 707. 1753),
Polygalaceae.
Dalbergia riedelii (Benth.) Sandwith, Bull.
Misc. Inform. 1931: 358. 1931.
—Ecastaphyllum monetaria var. riedelii
Benth., J. Proc. Linn. Soc., Bot.
4(suppl.): 50. 1860. —Ecastaphyllum
riedeli (Benth.) Radlk. in Köpff, Anat.
Charakt. Dalberg. 41. 1892.
Dalbergia pachycarpa Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 145. 1922.
Liana or climbing shrub; petals white. Ev-
ergreen lowland to montane forests, 50–1000
m; Bolívar (Caño Chiviripa near Río
Maniapure, La Escalera), Amazonas (Cerro
Cariche, La Esmeralda, Macuruco, Río
Asisa, Río Atabapo, Río Casiquiare, Santa
Barbara del Orinoco). Colombia, Guyana,
Suriname, French Guiana, Peru, Brazil
(Acre, Amazonas, Pará, Roraima). ŠFig. 266.
Dalbergia spruceana Benth., J. Proc.
Linn. Soc. Bot. 4(suppl.): 35. 1860.
Tree or climbing liana; flowers white. Ev-
ergreen lowland or seasonally dry forests,
100–300 m; Bolívar (El Pilón near Cerro
Marimarota, Río Parguaza), Amazonas
(Cerro Yutajé, Río Coro Coro). Brazil (Acré,
Amapá, Amazonas), Bolivia. ŠFig. 267.
Dalbergia subcymosa Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 144. 1922.
Liana. Evergreen lowland forests, near
sea level to 100 m; Delta Amacuro (San Vic-
tor near Guyana border). Suriname, French
Guiana, Peru, Brazil.
Dalbergia sp. A
Liana; flowers white. Granitic outcrops,
evergreen lowland forests, ca. 100 m; Bolívar
(Jabillal on Río Caura).
Dalbergia sp. A is based on Fernández
5267 (MO, PORT).
Dalbergia sp. B
Liana; flowers white. Deciduous forests,
50–100 m; Bolívar (Puerto Ordaz to San
Felíx).
Dalbergia sp. B is based on Aristeguieta
5314 (MO, VEN).
Dalbergia sp. C
Liana; flowers white. Gallery forests,
100–200 m; Amazonas (Río Mawarinuma).
Dalbergia sp. C is based on Davidse &
Miller 27422 (MO, NY, VEN).
Dalbergia sp. D. —Wakado (Yekwana).
Tree; flowers yellow. Evergreen forests,
1200–1300 m; Amazonas (upper Río Ven-
tuari).
Dalbergia sp. D is based on Chaviel 417
(MO, PORT).
 Dalbergia 299

Fig. 261. Dalbergia foliosa Fig. 262. Dalbergia inundata

Fig. 263. Dalbergia intermedia

300 F ABACEAE

Fig. 264. Dalbergia hygrophila Fig. 265. Dalbergia frutescens

var. frutescens

Fig. 266. Dalbergia riedelii

 Dalbergia 301

Fig. 267. Dalbergia spruceana

Fig. 268. Dalbergia monetaria

302 F ABACEAE

22. DERRIS Lour., Fl. Cochinch. 432. 1790, nom. cons.

by Hans-Helmut Poppendieck
Woody vines. Leaves alternate, odd-pinnate; stipules caducous, stipels absent,
leaflets opposite, entire. Inflorescence a lateral or terminal pseudoraceme composed
of short flower-bearing shoots to 3 mm long (to 10 mm long and secund in D.
amazonica); bracts caducous; bracteoles 2, borne on the middle of the pedicel. Calyx
dentate to truncate, cup-shaped; corolla white to light yellow (purple in D. ama-
zonica); standard ovate, clawed, not reflexed, the base attenuate, without callosities,
the apex obtuse (reflexed and truncate with basal callosities in D. amazonica); wing
petals adhering to keel by pressure and structural conformity; keel petals coherent
or connate; wing and keel petals about equal. Vexillar stamen free at base; anthers
uniform; nectary glands indistinct. Ovary pubescent; ovules 2–4; style glabrous.
Fruit indehiscent, flat, 1- or 2-seeded, with submarginal wings on either side close to
the vexillar margin.
Colombia, Venezuela, Guyana, Suriname, French Guiana, northern Amazo-
nian Brazil, Peru; 3 species, all in the flora area.
See Lonchocarpus for a discussion of the taxonomic problems between these
two genera.

Key to the Species of Derris

1. Corolla deep lavender, the standard with a light green to white spot at its
base and with basal callosities; short flower-bearing shoots 5–10 mm
long, secund; indumentum when present shortly sericeous, petiolule
usually whitish-sericeous ...................................................... D. amazonica
1. Corolla white to light yellow, the base of standard narrowed and without
callosities; short flower-bearing shoots 1–3 mm long, indumentum
when present brownish-rufescent ......................................................... 2
2(1). Legumes ovate .............................................................................. D. negrensis
2. Legumes oblong to narrowly oblong ........................................ D. pterocarpus

Derris amazonica Killip, J. Wash. Acad.
Sci. 24: 48. 1934. —Lonchocarpus ne-
grensis Benth., J. Proc. Linn. Soc., Bot.
4(suppl.): 98. 1860, non Derris negrensis
Benth. 1860.
Vine to 30 m high. Lower montane forests
on poor soil, 600–800 m; Amazonas (northern
slopes of Cerro Duida). Guyana, Suriname,
French Guiana, northern Amazonian Brazil.
ŠFig. 270.
The isolated position of this species within
Derris is evident from the generic description
and has already been noted by G. Amshoff
(Meded. Bot. Mus. Herb. Rijks Univ. Utrecht
52: 59. 1939). It agrees with Lonchocarpus
subgenus Phacelanthus Pittier in most char-
acters except for the winged fruit. For fur-
ther discussion, see Lonchocarpus.
Derris negrensis Benth., J. Proc. Linn.
Soc., Bot. 4(suppl.): 98. 1860.
Lonchocarpus killipii Ducke, Trop. Woods.
69: 5. 1942.
High-climbing, robust liana to 20 m high;
leaves coriaceous; flowers white, fragrant.
Flooded forests, occasionally in disturbed but
nonflooded evergreen lowland forests, 50–
300 m; Delta Amacuro (Sacupana), Bolívar
(lower Río Caura, Río Parguaza), Amazonas
(Río Casiquiare, Río Cunucunuma, Río Ne-
gro, Río Orinoco, Río Ventuari). Amazonian
Brazil (Rio Negro basin). ŠFig. 269.
This species is very similar to Derris
pterocarpus when in flower, but differs from
it by the fruit shape character presented in
the key.
 Derris 303

Fig. 269. Derris

negrensis

Fig. 270. Derris amazonica

304 F ABACEAE

Derris pterocarpus (DC.) Killip, J. Wash.

Acad. Sci. 26: 360. 1936. —Lonchocarpus
pterocarpus DC., Prodr. 2: 260. 1825.
Deguelia scandens Aubl., Hist. Pl. Guiane
750, t. 300. 1775. —Derris scandens
(Aubl.) Pittier, Contr. U.S. Natl. Herb.
20: 41. 1917; Pittier, Legumin. Venez. 1:
105. 1944, non Derris scandens Benth.,
J. Proc. Linn. Soc., Bot. 4(suppl.): 106.
1860.
Derris guyanensis Benth., J. Proc. Linn.
Soc., Bot. 4(suppl.): 106. 1860.
Glabrous liana, reaching high into the for-
est canopy. Flooded forests along rivers, dis-
turbed forests on higher ground, 50–300 m;
Bolívar (El Manteco, Puerto Ordaz, Serranía
de Imataca). Apure; Guyana, Suriname,
French Guiana, northern Amazonian Brazil.
ŠFig. 271.

Fig. 271. Derris pterocarpus

23. DESMODIUM Desv., J. Bot. Agric. 1: 112, pl. 5. 1813, nom. cons.
Meibomia Heist. ex Fabr., Enum. 168. 1759.
by Nidia L. Cuello A.
Herbs, subshrubs, or shrubs, prostrate, decumbent or scrambling to erect.
Leaves 1–3(–5)-foliolate, petiolate, stipulate; leaflets petiolulate and stipellate. In-
florescences axillary or terminal (fasciculate and leaf-opposed in D. triflorum), race-
mose or racemose-paniculate, or extremely dense and capitate; primary bracts stri-
ate and ciliate, each subtending 1 flower or a fascicle of 2–several flowers; secondary
bracts often present and similar to the primary bracts or depauperate, each sub-
tending a single flower; bracteoles rarely if ever present in American species. Flow-
ers pedicellate. Calyx 2-lobed, the upper lobe almost entire to bifid, the lower 3-
toothed with the central tooth longer than the laterals, or the calyx almost equally 5-
lobed; corolla exceeding the calyx; standard short-clawed and the wing petals
subsessile; wing petals often attached to keel petals by a small appendage; keel pet-
als long-clawed, partially fused above. Vexillar stamen free or partially fused; an-
thers elliptic. Ovary sessile or stipitate; ovules 2–many; style slender; stigma termi-
nal. Fruit a loment, transversely septate into 1–many articles, stipitate, indehiscent,
or tardily dehiscent, the articles nearly linear to almost saccate or circular and
notched at the apex, sometimes folded on each other in accordion fashion, the sur-
faces glabrous to densely pubescent with straight or hooked trichomes, or pubescent
only on the sutures, 1-seeded. Seeds oblong or subquadrate.
North America, tropics and subtropics (most diverse in eastern Asia, Mexico,
and Brazil), temperate Asia; ca. 300 species, ca. 25 in Venezuela, 17 of these in the
flora area.
 Desmodium 305

Key to the Species of Desmodium

1. Leaves all 1-foliolate, appearing simple .................................................... 2

1. Leaves all 3-foliolate, or both 3-foliolate and 1-foliolate on the same plant
................................................................................................................ 5
2(1). Leaflets linear to lance-ovate, narrowed to ± acute apex, rounded at base,
4–9.5 × 0.4–1.4 cm, prominently reticulate and soft-puberulent
on lower surface ............................................................... D. sclerophyllum
2. Leaflets obovate, ovate, or elliptic, the blades wider, shorter or longer
than above .............................................................................................. 3
3(2). Leaflets 4–8 cm wide, obovate to broadly ovate; inflorescence (2–)many-
branched; erect shrub > 1 m tall ............................................ D. distortum
3. Leaflets 1–3.5 cm wide, ovate to elliptic-lanceolate; inflorescence 1–3-
branched; creeping to ascending herb or suffrutescent, < 1 m tall ...... 4
4(3). Leaflets mostly ovate, the lowermost sometimes oblong to oblong-lan-
ceolate, the uppermost linear-lanceolate, the lower surface pilose to
glabrescent; articles of loment 3–3.5 mm long, ovate-oblong to
suborbiculate, hirtellous-pilose, the dorsal suture convex; isthmi (areas
between articles of loment) central ................................. D. pachyrrhizum
4. Leaflets all elliptic, the lower surface strigose; articles of loment 4–5 mm
long, ovate, uncinulate, pubescent, dorsal suture straight; isthmi
subcentral ........................................................................... D. hickenianum
5(1). Leaves both 3-foliolate and 1-foliolate on the same plant ........................ 6
5. Leaves all 3-foliolate .................................................................................. 7
6(5). Herb or shrublet to 50 cm tall; leaflets 0.8–3 × 0.6–2.1 cm, ovate, oblong or
elliptic, apex rounded to emarginate; stipules linear, acuminate, 4–7 mm
long; frequent on granitic outcrops ....................................... D. orinocense
6. Herb or shrub to 3 m tall; leaflets 3.7–17 × 2–11 cm, elliptic to ovate, apex
mostly obtuse and apiculate; stipules clasping stem, obliquely ovate,
6–15 × 3.5–6 mm; widespread in other habitats .................... D. distortum
7(5). Leaflets < 1 cm long; inflorescence (1)2–4 flowers, fasciculate, axillary or
leaf-opposed .............................................................................. D. triflorum
7. Leaflets mostly > 1 cm long; inflorescence of many more than 4 flowers,
not fasciculate, terminal or axillary ...................................................... 8
8(7). Inflorescence a short, dense raceme, ca. 1–2.5(–3.5) cm long, borne above
the uppermost leaf; calyx densely long silky-pilose and ciliate, the tri-
chomes to 2 mm long .............................................................. D. barbatum
8. Infloresecence a lax raceme, usually longer than 3.5 cm, or a sparingly to
many-branched panicle; calyx variously puberulent or pilose but not as
above ....................................................................................................... 9
9(8). Articles with dorsal suture straight or only slightly concave or notched at
the center ............................................................................................. 10
9. Articles with dorsal suture curved, convex, sinuate, or obtusely to acutely
angled ................................................................................................... 14
10(9). Loments with (1)2 or 3 articles, the articles with dorsal suture slightly
concave or notched at the center ......................................................... 11
10. Loments with 4–many articles, the articles with dorsal suture straight
.............................................................................................................. 12
306 F ABACEAE

11(10). Leaflets chartaceous to subcoriaceous, short- to long-pilose on lower sur-

face; terminal leaflet rhombic-ovate or ovate to elliptic-ovate, obtuse to
rounded at base, the margin not undulate; articles of loment 4–5.5 mm;
loment stipe 3–8 mm long .......................................................... D. axillare
11. Leaflets thin, membranous, sparsely pilose to glabrescent on lower sur-
face; terminal leaflet ovate, acuminate, truncate at base, the margin
undulate; articles of loment 2.5–3 mm wide; loment stipe 1–1.5 mm
long ..................................................................................... D. wydlerianum
12(10). Petioles usually slender, sparsely pilose to glabrescent; leaflets orbicular,
obcordate, or elliptic, secondary and tertiary veins slightly visible but
not prominent on lower surface; loments with 1–4 articles ...................
............................................................................................... D. adscendens
12. Petioles stout, with both uncinulate trichomes and moderately long
spreading trichomes; leaflets generally ovate to ovate-rhombic, second-
ary and tertiary veins easily visible and prominent on lower surface;
loments with 4–many articles ............................................................. 13
13(12). Stipules free from each other; petioles often equaling or longer than the
lateral leaflets, (2.5–)3–7 cm long; leaflets thin; loments with to 5 ar-
ticles, isthmi (area between articles) usually 1/4 or less of the width of
the articles ..................................................................................... D. affine
13. Stipules fused basally or overlapping; petioles usually much shorter than
the lateral leaflets, 1–2.5 cm long; leaflets thick; loments with to 8 ar-
ticles, isthmi usually 1/4–1/2 the width of the articles ............... D. incanum
14(9). Plants prostrate or semiprostrate; articles of loment elliptic or rhombic in
outline .................................................................................................. 15
14. Plants mostly erect, or erect to sprawling, usually becoming shrubby to
3 m or more; articles of loment orbicular or triangular in outline ..... 16
15(14). Articles of loment narrowly elliptic, sutures scarcely constricted at isth-
mi; loments with 7 or 8 articles ............................................. D. scorpiurus
15. Articles of loment mainly rhombic in outline, appearing slightly to much-
twisted, sutures markedly constricted at isthmi; loments with 5 ar-
ticles ..................................................................................... D. procumbens
16(14). Articles of loment mostly orbicular in outline, but sometimes with the
margins alternately revolute; isthmi between articles central .......... 17
16. Articles of loment triangular in outline; isthmi between articles excentric
.............................................................................................................. 18
17(16). Pedicels > 1 cm long, spreading; articles of loment 3–3.5(-4) × 2.5-3 mm;
lower surface of leaflets sparsely pilose mainly on the veins, margins
sparsely ciliate ........................................................................ D. tortuosum
17. Pedicels < 1 cm long; articles of loments 1.5–2.5(–3) × 0.8–1.5(–2) mm;
lower surface of leaflets densely soft-pilose throughout, margins
densely ciliate ......................................................................... D. distortum
18(16). Inflorescences terminal and axillary, densely and many-flowered, race-
mose to racemose-paniculate, and many-branched; pedicels 2.5–6 mm
long, erect at maturity; upper surface of leaflets usually lustrous,
sparsely puberulent ............................................................ D. cajanifolium
18. Inflorescences axillary, racemose, lax and few-flowered; pedicels mostly
12–20 mm long, slender, flexuous; upper surface of leaflets not lus-
trous, appressed-pilose on both surfaces ....................... D. campyloclados

Desmodium adscendens (Sw.) DC., Prodr.
2: 332. 1825. —Hedysarum adscendens
Sw., Prodr. 106. 1788. —Pega-pega.
Repent herb, subshrub, or shrub to 1 m
tall. Weedy areas, widespread in savannas,
forest borders, along rivers, 50–900 m; wide-
spread in Delta Amacuro, Bolívar, and
Amazonas. Widespread elsewhere in Ven-
ezuela; Mexico, Central America, Antilles,
Colombia, Guyana, Suriname, French Gui-
ana, Ecuador, Peru, Brazil, Bolivia, Argen-
tina, Africa, Asia, Melanesia.
Desmodium affine Schltdl., Linnaea 12:
312. 1838.
Procumbent herb; flowers white or pale li-
lac with white. Roadsides, near sea level to
300 m; Delta Amacuro (Caño Mánamo,
Pedernales), Bolívar (Isla Anacoco, Río
Caura). Widespread elsewhere in Venezuela;
Mexico, Central America, Antilles, Colombia,
Ecuador, Peru, Brazil, Bolivia, Paraguay, Ar-
gentina.
Desmodium axillare (Sw.) DC., Prodr. 2:
333. 1825. —Hedysarum axillare Sw.,
Prodr. 107. 1788. —Meibomia axillaris
(Sw.) Kuntze, Revis. Gen. Pl. 1: 195.
1891.
Neotropics; 4 varieties, 3 in Venezuela, 2
of these in the flora area.
Key to the Varieties of D. axillare
1. Stems with long, dense pubescence of
straight trichomes; leaflets ovate to el-
liptic-ovate, the lower surface long-pi-
lose, the apices acute to gradually acu-
minate ...................... var. acutifolium
1. Stems with rather inconspicuous pubes-
cence of short, hooked trichomes; leaflets
ovate, the lower surface short-pilose, the
apices abruptly short-acuminate ............
................................. var. stoloniferum
D. axillare var. acutifolium (Kunze) Urb.,
Symb. Antill. 4: 292. 1905. —Meibomia
axillaris var. acutifolia Kunze, Revis.
Gen. Pl. 1: 195. 1891. —Pega-pega.
Herb with repent or trailing stems; flow-
ers pink, pale-purple, or rose-purple. Shaded
places along forest borders, Trachypogon sa-
vannas, cultivated fields, 50–800 m; Bolívar
(lower Río Caura, near Santa Elena de
Desmodium 307
Uairén), Amazonas (Isla Ratón). Aragua,
Barinas, Carabobo, Lara, Mérida, Miranda,
Portuguesa, Sucre, Trujillo, Yaracuy, Zulia;
Mexico, Central America, Antilles, Colombia,
Guyana, Suriname, French Guiana, Ecua-
dor, Peru, Brazil, Bolivia, Paraguay.
D. axillare var. stoloniferum (Rich. ex
Poir.) B.G. Schub., J. Arnold Arbor. 44:
289. 1963. —Hedysarum stoloniferum
Rich. ex Poir. in Lam., Encycl. 6: 421.
1804 [1805].
Desmodium axillare var. sintenisii Urb.,
Symb. Antill. 2: 303. 1900.
Herb rooting at the nodes; flowers deep
pink. Shaded places along forest borders, 50–
200 m; Bolívar (lower Río Caura). Cojedes,
Monagas; Central America, Antilles,
Guyana, Suriname, French Guiana, Peru,
Brazil, Bolivia.
Desmodium barbatum (L.) Benth. in Miq.,
Pl. Jungh. 224. 1852. —Hedysarum
barbatum L., Syst. Nat. ed. 10, 1170.
1759. —Këmëhkë, Kuhpëkwamën (Pa-
nare).
Herb with ascending to erect stems or
shrub 0.2–1 m tall; flowers purple-pink,
white with some pink stripes, or white to
light purple. Frequent in sandy savannas
and along roadsides, 50–1000 m; widespread
in Delta Amacuro, Bolívar, and Amazonas.
Widespread elsewhere in Venezuela; Mexico,
Central America, Colombia, Guyana, Suri-
name, French Guiana, Ecuador, Peru, Brazil,
Bolivia, Paraguay, Argentina, Old World
Tropics. ŠFig. 273.
Desmodium barbatum is a polymorphic
species with variable habit. The leaves are
sometimes unifoliolate to trifoliolate on the
same plant, the leaflets are sometimes very
small and look like those of D. triflorum, and
the pubescence density on the lower leaf sur-
face is also quite variable.
Desmodium cajanifolium (H.B.K.) DC.,
Prodr. 2: 331. 1825. —Hedysarum caja-
nifoliun H.B.K., Nov. Gen. Sp. (quarto
ed.) 6: 525. 1823 [1824].
Suffrutescent to 3 m tall or erect herb to
1.5 m; flowers rose turning blue-purple. Fre-
quent in Mauritia palm swamps, 50–400 m;
Bolívar (Ciudad Piar, Los Pijiguaos, Río
Chicanán, Río Parguaza, lower Río Sua-
308 F ABACEAE
pure), Amazonas (near Puerto Ayacucho).
Widespread elsewhere in Venezuela; Mexico,
Central America, Antilles, Colombia, Guy-
ana, Suriname, Peru, Brazil, Bolivia.
Desmodium campyloclados Hemsl., Biol.
Cent.-Amer., Bot. 1: 276. 1880. —Mei-
bomia campyloclada (Hemsl.) Kuntze,
Revis. Gen. Pl. 1: 197. 1891.
Desmodium flexuosum Pittier, Bol. Técn.
Minist. Agric. 5: 26. 1944.
Desmodium dubium Pittier, Bol. Soc.
Venez. Ci. Nat. 11: 17. 1947.
Vine, sprawling herb, or slender strag-
gling shrub to 3 m tall or more; flowers
lavender. Tepui slopes, 1500–1800 m; Ama-
zonas (west of Cerro Yutajé). Barinas,
Lara, Mérida, Táchira, Trujillo, Yaracuy;
Central America, Colombia, Ecuador, Pe-
ru, Bolivia.
Desmodium distortum (Aubl.) J.F. Macbr.,
Field Mus. Nat. Hist., Bot. Ser. 8: 101.
1939. —Hedysarum distortum Aubl.,
Hist. Pl. Guiane 774. 1775. —Ihchan
(Panare).
Desmodium asperum (Poir.) Desv., J. Bot.
Agric. 1: 122. 1813. —Hedysarum as-
perum Poir. in Lam., Encycl. 6: 408. 1804
[1805].
Herb to subshrub 1–3 m tall; flowers fra-
grant, purple or rose-lavender. Rocky or
sandy savannas, sandy floodplains, ca. 50–
900 m; Bolívar (Los Pijiguaos, Río Par-
guaza), Amazonas (Puerto Ayacucho). Wide-
spread elsewhere in Venezuela except the
Andes; Mexico, Costa Rica, Antilles, Colom-
bia, Guyana, Suriname, French Guiana, Ec-
uador, Peru, Brazil, Paraguay.
Desmodium hickenianum Burkart, Dar-
winiana 3: 217. 1937.
Herb to suffrutescent to 1 m tall; flowers
purple. Savannas, 100–200 m; Bolívar (near
El Manteco, near Moitaco). Aragua; Bolivia,
Paraguay, Argentina, Uruguay.
Desmodium incanum DC., Prodr. 2: 332.
1825. —Hedysarum incanum Sw., Prodr.
107. 1788.
Hedysarum canum J.F. Gmel., Syst. Nat.
1124. 1792. —Desmodium canum (J.F.
Gmel.) Schinz & Thell., Mém. Soc. Sci.
Nat. Neuchâtel 5: 371. 1914 [1913].
Stems procumbent to ascending or erect-
ascending; flowers rose, bluish gray, or lav-
ender-rose. Weedy areas near towns, along
roadsides, near sea level to 300 m; Delta
Amacuro (Pedernales, Tucupita), Bolívar (El
Dorado, northwest of El Manteco, lower Río
Caura, Upata), Amazonas (Río Puruname).
Widespread elsewhere in Venezuela; U.S.A.,
Mexico, Central America, Antilles, Colombia,
Guyana, Suriname, French Guiana, Ecua-
dor, Peru, Brazil, Bolivia, Paraguay, Argen-
tina, widespread in the paleotropics.
Desmodium orinocense (DC.) Cuello,
Novon 4: 98. 1994. —Desmodium ad-
scendens var. orinocense DC., Prodr. 2:
332. 1825. —Nicolsonia orinocensis
(DC.) Schindl., Repert. Spec. Nov. Regni
Veg. 23: 358. 1927. —Chapere (Gua-
hibo).
Hedysarum adscendens auct. non Sw.
1788: sensu H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 520, t. 597. 1823 [1824].
Shrublet to 50 cm tall, or prostrate shrub
to 1.3 m tall, or spreading herb with
divaricate stems; flowers purplish blue, lav-
ender, or lilac. Herbaceous vegetation around
igneous outcrops, 50–300 m; Amazonas (near
Puerto Ayacucho, Río Atabapo, Río Baría, Río
Casiquiare, Río Cuao, Río Pasimoni). Colom-
bia (Vaupés). ŠFig. 272.
Desmodium pachyrrhizum Vogel, Lin-
naea 12: 97. 1838. —Pega-pega.
Herb, creeping to ascending. Savannas,
50–300 m; Bolívar (Altiplanicie de Nuria,
north of El Manteco, near Represa Guri,
lower Río Caura, Río Paragua). Anzoátegui,
Apure, Aragua, Barinas, Carabobo, Cojedes,
Distrito Federal, Guárico, Miranda, Mona-
gas, Portuguesa, Yaracuy; Colombia, Brazil,
Bolivia, Paraguay, Argentina.
Desmodium procumbens (Mill.) A.
Hitchc., Annual Rep. Missouri Bot.
Gard. 4: 76. 1893. —Hedysarum pro-
cumbens Mill., Gard. Dict. ed. 8, no. 10.
1768.
Pantropics; 3 varieties, all in Venezuela, 1
in the flora area.
D. procumbens var. procumbens
Stems procumbent; flowers white or rose.
Savannas, near sea level to 200 m; Delta
 Desmodium 309

Amacuro (Tucupita), Bolívar (lower Río

Caroní). Anzoátegui, Aragua, Carabobo,
Distrito Federal, Guárico, Lara, Miranda,
Portuguesa, Sucre, Trujillo, Zulia; U.S.A.,
Mexico, Central America, Antilles, Colombia,
Guyana, Suriname, Ecuador, Brazil, Africa.

Desmodium sclerophyllum Benth. in

Mart., Fl. Bras. 15(1): 102. 1859.
Suffrutescent to 1 m tall; flowers red-
purple. Wet savannas dominated by Cura-
tella americana, 100–400(–600) m; Bolívar
(La Paragua, Río Chiguao, Río Paragua, base
of Roraima-tepui), Amazonas (Río Parucito).
Aragua; Colombia, Guyana, Suriname, Peru,
Brazil, Bolivia, Paraguay.

Desmodium scorpiurus (Sw.) Desv., J. Bot.

Agric. 1: 122. 1813. —Hedysarum scor-
piurus Sw., Prodr. 107. 1788. —Amjadi
(Panare), Pega-pega.
Subprostrate herbaceous vine; stems
procumbent; flowers pinkish white or laven-
der. Savannas, near sea level to 300 m; Delta
Amacuro (Tucupita), Bolívar (Ciudad Bolí-
var, La Paragua, around Represa Guri, lower
Río Caroní), Amazonas (Puerto Ayacucho,
Río Ocamo). Widespread elsewhere in Ven-
ezuela; U.S.A., Mexico, Central America, An-
tilles, Colombia, Guyana, French Guiana,
Ecuador, Peru, Brazil (Rondônia), Bolivia,
Africa.

Desmodium tortuosum (Sw.) DC., Prodr. 2:

332. 1825. —Hedysarum tortuosum Sw.,
Prod. 107. 1788. —Arestín, Pega-pega.
Woody herb or shrub to 1 m tall or a vine;
flowers purple. Roadsides, borders of forests,
50–300 m; Delta Amacuro (Los Castillos), Bo-
lívar (Kilómetro 88, Maripa, lower Río Caroní,
Upata), Amazonas (Puerto Ayacucho). Aragua,
Distrito Federal, Falcón, Mérida, Miranda,
Monagas, Portuguesa, Sucre, Táchira, Zulia;
U.S.A., Mexico, Guatemala, Belize, Panama,
Antilles, Colombia, Suriname, Ecuador, Peru,
Brazil, Bolivia, Africa.

Desmodium triflorum (L.) DC., Prodr. 2:

334. 1825. —Hedysarum triflorum L.,
Sp. Pl. 749. 1753.
Low creeping herb; flowers blue. Savan-
nas, 50–200 m; Bolívar (Isla Anacoco, La Ver-
gareña), Amazonas (Puerto Ayacucho). Anzo-
átegui, Apure, Barinas, Carabobo, Cojedes, Fig. 272. Desmodium orinocense
310 F ABACEAE

Miranda, Monagas, Portuguesa, Sucre;

U.S.A., Mexico, Central America, Antilles,
Colombia, Guyana, Suriname, French Gui-
ana, Peru, Brazil, Bolivia, Asia.

Desmodium wydlerianum Urb., Symb.

Antill. 2: 302. 1900.
Slender creeping herb, often a subshrub;
flowers purple. Lower montane forests, in
patches on ground, 300–700 m; Bolívar
(Altiplanicie de Nuria, Río Chiguao), Ama-
zonas (Río Ugueto). Distrito Federal, Sucre;
Guatemala, Costa Rica, Panama, Antilles,
Colombia, Guyana, Suriname, Peru, Brazil.

Fig. 273. Desmodium barbatum

24. DIOCLEA H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 437, fig. 576. 1823 [1824].
by Richard H. Maxwell
Woody vines or lianas, occasionally small, erect subshrubs. Leaves alternate, 3-
foliolate; leaflets ovate, oval, elliptic, to broadly lanceolate; stipules produced below
insertion (i.e., extending above and below point of attachment) or not, rarely absent;
leaflets ovate, oval, elliptic, to broadly lanceolate, entire, unlobed, stipellate. Inflo-
rescence axillary, erect, usually single, pseudoracemose; bracts and bracteoles per-
sistent to caducous. Flowers numerous. Calyx tube 4-lobed, the upper lobe entire or
emarginate; corolla usually shades of purple, occasionally white; standard reflexed
or occasionally straight, mostly emarginate, basally biauriculate, usually bicallose;
wing petals free, occasionally with a spur; keel petals fused distally, beaked or
unbeaked. Stamens 10, pseudomonadelphous; anthers dimorphic or monomorphic.
Pistil geniculate or sigmoidal; ovary usually villous, 1–many-ovulate; style flat or
terete distally; stigma terminal and capitate or somewhat subterminal. Fruit a le-
gume, oblong to obovate, flat or turgid, dry or fleshy, dehiscent or indehiscent,
mostly pubescent. Seeds mostly oval-oblong or cuboid, variable in size, soft or hard,
the hilum linear, nearly 1/2–3/4 encircling the seed or short-oblong.
Pantropics, mostly New World; ca. 55 species, ca. 20 species in Venezuela, 13 of
these in the flora area.
The genus is currently undergoing revision.

Key to the Species of Dioclea

1. Stipules produced below insertion (extending above and below point of

attachment); bracts 6–12 mm long, mostly caducous or semipersistent;
anthers dimorphic (5 + 5) ...................................................................... 2
1. Stipules not produced below attachment; bracts to ca. 3 mm long, mostly
persistent; anthers 10, monomorphic or dimorphic (5 + 5) .................. 3
 Dioclea 311

2(1). Leaflets with secondary veins in ca. 8 pairs, oval or ovate, the apices
rounded or abruptly acute; legume compressed; seeds hard, the hilum
2/3–3/4 encircling; bracts ca. 10 mm long, reflexed, semipersistent or ca-

ducous; flowers buds somewhat oblong, the calyx lobes straight ..........
....................................................................................................... D. reflexa
2. Leaflets with secondary veins in ca. 12 pairs, broadly ovate to somewhat
oval, the apices with abruptly acuminate tips; legume turgid; seeds
soft, the hilum 1/2 encircling; bracts ca. 6 mm long, erect; flower buds
slender, the calyx lobes upcurved ...................................... D. malacocarpa
3(1). Legume 6–16-seeded; seeds hard, the hilum linear, nearly 1/2 encircling;
anthers 10, monomorphic; standard and keel petals mostly puberulent
apically; keel petals oblong or obliquely oblong, the upper margin par-
tially toothed .......................................................................................... 4
3. Legume 1–6(–8)-seeded; seeds soft, the hilum oblong, much less than
1/2 encircling; anthers 10, dimorphic (5 + 5); petals glabrous; keel petals

triangular or somewhat semilunar or rostrate, the upper margin entire

................................................................................................................ 9
4(3). Calyx tube pubescent, sometimes basally or only on the midribs; bracte-
oles 1.5–3 mm long, ovate, acute or lanceolate, mostly persistent, occa-
sionally caducous ................................................................................... 5
4. Calyx tube glabrous; bracteoles 6–16 mm long, broadly ovate or elliptic,
mostly caducous ..................................................................................... 8
5(4). Flowers 1.2–2.5(–3) cm long; standard as wide as or almost as wide as
long; calyx lobes about equal the tube length ....................... D. guianensis
5. Flowers 2.6–3.5 cm long; standard longer than wide; calyx lobes exceed-
ing the tube length ................................................................................. 6
6(5). Legume oblong, 2–2.5 cm wide; calyx tube glabrous or glabrescent, except
along the midribs; bracteoles ovate-orbicular ......................... D. apurensis
6. Legume linear-oblong or oblong, ca. 1.3–2 cm wide; calyx tube pubescent;
bracteoles ovate, acute, or lanceolate .................................................... 7
7(6). Legume linear-oblong, ca. 1.5 cm wide; leaflets oval, elliptic, or broadly
ovate, the apices rounded, both sides canescent-velutinous; calyx
sparsely strigulose, occasionally almost glabrous, except along the mid-
ribs; flowers 3–3.5 cm long, mostly white, occasionally lilac; each wing
petal with a spur ....................................................................... D. albiflora
7. Legume oblong, ca. 2 cm wide; leaflets mostly ovate, oval, to broadly lan-
ceolate, the apices obtuse or acute, the lower surface villous; calyx
sparsely pubescent basally as well as along the midribs; flowers 2.5–3.5
cm long, mostly purple, lilac, rosy white, or occasionally white; wing
petal without a spur ................................................................... D. holtiana
8(4). Flowers > 4 cm long; keel oblanceolate, the upper margin weakly serrate;
legume 10–16-seeded; standard straight ............................. D. macrantha
8. Flowers 2–3 cm long; keels somewhat oblong, the upper margin basally
fimbriate; legume 7–10-seeded; standard reflexed ..................... D. virgata
9(3). Leaflets glabrous; mostly high-climbing forest lianas ............................ 10
9. Leaflets pubescent; lianas, woody vines, or shrublets of open woods and
savannas ............................................................................................... 11
10(9). Anthers 5 + 5; ovary 2-ovulate; stems with conspicuous raised elliptic len-
312 F ABACEAE

ticels; calyx lobes strongly upcurved; legumes obovate-oblanceolate,

2-seeded; seeds compressed; leaflets coriaceous .......................... D. scabra
10. Anthers 10, monomorphic; ovary 4–8-ovulate; stems without raised len-
ticels; calyx lobes mostly straight; legumes mostly oblong, ca. 5-seeded;
seeds compressed or rarely overgrown; leaflets papyraceous to some-
what membranaceous .......................................................... D. macrocarpa
11(9). Legume turgid, oblong, the beak upcurved; seeds cuboid; leaflets subco-
riaceous, with drip tips; mostly forest lianas; flowers to ca. 2 cm long;
anthers monomorphic; inflorescences occasionally branched ... D. ruddiae
11. Legume compressed, elliptic or oblanceolate, the beak downcurved; seeds
compressed, oval-elliptic; leaflets rigidly coriaceous; small vines or
erect shrublets of open woods and savannas; flowers ca. 1.5 cm long or
unknown; anthers monomorphic or unknown; inflorescences single
.............................................................................................................. 12
12(11). Leaflets subconduplicate, the bases cordate; woody vine to 1 m long; flow-
ers unknown; known only from west rim of Cerro Parú, Amazonas
....................................................................................................... D. rigida
12. Leaflets unfolded, the bases mostly round; erect shrublets; flowers ca.
1.5 cm long; plants of open woods and savannas ............... D. steyermarkii

Dioclea albiflora R.S. Cowan, Mem. New
York Bot. Gard. 10(3): 150. 1958.
Vine sprawling over low shrubs. Open for-
ests, river banks, 100–200 m; Bolívar (Río
Orinoco at Piedra Marimare), Amazonas
(Isla Ratón on Río Orinoco, near Puerto
Ayacucho). Endemic.
Dioclea apurensis H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 438. 1823 [1824].
Small to medium-sized vine. Mostly open
areas, ca. 100 m; Bolívar (along Río Orinoco).
Apure; Suriname, Amazon basin.
This species is poorly known. The syntype
has fruit but lacks flowers. Its legume is
larger than those of the Dioclea guianensis
complex. Collections with flowers that have
been assigned to D. apurensis have small or-
bicular or acute bracteoles similar to D.
guianesis s. lat., but frequently the calyx is
pubescent only along the dorsal and ventral
midlines, or the calyx is glabrous.
Dioclea guianensis Benth., Comm. Legum.
Gen. 70. 1837. —Frijolillo.
Dioclea broadwayana Pittier, Bol. Técn.
Minist. Agric. 5: 86. 1944, pro parte,
nom. nud.
Vine. Open areas, extending into forests,
50–800 m; Delta Amacuro (near Los Castillos
de Guayana), widespread in Bolívar and Ama-
zonas. Widespread in the rest of Venezuela;
Mexico, Central America, Colombia, Guyana,
Suriname, French Guiana, Amazonian Brazil.
Pittier’s invalid Dioclea broadwayana
was published without Latin. Pittier contin-
ued to segregate species from his concept of
Dioclea guianensis as well as from that of his
concept of D. broadwayana within the
Guayana area, but he did not validly publish
the names. These taxa are primarily segre-
gates of D. guianensis s. lat., and since their
significance extends beyond the Guayana
area, their elucidation has been deferred
pending further study.
Dioclea holtiana Pittier ex R.H. Maxwell,
Ann. Missouri Bot. Gard. 77: 584. 1990.
—Dioclea holtiana Pittier, Bol. Técn. Mi-
nist. Agric. 5: 84, fig. 36. 1944, nom. nud.
Vine or liana climbing over small trees.
Lowland forests, along rivers and roads, also
associated with granitic outcrops, 50–200 m;
Amazonas (near Puerto Ayacucho, Río Ca-
taniapo, near the mouth of Río Vichada on
the Río Orinoco). Trujillo; Central America,
possibly Mexico, Colombia.
This species is similar in many respects to
Dioclea albiflora and is found in the same lo-
calities in the flora.
Dioclea macrantha Huber, Bol. Mus.
Goeldi Paraense Hist. Nat. Ethnogr. 5:
408. 1909.

Climbing, twining vine over shrubs. Sec-
ondary growth, 100–200 m; Bolívar (near El
Palmar on the Río Grande). Colombia,
Guyana, Suriname, French Guiana, Brazil
(Amapá, Pará).
Dioclea macrocarpa Huber, Bol. Mus.
Goeldi Paraense Hist. Nat. Ethnogr. 5:
410. 1909. —Bejuco de jabón.
High-climbing forest liana. Flooded low-
land riparian forests, 50–500 m; Delta
Amacuro (vicinity of Curiapo, mouth of Río
Amacuro, Río Grande), Bolívar (middle Río
Chiguao, Serranía de Imataca), Amazonas
(Río Mawarinuma). Colombia, Guyana, Suri-
name, Amazonian Ecuador, Peru, Brazil.
Some collections contain legumes with
twisting dehiscence and overgrown cuboid
seeds that do not match the seeds of the type.
Further investigation may warrant the nam-
ing of a variety of Dioclea macrocarpa or D.
ruddiae, an additional variety of D. scabra,
or possibly a new species.
Dioclea malacocarpa Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 170. 1922. —Ojo
de zamuro, Yawade ansajüdü (Yekwana).
Sprawling, coarse vine or forest liana.
Flooded, riparian, and rain forests, 50–500
m; Delta Amacuro (northeast of El Palmar),
Bolívar (Caño Adaua west of Cerro
Guanacoco, southeast of El Dorado, Río
Caura, upper Río Paragua), Amazonas (Río
Atabapo, Río Casiquiare, Río Ventuari, Río
Yaciba in Río Yatúa basin). Trinidad,
Guyana, Suriname, French Guiana, Amazo-
nian Brazil.
Reports of this species from Central
America, Colombia, western Venezuela, Ec-
uador, and Peru should probably be referred
to the closely related Dioclea pulchra
Moldenke.
Dioclea reflexa Hook. f. in Hook., Niger Fl.
306. 1849. —Bejuco des amni, Bejuco de
zamuro, Ojo de cari cari, Ojo de zamuro,
Tangranta-moi (Taurepán).
Coarse climbing or sprawling vine. Low-
land wet forests, near sea level to 400 m;
Delta Amacuro (Río Orinoco basin), northern
Bolívar, Amazonas (upper Río Orinoco).
Pantropics (distribution above the ocean
drift line and upriver due to a flotation adap-
tation in the seed). ŠFig. 274.
Dioclea 313
The name Dioclea hexandra (Ralph)
Mabberly was published (David Mabberley.
Taxon 29: 605. 1980) based on Mucuna
hexandra Ralph (Icon. Carpolog. 30, t. 34, fig.
5. 1849). Mabberley placed the name D.
reflexa in synonymy, but it is retained here,
because it is believed that the Asian D.
hexandra (= D. javanica Benth.) is not the
same species as the western African, north-
ern South American, and Central American
D. reflexa.
Dioclea rigida R.S. Cowan, Mem. New York
Bot. Gard. 10(3): 150. 1958.
Woody vine to 1 m long. Occasional along
upper ridges of summits, ca. 2000 m;
Amazonas (Cerro Parú). Endemic.
The species was described without flowers
from a single collection only, but fits into
Dioclea section Platylobium Benth. Cowan
notes, “the rigidly coriaceous, venose, cor-
date leaflets are distinctive.”
Dioclea ruddiae R.H. Maxwell, Ann. Mis-
souri Bot. Gard. 75: 730. 1988. —Pa-
ronta-yek (Arekuna).
High-climbing forest liana. Roadsides,
savannas, wooded hills, slopes to upland, hu-
mid forests, 800–1600 m; Bolívar (near Cerro
Venamo and La Escalera, Gran Sabana),
Amazonas (Cerro Huachamacari, Sima-
rawochi). Endemic.
Dioclea scabra (Rich.) R.H. Maxwell, Ann.
Missouri Bot. Gard. 77: 578. 1990.
—Dolichos scabra Rich., Actes Soc. Hist.
Nat. Paris 1: 111. 1792.
Usually a high-climbing forest liana,
woody vine, or shrublet. Venezuela, Guyana,
Suriname, French Guiana, Amazonian Bra-
zil; 3 varieties, 2 in Venezuela, both in the
flora area.
Key to the Varieties of D. scabra
1. Flowers 2.3–3 cm long ............ var. scabra
1. Flowers ca. 2 cm long ............ var. brownii
D. scabra var. brownii R.H. Maxwell, Ann.
Missouri Bot. Gard. 77: 579. 1990.
High-climbing liana. Evergreen forests,
ca. 100 m; Amazonas (known from a single
collection from Caño Yagua). Endemic.
314 F ABACEAE

Fig. 274. Dioclea reflexa

 Diplotropis 315

D. scabra var. scabra. —Bejuco de sapo, Neotropics, introduced into Paleotropics; 2

Bejuco de zamuro.
Dioclea elliptica R.H. Maxwell, Ann. Mis-
souri Bot. Gard. 77: 578. 1990, as syn-
onym.
Dioclea glabra auct. non Benth. 1837:
sensu Pittier, Bol. Técn. Minist. Agric. 5:
79. 1944.
Mostly high-climbing liana. Forests, ex-
tending into open areas, 100–400 m; Bolívar
(Río Cuyuní near border with Guyana), Ama-
zonas (widespread). Guyana, Suriname, French
Guiana, Brazil (Amapá, Amazonas, Pará).
Dioclea steyermarkii R.H. Maxwell, Ann.
Missouri Bot. Gard. 77: 585. 1990.
Shrublet or vine to ca. 1 m tall. Savannas
of hills and ridges, 100–600 m; Amazonas
(south and southeast of Cerro Camani). En-
demic.
Dioclea steyermarkii is close to D. coriacea
Benth., but with 10 perfect anthers instead
of 5 perfect and 5 imperfect. It also shows an
affinity to the imperfectly known D. rigida.
Dioclea virgata (Rich.) Amshoff, Meded.
Bot. Mus. Herb. Rijks. Univ. Utrecht 52:
59. 1939. —Dolichos virgatus Rich.,
Actes Soc. Hist. Nat. Paris 1: 111. 1792.
Trailing or climbing roadside vine, or sub-
shrub, sometimes forming dense clumps.
varieties, both in the flora area.
Key to the Varieties of D. virgata
1. Keel petals with part of the upper margin
crenate; bracteoles to ca. 6 mm long,
semipersistent; fruit exocarp with short,
curly, mostly canescent pubescence ........
......................................... var. crenata
1. Keel petals with part of the upper margin
long-fimbriate; bracteoles to ca. 10 mm
long, caducous; fruit exocarp with long,
stiff ferruginous, fugaceous pubescence
.......................................... var. virgata
D. virgata var. crenata R.H. Maxwell, Ann.
Missouri Bot. Gard. 77: 585. 1990.
Woody vine or liana. Margins of humid
forests, 100 m; Amazonas (Río Casiquiare).
Suriname, Brazil (Amapá, Amazonas, Pará).
D. virgata var. virgata. —Ero-cuaja.
Vine. Open areas, river banks, forest
edges, near sea level to 200 m; Delta Ama-
curo (Caño Araguabisi between Caño Ara-
guao and Caño Güiniquina, northeast of
Tucupita), Bolívar (Río Parguaza), scattered
in Amazonas. Throughout the rest of Venezu-
ela; Neotropics, introduced into Paleotropics
through botanical gardens.

25. DIPLOTROPIS Benth., Comm. Legum. Gen. 24. 1837.

by Haroldo Cavalcante de Lima and Gerardo A. Aymard C.
Trees. Leaves alternate, odd-pinnate; stipules small, caducous; stipels none.
Inflorescence a terminal or corymbose panicle or terminal raceme. Flower buds club-
shaped, curved. Calyx 5-dentate, the uppermost 2 joined higher up; petals rose to
violet, plicate-corrugated; standard nearly hastate and auriculate above the claw;
keel petals free. Stamens 10 and dimorphic, or 5 with 5 staminodes; filaments free,
alternately long and short. Ovary subsessile or short-stipitate; ovules 3–6. Fruit in-
dehiscent, samaroid or nut-like, oblong to oblong-orbiculate, membranous to woody.
Seeds 1–few; radicle straight.
Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil, Bo-
livia; 12 species, 7 in Venezuela, all of these in the flora area.

Key to the Species of Diplotropis

1. Leaves 12–20-foliolate, leaflets 1–3 cm wide, oblong to oblong-elliptic; in-
florescences terminal racemes; androecium with 5 stamens and 5 sta-
minodes .................................................................................... D. racemosa
1. Leaves 5–16-foliolate, leaflets 4–9 cm wide, ovate, obovate, elliptic, or
combinations of these; inflorescences terminal panicles; androecium
with 10 stamens ..................................................................................... 2
316 FABACEAE

2(1). Pedicels 4–5 mm long; calyx rusty-villose or appressed-ferruginous-pu-

bescent .................................................................................................... 3
2. Pedicels 0.2–3 mm long; calyx sericeous or tawny-tomentose ................. 5
3(2). Branches and branchlets strigose; leaves 8–11-foliolate; leaflets strigose
on the lower surface; calyx rusty-villose; standard 11–13 mm long
................................................................................................. D. strigulosa
3. Branches and branchlets glabrescent; leaves 5–7-foliolate; leaflets gla-
brous on the lower surface; calyx appressed-ferruginous-pubescent;
standard 7–9 mm long ........................................................................... 4
4(3). Leaflets with dull black on the lower surface; ovary sericeous; fruit woody,
without wings .................................................................................. D. sp. A
4. Leaflets without dull black on the lower surface; ovary ferrugineous; fruit
a samara ....................................................................................... D. triloba
5(2). Leaflets rigid-coriaceous; fruit woody, without wings ................. D. martiusii
5. Leaflets membranaceous to coriaceous; fruit a flat samara, membranous
to papery ................................................................................................ 6
6(5). Secondary and tertiary venation evident on the lower surface; calyx
6–10 mm long; fruits 8–12 × 3–5 cm ....................................... D. purpurea
6. Secondary and tertiary venation strongly reticulate on the lower sur-
face; calyx 4–6 mm long; fruits 5–6 × 1.5–2 cm ................... D. brasiliensis
Diplotropis brasiliensis (Tul.) Benth. in D. purpurea var. purpurea. —Amiña,
Mart., Fl. Bras. 15(1): 320. 1862. —Di- Amuña (Yekwana), Aru-ca-yu-dek (Are-
brachion brasiliensis Tul., Ann. Sci. Nat. kuna), Cachicamo montañero, Congrio,
Bot. sér. 2, 20: 139. 1843. —Bowdichia Congrio blanco, OruKaiyet (Arekuna).
brasiliensis (Tul.) Ducke, Arch. Jard. Bot. Tree 20–40 m tall; flowers pink. Evergreen
Rio de Janeiro 1: 31. 1915. lowland to montane forests, 100–1100 m;
Tree 6–18(–25) m tall; flowers pink; fruit a Delta Amacuro (Río Toro), Bolívar (wide-
samara. Flooded forests, 50–200 m; Ama- spread), Amazonas (Puerto Ayachucho, Si-
zonas (San Carlos de Río Negro). Brazil (Acre, erra Parima, Simarawochi, Río Coro Coro,
Amapá, Amazonas, Pará, Rondônia). upper Río Cunucunuma, Río Jenita on Río
Ocamo, Yutajé). Sucre; Colombia, Guyana,
Diplotropis martiusii Benth., Comm. Le-
Suriname, French Guiana, Ecuador, Peru,
gum. Gen. 24. 1837. —Bowdichia
Bolivia. ŠFig. 275.
martiusii (Benth.) Ducke, Arch. Jard. Bot.
Rio de Janeiro 3: 131. 1922. —Congrio, Diplotropis racemosa (Hoehne) Amshoff,
Iyoynaji (Yanomami). Meded. Bot. Mus. Herb. Rijks Univ.
Dibrachion riparium Spruce ex Benth. in Utrecht 52: 43. 1939. —Bowdichia
Mart., Fl. Bras. 15(1): 321. 1861. racemosa Hoehne, Comiss. Linhas Telegr.
Tree 6–18(–25) m tall; flowers pink; fruit Estratég. Mato Grosso Amazonas, anexo
woody. Flooded forests, 50–200 m; Amazonas 5, Bot. 8: 55. 1919.
(Río Atacavi, Río Baría, Río Casiquiare, Río Venezuela, Guyana, Suriname, Brazil; 2
Pamoni, Río Siapa, Río Sipapo, Río Yatúa). varieties, 1 in Venezuela.
Colombia, Guyana, Suriname, French Gui-
D. racemosa var. racemosa
ana, Peru, Brazil. ŠFig. 276.
Diplotropis racemosa var. kaieteurensis
Diplotropis purpurea (Rich.) Amshoff, Amshoff, Bull. Torrey Bot. Club 75: 393.
Meded. Bot. Mus. Herb. Rijks Univ. 1948.
Utrecht 52: 44. 1939. —Tachigali Tree 6–10(–30) m tall; flowers pinkish
purpurea Rich., Actes Soc. Hist. Nat. white. Borders of evergreen lowland forests,
Paris 1: 108. 1792, "Tachigalia." seasonally flooded forests near black-water
Colombia, Venezuela, Guyana, Suriname, rivers, 100–200 m; Amazonas (Piedra Gua-
French Guiana, Peru, Brazil; 2 varieties, 1 in nare, Raudal Ceguera on Río Autana, Río
the flora area. Casiquiare). Guyana, Suriname, Brazil.
 Diplotropis 317

Fig. 275. Diplotropis purpurea var. purpurea

318 FABACEAE

Fig. 276. Diplotropis martiusii

Diplotropis strigulosa R.S. Cowan, Mem.
New York Bot. Gard. 10(1): 151. 1958.
Tree to 20 m tall; flowers pink. Montane
forests, 1800–2000 m; Amazonas (Cerro Pa-
rú). Endemic.
Diplotropis triloba Gleason, Bull. Torrey
Bot. Club 60: 355. 1933.
Tree to 20 m tall; fruit membranous. Ever-
green lowland forests, 100–200 m; Amazonas
(San Carlos de Río Negro). Brazil (Amazonas,
Mato Grosso, Rondônia).
Diplotropis sp. A
Tree 6–8 m tall; flowers pink. Evergreen
lowland forests, seasonally flooded forests
near black-water rivers 100–200 m; Ama-
zonas (Raudal Ceguera on Río Autana). En-
demic.

26. DIPTERYX Schreb., Gen. Pl. 2: 485. 1791, nom. cons.

Coumarouna Aubl., Hist. Pl. Guiane 740. 1775.
Oleiocarpus Dwyer, Ann. Missouri Bot. Gard. 52: 51. 1965.
by Haroldo Cavalcante de Lima
Trees. Leaves alternate, even-pinnate; leaflets opposite or alternate, often pel-
lucid-punctate; rachis flattened or winged, with apex projected beyond the leaflets;
stipules small, caducous; stipels none. Inflorescence a terminal panicle. Calyx pellu-
cid-punctate, bilabiate, upper lip with 2 large lobes and lower lip very short, entire, or
3-dentate; petals violet to whitish mauve; standard emarginate; keel petals adnate on
lower side. Stamens 10, monadelphous. Ovary stalked; ovule 1. Fruit drupaceous,
 Dipteryx 319

ovoid to oblong-fusiform; mesocarp thick-fibrous; endocarp bony or woody, tardily

opening on ground. Seed cylindric-fusiform; radicle short, straight.
Central America, South America; 9 or 10 species, 4 in Venezuela, all in the flora
area.

Key to the Species of Dipteryx

1. Leaflets 10–18, thin-coriaceous; calyx membranous, glabrous or pubes-

cent; fruit ovoid, with sparse mesocarp and bony endocarp ... D. magnifica
1. Leaflets 2–8, coriaceous; calyx coriaceous, tomentose; fruit ovoid-oblong to
oblong-fusiform, with abundant mesocarp and woody endocarp .......... 2
2(1). Upper lip of calyx 18–25 mm long; leaflets mostly 14–20 × 6–8 cm ...... D. rosea
2. Upper lip of calyx 10–15 mm long; leaflets mostly 7–15 × 3–7 cm ........... 3
3(2). Calyx rusty-tomentose with upper lip 10–15 mm long; petals whitish
mauve .......................................................................................... D. odorata
3. Calyx tawny-tomentose (violet-tomentose when fresh) with upper lip
6–9 mm long; petals violet ........................................................ D. punctata

Dipteryx magnifica (Ducke) Ducke, Trop.
Woods 61: 8. 1940. —Coumarouna mag-
nifica Ducke, Rev. Int. Bot. Appl. Agric.
Trop. 14: 405. 1934. —Maikonaji
(Yanomami), Sarrapia mona.
Taralea casiquiarensis Pittier, Bol. Soc.
Venez. Ci. Nat. 8: 262. 1943.
Tree 20–45 m tall. Forests on white sand
or on granitic outcrops, 100–300 m; Ama-
zonas (Caño Caname, Río Casiquiare, Río
Cunucunuma, Río Padamo, Santa Bárbara
del Orinoco). Brazil.
Dipteryx odorata (Aubl.) Willd., Sp. Pl. 3(2):
910. 1802. —Coumarouna odorata Aubl.,
Hist. Pl. Guiane 740. 1775. —Sarrapia.
Tree 10–30 m tall. Forests on river banks,
100–400 m; Delta Amacuro (Serranía de
Imataca), cultivated in Bolívar and Ama-
zonas. Colombia, Guyana, Suriname, French
Guiana, Peru, Brazil. ŠFig. 277.

Fig. 277. Dipteryx odorata

320 FABACEAE

Dipteryx punctata (S.F. Blake) Amshoff,
Meded. Bot. Mus. Herb. Rijks Univ.
Utrecht 52: 60. 1939. —Coumarouna
punctata S.F. Blake, Contr. U.S. Natl.
Herb. 20: 525. 1924. —Sarrapia, Sar-
rapia real.
Dipteryx trifoliolata Ducke, Trop. Woods
61: 7. 1940.
Tree 8–20 m tall. Evergreen lowland to
lower montane forests, 100–500 m; Bolívar
(Ciudad Bolívar, El Palmar, Río Caura), Ama-
zonas (Río Guainía, San Carlos de Río Negro).
Widely cultivated in Venezuela; Colombia,
Guyana, Suriname, French Guiana, Brazil.
A cultivated form of Dipteryx punctata with
edible fruits and 2 or 3 leaflets was described
as D. trifoliolata Ducke. The fruits are used to
treat stomach pain.
Dipteryx rosea Spruce ex Benth., J. Proc.
Linn. Soc., Bot. 4(suppl.): 125. 1860.
—Sarrapia, Sarrapia mona.
Tree 10–30 m tall. Riparian forests, 50–
300 m; Bolívar (Serranía de Imataca), Ama-
zonas (near San Carlos de Río Negro). Colom-
bia, Peru, Brazil.

27. ERIOSEMA (DC.) Desv., Ann. Sci. Nat. (Paris) 9: 421. 1826. —Rhynchosia sect.
Eriosema DC., Prodr. 2: 388. 1825, spelling variant: Euriosma.
by Renée H. Fortunato
Suffrutescent herbs or subshrubs, erect, ascending to prostrate. Roots woody,
turnip-shaped or spindle-shaped. Stems simple or few-branched, glabrous or pubes-
cent with various pubescence types, glandular or eglandular. Leaves alternate, unifo-
liolate or pinnate-trifoliolate (commonly the 1 or 2 older ones near base of stem uni-
foliolate); stipules free or connate, persistent, rarely deciduous; petioles canaliculate,
short to ± lacking; leaflets membranous, chartaceous, or coriaceous, glabrous to pu-
bescent, glandular on both surfaces or only on lower surface, seldom eglandular,
glandular trichomes usually yellowish or reddish brown; stipels lacking. Inflores-
cences racemose, lax or congested, axillary, solitary, 1–many-flowered; peduncles
surpassing or not surpassing the main cauline leaves; bracts small, persistent or de-
ciduous; bractlets lacking. Calyx 5-lobed, externally pubescent and glandular, the
lobes deltoid, triangular, or subulate, not exceeding corolla, the keel lobe frequently
longer; corolla yellow, sometimes with the standard veins reddish purple; standard
obovate, elliptic or roundish, emarginate, biauriculate, clawed, externally pubescent
and glandular; wing petals narrowly oblong, ± pubescent and glandular at apex; keel
petals falcate, incurved and glandular-pubescent at apex. Stamens 10, diadelphous;
anthers 2-locular, ellipsoid, dorsifixed, uniform. Gynoecium sessile to subsessile;
ovary 2-ovulate, pubescent; style incurved, glabrous; stigmas small, subcapitate. Le-
gumes 2-seeded, obovoid, ellipsoid, compressed, dehiscent, obliquely beaked. Seeds
reniform or ellipsoid, ± oblique in the legumes, lustrous, brown to black, with
strophiole, the hilum long, linear; funicular attachment at extreme apex of the
hilum.
Pantropics; ca. 100 species, 6 in Venezuela, 4 of these in the flora area.

Key to the Species of Eriosema

1. Leaves all unifoliolate ........................................................... E. simplicifolium

1. Leaves trifoliolate (usually 1 or 2 leaves unifoliolate at base of stems) ... 2
2(1). Middle leaflets < 5 times longer than wide, elliptic to obovate; stipules
mostly free, ovate ........................................................................... E. rufum
2. Middle leaflets > 5 times longer than wide, oblong to narrowly elliptic;
stipules connate, rarely free, lanceolate ............................................... 3

3(2). Inflorescence pseudocorymbose, congested, when fully expanded equal to
or shorter than the main cauline leaves; flowers > 9 mm long .............
.................................................................................................... E. crinitum
3. Inflorescence racemose, lax, when fully expanded equaling or exceeding
the main cauline leaves; flowers 4.5–7.5 mm long ................ E. violaceum
Eriosema crinitum (H.B.K.) G. Don, Gen.
Hist. 2: 348. 1932. —Glycine crinita
H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 421.
1823 [1824]. —Rhynchosia crinita
(H.B.K.) DC., Prodr. 2: 389. 1825.
Erect to decumbent subshrub; stems and
leaflets densely reddish-, yellow-, or white-pi-
lose, puberulent, or glabrate; inflorescences
usually in roundish clusters at end of pe-
duncles, 1–8-flowered. Sandy and rocky sa-
vannas, wet forests along rivers, forest edges,
50–1200 m; Delta Amacuro, Bolívar, Ama-
zonas. Mexico, Central America, Greater
Antilles, tropical and subtropical South
America; 4 varieties, 2 in Venezuela, both in
the flora area.
Key to the Varieties of E. crinitum
1. Stems and leaflets densely reddish-pilose
or yellow-pilose, or glabrate; inflores-
cences 2–8-flowered ....... var. crinitum
1. Stems and leaflets white-pilose; inflores-
cences 1–4-flowered ...... var. stipulare
E. crinitum var. crinitum
Ca. 20–40 cm tall. Savannas, edge of for-
ests in open grassland, 50–1200 m; Bolívar
(widespread), Amazonas (near Puerto Aya-
cucho, Sierra Parima). Widespread elsewhere
in Venezuela; Mexico, Guatemala, Belize,
Honduras, Greater Antilles, Colombia, Guy-
ana, Suriname, French Guiana, Peru, Brazil,
Bolivia, Paraguay.
E. crinitum var. stipulare (Benth.) For-
tunato, Novon 3: 25. 1993. —Eriosema
stipulare Benth., Linnaea 22: 519. 1849.
20–100 cm tall. Savannas, 50–300 m;
Delta Amacuro (between Los Castillos de
Guayana and Piacoa), northern Bolívar, cen-
tral and northern Amazonas. Apure, Cojedes,
Guárico, Monagas, Portuguesa, Zulia; Costa
Rica, Colombia, Guyana, Suriname, Brazil.
ŠFig. 279.
Eriosema rufum (H.B.K.) G. Don, Gen. Hist.
2: 347. 1823. —Glycine rufa H.B.K., Nov.
Eriosema 321
Gen. Sp. (quarto ed.) 6: 423, t. 574. 1823
[1824]. —Rhynchosia rufa (H.B.K.) DC.,
Prodr. 2: 384. 1825.
Subshrub 50–100 cm tall. West Indies,
Colombia, Venezuela, Trinidad, Guyana, Su-
riname, Colombia, Ecuador, Peru, Brazil, Bo-
livia, Paraguay; 2 varieties, 1 in Venezuela.
E. rufum var. rufum
Erect stems reddish-pubescent; stipules
commonly free; leaflets short-pilose on both
surfaces. Dry savannas, 50–400 m; Delta
Amacuro (between San Félix and Los Cas-
tillos), common in northern Bolívar. Aragua,
Apure, Cojedes, Guárico, Lara, Monagas,
Portuguesa, Sucre, Trujillo, Yaracuy, Zulia;
Antilles, Colombia, Trinidad, Guyana, Suri-
name, Peru, Brazil, Bolivia. ŠFig. 278.
Eriosema simplicifolium (H.B.K.) G. Don,
Gen. Hist. 2: 348. 1832. —Glycine
simplicifolia H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 419. 1823 [1824]. —Rhyn-
chosia simplicifolia (H.B.K.) DC., Prodr.
2: 389. 1825.
Eriosema simplicifolium var. micranthum
Grear, Mem. New York Bot. Gard. 20(3):
88. 1970.
Decumbent to prostrate, rarely erect;
stems simple or few-branched, 50–80 cm
long; stipules connate. Savannas, edges of
gallery forests, 50–400(–1200); Delta Ama-
curo (between Los Castillos and Piacoa),
Bolívar (scattered), northern and central
Amazonas. Anzoátegui, Apure, Barinas,
Guárico, Monagas, Portuguesa, Zulia; Costa
Rica, Panama, Colombia, Trinidad, Guyana,
Suriname, French Guiana, Brazil, Bolivia,
Paraguay.
Eriosema violaceum (Aubl.) G. Don, Gen.
Hist. 2: 347. 1832. —Cytisus violaceous
Aubl., Hist. Pl. Guiane 766. 1775.
—Rhynchosia violacea (Aubl.) DC., Prodr.
2: 388. 1825.
Stems erect to ascending; stipules connate
or the older ones often free; leaflets ap-
pressed-villous or tomentose on both sur-
322 FABACEAE

Fig. 278. Eriosema rufum var. rufum Fig. 279. Eriosema crinitum var. stipulare

faces. Wet savannas, edges of forests along Monagas; Mexico, Central America, Colom-
rivers, 50–900 m; scattered in Bolívar and bia, Trinidad, Guyana, Suriname, French
Amazonas. Anzoátegui, Apure, Barinas, Guiana, Brazil, Paraguay.

28. ERYTHRINA L., Sp. Pl. 706. 1753.

by David A. Neill
Trees, shrubs, or (outside the flora area) herbaceous perennials with woody
rootstocks, often deciduous and flowering when leafless; bark usually armed with
conical prickles. Leaves pinnately 3-foliolate, with glandular stipels on petiole below
lateral leaflets; lower surface of leaflets glabrous, pubescent, or glaucous with epicu-
ticular wax. Inflorescence racemose, terminal or pseudo-terminal, rarely axillary,
erect or horizontal, flowers in fascicles of (2)3(–6). Flowers showy, red, orange, or
pink. Calyx tubular or campanulate, thick and fleshy or chartaceous, the apex trun-
cate or 1–5-dentate, sometimes cleft or bilabiate; standard larger than wing and keel
petals, sometimes conduplicate and completely enclosing the latter; keel petals co-
herent or free. Stamens usually monadelphous, the vexillar one free for its upper 3/4
 Erythrina 323

and the other 9 free for the upper 1/4. Ovary stipitate, usually tomentose; stigma capi-
tate. Fruit linear, sometimes constricted between the seeds, dehiscent. Seeds red,
bicolored (red and black) or brown, often persistent on pods long after dehiscence.
Tropics, subtropics, and warm-temperate regions worldwide; ca. 115 species, 8
in Venezuela, 4 of these in the flora area.
Erythrina pallida Britton & Rose, known from coastal Venezuela, Trinidad, and
Tobago, is closely related to E. mitis. It may occur in drier forests of the flora area but
flowering material or mature seeds, necessary for positive identification, have not
been collected.

Key to the Species of Erythrina

1. Inflorescence erect; calyx tubular; standard tubular, at least 3 times longer

than wide, conduplicately folded with wing and keel petals and sexual
organs concealed within the standard; seeds red .................................. 2
1. Inflorescence horizontal or arching; calyx campanulate; standard spread-
ing, revealing the keel petals, not conduplicate, < 2 times longer than
wide; seeds brown .................................................................................. 3
2(1). Corollas scarlet, blades of the keel petals free from each other and ob-
tusely rounded at apex .................................................................... E. mitis
2. Corollas dark pink to orange-red, blades of the keel united by their exte-
rior margins or, if free, then acute to acuminate at tip ....... E. rubrinervia
3(1). Evergreen trees, usually growing in swampy or riparian habitats; leaf with
small (ca. 1 mm diameter) glandular, not cupular, stipel; leaflets elliptic
with revolute margins; corollas relatively thick and fleshy, the standard
with a claw about 1/3 the length of the blade, the wings relatively large,
about 1/2 the length of the ovate keel; standard pale orange or salmon-
colored, the wing and keel petals ivory at base, red at apex ......... E. fusca
3. Deciduous trees, usually growing on alluvial terraces but not in swampy
habitats; leaf with large cupular stipel to 4 mm long and 3 mm diameter
on petiole below lateral leaflets; leaflets rhombic to rhombic-ovate; co-
rollas thin-chartaceous, the standard with a barely discernible claw, the
wing petals minute, barely exceeding the calyx; petals uniformly deep
orange .................................................................................... E. poeppigiana

Erythrina fusca Lour., Fl. Cochinch. ed. 1,
427. 1790. —Bucare, Oparu.
Erythrina glauca Willd., Ges. Naturf.
Freunde Berlin Neue Schriften 3: 428.
1801.
Tree to 25 m tall. Usually swampy or ripar-
ian habitats, sometimes forming extensive
monospecific stands, near sea level to 200 m;
Delta Amacuro (road between Caño Guará
and La Horqueta, Caño Rico between Caño
Macareo and Río Grande, Río Orocoima), Bo-
lívar (La Paragua). Anzoátegui, Apure, Ara-
gua, Carabobo, Distrito Federal, Guárico, La-
ra, Mérida, Zulia; Central America, West
Indies, Colombia, coastal Ecuador, eastern
Peru, Brazil, Bolivia, widespread in tropical
southeast Asia, Mascarene Islands, western
Pacific islands, New Guinea. ŠFig. 281.
This is the only species of Erythrina
known from both the Old World and New
World tropics.
Erythrina mitis Jacq., Pl. Hort. Schoenbr. 2:
47. 1797. —Peniolilla, Pionía.
Deciduous tree to 8 m tall, flowering when
leafless; legumes strongly constricted be-
tween the seeds. Semideciduous and lower
montane forests, 50–500 m; Bolívar (Alti-
planicie de Nuria, Caño Maracapra on lower
Río Paragua, Río Botanamo basin), Ama-
324 FABACEAE
zonas (Salto Salas on upper Río Orinoco).
Barinas, Carabobo, Cojedes, Distrito Fed-
eral, Falcón, Lara, Mérida, Miranda, Portu-
guesa, Sucre, Trujillo, Yaracuy, Zulia.
Erythrina poeppigiana (Walp.) Cook,
U.S.D.A. Div. Bot. Bull. 25: 57. 1901.
—Micropteryx poeppigiana Walp., Lin-
naea 23: 740. 1850. —Pericoco.
Tree to 30 m tall. Semideciduous lowland
forests and seasonally flooded riparian for-
ests, 100–400 m; Bolívar (Río San Pedro 200
km south of Caicara), Amazonas (Raudal
Arata on Río Ocamo). Anzoátegui, Barinas,
Carabobo, Distrito Federal, Falcón, Guárico,
Lara, Mérida, Miranda, Portuguesa, Sucre,
Táchira, Trujillo, Yaracuy, Zulia; Panama,
Fig. 280. Erythrina poeppigiana
along base of Andes in Colombia, Ecuador,
eastern Peru, and eastern Bolivia, widely
planted as shade tree for coffee and cacao
(and naturalized in places) in Nicaragua,
Costa Rica, western Panama, and West
Indies. ŠFig. 280.
Erythrina rubrinervia H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 434. 1823 [1824].
Deciduous tree to 25 m tall, flowering when
leafless; branchlets spinose, pith chambered;
legumes strongly constricted between the
seeds. Semideciduous lowland forests, ca.
200 m; Amazonas (Raudal Arata on Río Oca-
mo). Barinas, Lara, Mérida, Táchira, Trujillo;
Belize, Panama, Colombia, Ecuador, Peru,
Bolivia.
Fig. 281. Erythrina fusca
 Etaballia 325

29. ETABALLIA Benth., J. Bot. (Hooker) 2: 99. 1840.

by Nidia L. Cuello A.
Trees. Young stems glabrous, unarmed. Leaves alternate, unifoliolate, the leaf-
let large, entire, pinnately veined, ovate to ovate-oblong, acute to acuminate,
rounded or subcordate at the base; stipules small, deciduous; petiolule very short.
Inflorescences axillary spikes; bracts and bracteoles small, deciduous. Flowers 10–15
mm long, not papilionaceous. Calyx tubular, closed in bud, subregularly 2–5-toothed;
petals 5(6), free, subequal, linear, imbricate. Stamens 10, rarely 11; filaments alter-
nately long and short, connate at the base into a tube free or shortly adnate to the
petals; anthers uniform, short, didymous, opening lengthwise. Ovary subsessile, 1–3
ovulate; style 1 mm long; stigma truncate, oblique. Fruit obovoid, 2-valved but drupe-
like, indehiscent. Seed usually 1; cotyledons fleshy.
Venezuela, Guyana, Brazil; 1 species.

Fig. 282. Etaballia dubia

Etaballia dubia (H.B.K.) Rudd, Phytologia

20: 427. 1970. —Hecastophyllum dubium
H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 388.
1823 [1824]. —Brasil, Sangrito.
Etaballia guianensis Benth., J. Bot.
(Hooker) 2: 99. 1840.
Tree to ca. 30 m tall; inner bark with red
sap; petals linear, yellow to orange. Semide-
ciduous to evergreen lowland forests, 50–200
m; Bolívar (Caicara, Las Bonitas, Puerto
Ordaz to San Félix), Amazonas (Caño Yapa-
cana, Puerto Ayacucho, Río Ventuari). Anzo-
átegui, Apure, Guárico; Guyana, Brazil (Ama-
zonas, Pará). ŠFig. 282.
326 F ABACEAE

30. FISSICALYX Benth., J. Proc. Linn. Soc., Bot. 5: 79. 1861.

by Gerardo A. Aymard C.
Trees. Leaves alternate, odd-pinnate; stipules deciduous; leaflets opposite or
subopposite, without stipels. Inflorescences terminal panicles; bracts and bracteoles
small, the latter persistent. Calyx tube turbinate, 1-lipped, the limb acuminate, en-
tire or shortly 2-dentate, splitting in flower and spathaceous; petals yellow to or-
ange, inserted with the stamens at the top of the calyx tube; standard ovate; wing
petals obliquely oblong, free; keel petals ± similar to the wing petals, free. Stamens
connate into a sheath split above; anthers versatile, opening by 2 apical pores. Ovary
shortly stipitate, 2-ovulate; style filiform; stigma minute, terminal. Fruit flat, ovate-
elliptic including the broad wings, indehiscent. Seed pendent, the hilum small.
Panama, Venezuela, Guyana, Brazil, Paraguay; 1 species.

Fissicalyx fendleri Benth., J. Proc. Linn.

Soc., Bot. 5: 79. 1861.
Monopteryx jahnii Pittier, Bull. Torrey
Bot. Club 42: 626. 1915.
Tree to 20 m tall; petals orange to yellow.
Semideciduous to evergreen lowland and
lower montane forests, 100–500 m; Bolívar
(Cerro Bolívar, La Paragua, lower Río Caro-
ní), Amazonas (Río Manapiare). Aragua, An-
zoátegui, Barinas, Carabobo, Cojedes, Guá-
rico, Lara, Miranda, Portuguesa, Táchira;
Panama, Guyana, Brazil, Paraguay. ŠFig.
283.

Fig. 283. Fissicalyx fendleri

 Galactia 327

31. GALACTIA P. Browne, Civ. Nat. Hist. Jamaica 298. 1756.

by Richard H. Maxwell
Trailing, twining, or climbing, woody or herbaceous perennials, occasionally
small, erect subshrubs. Leaves 3-foliolate or simple; stipules and stipels present;
leaflet blade with entire, unlobed margins. Inflorescence axillary or terminal, erect,
pseudoracemose; bracts and bracteoles present. Calyx 4-lobed; corolla pink, pur-
plish, or white; standard reflexed; wing petals free; keel petals fused distally. Sta-
mens 10, monadelphous or the vexillar one free. Ovary many-ovulate; style fre-
quently exserted during anthesis; stigma minute, capitate. Fruit a legume, oblong to
linear, dehiscent, somewhat compressed. Seeds ovoid, the hilum small, oval.
Pantropics, extending into temperate areas; 50 species, ca. 7 in Venezuela, 3 of
these in the flora area.

Key to the Species of Galactia

1. Leaves digitately 3-foliolate; leaflets linear .............................. G. gracillima

1. Leaves pinnately 3-foliolate; leaflets variable, but never linear .............. 2
2(1). Herbaceous or somewhat woody twining, climbing, or prostrate vines;
leaflets elliptic-ovate ..................................................................... G. striata
2. Woody erect shrubs or subshrubs; leaflets oval, ovate, oblong, or narrowly
elliptic ..................................................................................... G. jussiaeana

Galactia gracillima Benth. in Mart., Fl.
Bras. 15(1): 142. 1859. —Chinak guai-
quin-chinaten (Pemón).
Herbaceous, slender twiner to ca. 50 cm
long. Moist savannas, 1000–1100 m; Bolívar
(Río Kukenán). Suriname, southern Brazil,
Paraguay, Argentina, Uruguay.
Glycine tenuiflora Klein ex Willd., Sp. Pl.
3(2): 1059 1802. —Galactia striata var.
tenuiflora (Klein ex Willd.) Burkart,
Darwiniana 16: 721. 1971. —Galactia
tenuiflora (Klein ex Willd.) Wight & Arn.
1834, pro parte: sensu Benth. in Mart.,
Fl. Bras. 15(1): 143. 1859.

Galactia jussiaeana Kunth, Mimoses 196,

t. 55. 1819 [1824]. —Alcornoquillo, Juan
Zamora.
Galactia angustifolia Kunth, Mimoses
201, t. 56. 1819 [1824]. —Galactia jus-
siaeana var. angustifolia (Kunth) Bur-
kart, Darwiniana 16: 714. 1971.
Galactia camporum Sprague, Trans. &
Proc. Bot. Soc. Edinburgh 22: 430. 1905
[1904].
Shrub; pubescence sericeous to tomen-
tose. Savannas, 50–600 m; Delta Amacuro
(between San Felíx and Los Castillos), north-
ern Bolívar, northern Amazonas. Sucre,
widespread in the Llanos; West Indies, Co-
lombia, Guyana, Suriname, French Guiana,
northern Brazil. ŠFig. 284.

Galactia striata (Jacq.) Urb., Symb. Antill.

2: 320. 1900. —Glycine striata Jacq.,
Hort. Bot. Vindob. 1: 32. 1770, “Glicine.” Fig. 284. Galactia jussiaeana
328 F ABACEAE

Prostrate, trailing, or low twining vine.
Open disturbed areas, 100–500 m; Bolívar
(near Caicara, Moitaco), Amazonas (El Por-
venir 54 km south of Puerto Ayacucho, Santa
Rosa de Ucata). Throughout much of Venezu-
ela; U.S.A. (Florida), Central America, West
Indies, tropical South America, Paraguay,
Argentina.
This species has been introduced world-
wide as a forage crop for tropical pasture im-
provement and has become naturalized, ex-
tending its distribution.

32. GLIRICIDIA H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 393. 1823 [1824], spelling
variants: Glyricidia, Gliciridia.
Hybosema Harms, Repert. Spec. Nov. Regni Veg. 19: 66. 1923.
by Nidia L. Cuello A.
Shrubs or medium-sized trees. Leaves alternate, sometimes partly subopposite
or opposite, odd-pinnate; leaflets 5–19, entire, often drying with purple mottling be-
neath, without stipels; stipules small, caducous. Inflorescences axillary and/or
cauline, of clustered racemes, appearing before or with the leaves; bracts inconspicu-
ous, caducous; bracteoles absent. Flowers often rose or rose-tinged, pedicellate, hy-
panthium distinctively cup-shaped. Calyx teeth short and broad, or absent; petals 5,
of similar length, free except the basally connate keel petals, short-clawed; standard
nearly orbicular, erect or reflexed. Stamens 10, diadelphous, the vexillar stamen
free; anthers uniform. Ovary short-stipitate, slender, straight, flattened, glabrous;
ovules several to many; style glabrous, ca. 1/2 the length of the ovary; stigma capi-
tate. Fruit a legume, dehiscent, stipitate, flattened, often wider toward the apex, the
margins sometimes slightly thickened, nonseptate, exocarp smooth or only faintly
indented between the seeds, glabrous, the valves hard and often coiling in dehis-
cence. Seeds to 10 mm long, nearly round to oblong, compressed.
Neotropics; ca. 6 species, 1 in Venezuela.

Gliricidia sepium (Jacq.) Kunth ex Walp.,

Repert. Bot. Syst. 1: 679. 1842. —Ro-
binia sepium Jacq., Enum. Syst. Pl. 28.
1760. —Mata ratón.
Tree to 6 m tall. Cultivated and perhaps a
locally naturalized escape, 100–300 m;
northeastern Bolívar, Amazonas (Puerto
Ayacucho). Aragua, Carabobo, Cojedes, Dis-
trito Federal, Falcón, Guárico, Lara, Mérida,
Miranda, Portuguesa, Sucre, Yaracuy; native
to Mexico, Central America, and Colombia,
introduced and naturalized in West Indies,
Guyana, Ecuador, Brazil, and Paleotropics.
ŠFig. 285.
Gliricidia sepium is cultivated for forage.
It is also used as a living fence or as a mouse
poison.

Fig. 285. Gliricidia sepium

 Hymenolobium 329

33. HYMENOLOBIUM Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 84. 1860.
by Haroldo Cavalcante de Lima and Gerardo A. Aymard C.
Large or small trees or shrubs. Leaves alternate, crowded at the end of the
branches, deciduous; leaflets 15–55, opposite on the rachis; stipules linear or lan-
ceolate, caducous; stipels minute. Inflorescence a terminal panicle. Calyx campanu-
late, apex truncate, margin obscurely 5-dentate; petals rose or rose-violet; standard
orbiculate and emarginate; keel petals adnate on lower side. Stamens 10, connate
into a sheath open on the upper side. Ovary with many ovules. Fruit indehiscent,
orbicular to oblong-elliptic, samaroid or nut-like, reticulate-veined, with 2 parallel
prominent veins near the margins, membranous or coriaceous; seed chambers cen-
tral. Seeds 1(2), with short straight radicle.
Central America, South America; 12 species, 2 in Venezuela, both in the flora
area.

Key to the Species of Hymenolobium

1. Leaflets pilose to densely pubescent on lower surface, the apex apiculate;

calyx 8–9 mm long; fruit orbicular to oblong, nut-like, coriaceous, 4–7 ×
3–4 cm ............................................................................... H. heterocarpum
1. Leaflets glabrous on both surfaces, the apex emarginate; calyx 4–6 mm
long; fruit oblong-elliptic, samaroid, membranous, 5–12 × 1.5–2.5 cm
.................................................................................................. H. petraeum

Fig. 286. Hymenolobium heterocarpum

330 F ABACEAE

Hymenolobium heterocarpum Ducke, Hymenolobium petraeum Ducke, Arch.

Trop. Woods 47: 6. 1936. —Anzuelito,
Caraota montañera, Mañu, Siñate (Ye-
kwana).
Tree 10–30 m tall; flowers pinkish red.
Riparian forests, 50–200 m; Bolívar (Río
Tabaro near Río Nichare), Amazonas (Ma-
roa to Yavita, Río Cuao, Río Mawarinuma,
Río Sipapo, near San Fernando de Ata-
bapo). Suriname, Ecuador, Brazil. ŠFig.
286.
Jard. Bot. Rio de Janeiro 1: 36. 1915.
—Alcornoque, Alcornoque montañero, Ate-
brino, Pelote, Tanyeechemen (Panare).
Tree 20–40 m tall; flowers pink; fruit red.
Evergreen lowland and semideciduous forests,
granitic slopes, ecotone of savannas, 100–400
m; Bolívar (Río Maniapure, Río Parguaza, Río
Toro), Amazonas (Caicara to Puerto Ayacucho
road, Río Cataniapo). Guárico; Colombia,
Guyana, Suriname, French Guiana, Brazil.

34. INDIGOFERA L., Sp. Pl. 751. 1753.

Anila Mill., Gard. Dict. Abr. ed. 4. 1754.
by Gerardo A. Aymard C.
Perennial herbs or small woody shrubs, strigose with pale, appressed, T-shaped
(malpighiaceous) trichomes. Leaves odd-pinnate, pinnately 3-foliolate, or 1-foliolate;
leaflets entire; stipels present, often inconspicuous; stipules setaceous, deciduous,
apparently adnate to the petioles. Inflorescences axillary racemes or spikes; pedicels
short or apparently absent; bracteoles absent. Flowers usually reddish or white; hy-
panthium campanulate. Calyx 5-dentate, the teeth subequal; standard broad, circu-
lar, subsessile, strigose without; wing petals oblong, somewhat adherent to the keel
petals, auriculate; keel petals coherent, laterally spurred. Stamens 10, diadelphous,
vexillar stamen free, anthers with an apical projection. Ovary slender, sessile, usu-
ally strigose; ovules few to many; style short, glabrous; stigma capitate. Fruits
oblong or linear, terete or 4-angled, curved or straight, dehiscent or apparently
indehiscent, septate. Seeds globose to cylindric and truncate, attached at the
middle.
Pantropics (most diverse in Africa and Asia); ca. 700 species, 11 in Venezuela, 6
of these in the flora area.
Key to the Species of Indigofera
1. Leaves 1-foliolate, linear-lanceolate, 1.3–5 mm wide ............. I. bongardiana
1. Leaves with 3–20 leaflets, rarely 1-foliolate, oblong-elliptic, leaflets 6–15
mm wide ................................................................................................. 2
2(1). Leaflets acute at the base; fruits curved ................................................... 3
2. Leaflets cuneate at the base; fruits straight ............................................. 4
3(2). Fruits 1–1.5 cm long, strongly curved, finely pubescent; leaflets pubescent
on both surfaces .................................................................... I. suffruticosa
3. Fruits 3–3.5 cm long, slightly curved, glabrous or sparsely pubescent;
leaflets usually glabrous on upper surface, pubescent on lower surface
..................................................................................................... I. tinctoria
4(2). Prostrate herb; leaflets 0.2–0.8 cm long; flowers purple; fruits 0.4–1 cm
long ......................................................................................... I. microcarpa
4. Erect or procumbent herb or shrub; leaflets 1.5–5 cm long, eglandular;
flowers red or pink; fruits 1.5–4 cm long .............................................. 5
5(4). Stems, petioles, inflorescence rachis, and fruits appressed-sericeous to
glabrescent ......................................................................... I. lespedezioides
5. Stems, petioles, inflorescence rachis, and fruits densely hirsute with long
brown-red, erect trichomes ........................................................... I. hirsuta
 Indigofera 331

Indigofera bongardiana (Kuntze) Bur-

kart, Darwiniana 4: 171. 1942. —Anila
bongardiana Kuntze, Revis. Gen. Pl. 2:
938. 1891.
Suffrutescent ca. 60 cm tall; flowers pur-
ple. Periodically flooded savannas, 100–200
m; Amazonas (Río Parucito). Southern Bra-
zil, Paraguay, northern Argentina, Uruguay.

Indigofera hirsuta L., Sp. Pl. 1062. 1753.

—Aristín.
Perennial herb or shrub, sometimes with
procumbent branches; flowers pink. Savan-
nas, roadsides, 100–200 m; Bolívar (Maripa).
Widespread elsewhere in Venezuela; Mexico,
Central America, Antilles, Colombia, Guy-
ana, Brazil, Bolivia, Paraguay.

Indigofera lespedezioides H.B.K., Nov.

Gen. Sp. (quarto ed.) 6: 457. 1823 [1824].
—Añilito, Juan Zambrano, Juan Zamo-
ra, Mato zabandu, Raíz de zamuro,
Romero de sabana.
Indigofera pascuorum Benth., Ann. Nat.
Hist. 3: 431. 1839.
Perennial herb or shrub; flowers pink.
Wet savannas, roadsides, 50–400 m; north-
ern Bolívar, Amazonas (near Puerto Aya-
cucho). Widespread elsewhere in Venezuela;
Mexico, Central America, Antilles, Colombia,
Guyana, Brazil, Bolivia, Paraguay. ŠFig. 287.
This species is locally cultivated for its
roots, which are used to treat diarrhea.

Indigofera microcarpa Desv., J. Bot. (Des-

vaux) 3: 79. 1814. —Pega pega rosada.
Prostrate herb; flowers purple. Open sa-
vannas, 50–200 m; Bolívar (northeast of
Cerro Cuchivero, El Tigre on Río Cuchivero).
Guárico, Monagas, Zulia; Peru, Brazil, Bo-
livia, Paraguay, Uruguay.

Indigofera suffruticosa Mill., Gard. Dict.

ed. 8, no. 2. 1768. —Añilito, Inchan-
pichanemen (Panare).
Suffrutescent or shrub to 3 m tall; flowers
pink. Open and disturbed areas, near sea level
to 600 m; Delta Amacuro (Isla Iduburojo near
Misión San Francisco de Guayo, Pedernales)
Bolívar (Ciudad Bolívar, Icabarú, Kilómetro
88, Río Caura, Río Suapure, Upata). Wide-
spread elsewhere in Venezuela; U.S.A., Mex-
ico, Central America, Antilles, Colombia, Guy-
ana, Suriname, French Guiana, Ecuador,
Peru, Brazil, Bolivia, Paraguay, Argentina. Fig. 287. Indigofera lespedezioides
332 F ABACEAE

Indigofera suffruticosa is used as a fish 50 m; Bolívar (Ciudad Bolívar). Monagas;

poison in the lower Río Caura basin.
Indigofera tinctoria L., Sp. Pl. 751. 1753.
Erect or sprawling shrub to 1.5 m tall;
flowers pink. Open and disturbed areas, ca.
U.S.A. (Florida), Mexico, Central America,
Antilles, native of the Old World (India) in-
troduced and naturalized in Neotropics as
the source of blue indigo dye.

35. LECOINTEA Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 128. 1922.
by Basil Stergios
Trees. Leaves 1-foliolate, variously serrate to crenate, glabrous; stipules cadu-
cous. Flowers not papilionaceous, rather small, pedicellate, borne on axillary, often
branched racemes; bracts persistent; bracteoles small, persistent. Calyx gamo-
sepalous, tube campanulate to cotyliform (disk-shaped but with raised or ascending
border) with 5 obscure lobes; petals 5, separate, spatulate, clawed, imbricate, soon
deciduous, outer one much wider than inner 4. Stamens 9–13, free, ± unequal; an-
thers basifixed, longitudinally dehiscent, pilosulous at tip. Ovary stipitate; ovules 4–
6; style straight or slightly bent, stout and exerted; stigma small, oblique. Fruit
thick, globose to oval or broadly turbinate, indehiscent. Seeds 1 or 2, thick and
semicompressed, ovoid to orbicular.
Central America, Venezuela, Suriname, Ecuador, Peru, Brazil, Bolivia; 6 spe-
cies, 2 in Venezuela, 1 of these in the flora area.

Fig. 288. Lecointea amazonica

 Lonchocarpus 333

Lecointea amazonica Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 129, pl. 8. 1922.
—Níspero, Palo de arco, Pilón.
Tree 15–25 m tall; flowers white. Ever-
green lowland and flooded forests, 50–600 m;
Delta Amacuro (upper Río Toro), Bolívar
(Represa Guri, Río Nichare), Amazonas
(southern Río Manapiare basin). Guatemala,
Belize, Costa Rica, Suriname, Ecuador, Peru,
Brazil. ŠFig. 288.

36. LONCHOCARPUS H.B.K., Nov. Gen. Sp. (quarto ed.) 6: 383. 1823 [1824], nom.
cons.
by Hans-Helmut Poppendieck
Trees, shrubs, or vines. Leaves alternate, odd-pinnate; leaflets opposite, entire;
stipules caducous; stipels absent (present in L. densiflorus). Inflorescence a lateral
or terminal pseudoraceme, the flowers either in pairs on distinct or short, lateral
peduncles, or numerous on lateral short shoots, or arising directly from the rachis
and appearing verticillate, paired, or solitary; peduncular and pedicellar bracts ca-
ducous; bracteoles 2, borne on the pedicel or at the base of the calyx. Calyx dentate
to truncate, cup-shaped; corolla purple or white; standard blade broadly ovate to
obovate, clawed, the base either narrow and without callosities, or truncate to cor-
date with callosities; wing petals adhering to the keel petals by pressure and struc-
tural conformity; keel petals touching or connate. Vexillar stamen free at base; an-
thers uniform; nectary glands indistinct. Ovary pubescent; ovules 2–12; style gla-
brous. Fruit generally indehiscent, occasionally dehiscent along the vexillar suture,
or separating into 1-seeded loments, papery to thin or thick woody, 1–12-seeded,
vexillar margin occasionally winged, but wings never submarginal.
Mexico, Central America, West Indies, South America south to Peru and north-
ern Argentina, 1 species in western Africa; ca. 150 species, 21 in Venezuela, 16 of
these in the flora area.
The circumscription adopted here for Derris and Lonchocarpus is conventional,
but far from satisfactory. It is maintained following earlier treatments by Benth. (in
Carl Friedrich Philipp von Martius, Flora Brasiliensis 15(1): 275–287. 1862), G.
Amshoff (in A. Pulle, Flora of Suriname 2(2): 141–147. 1939), and H. Pittier (Bol.
Tec. Minist. Agric. Cría, Caracas 5: 96–106. 1944), and because it is consistent with
some recent chemical and other surveys and avoids nomenclatural changes with far-
reaching consequences. There is a close connection between Lonchocarpus subgenus
Phacelanthus Pittier and the controversial American species of Derris. The winged
fruit, which is the diagnostic character differentiating the two genera, has certainly
evolved more than once in a parallel manner.
Lonchocarpus utilis is a rotenone-yielding, cultivated species that is often
found in cultivation or escaped. It has never been collected in flower or fruit and
therefore is not included in the key. It has oblong-elliptic leaflets with a gradually
acuminate apex and a round to acute base, whereas the other known rotenone-con-
taining species, L. urucu, has obovate-oblong leaflets with an abruptly acuminate
apex and a roundish to obtuse base.

Key to the Species of Lonchocarpus (except L. utilis; see above)

1. Flowers numerous in short lateral shoots; generally lianas, rarely trees or

shrubs (subgenus Phacelanthus) ........................................................... 2
1. Flowers paired on a common peduncle, or directly arising from the spike;
trees or shrubs, rarely vines (subgenus Lonchocarpus) ....................... 6
334 F ABACEAE

2(1). Deciduous trees; legumes flattened, lanceolate to linear, fragile and

sublomentaceous, 1–8-seeded ........................................................ L. pictus
2. Evergreen plants, initially shrubby but then becoming large lianas; le-
gumes either flattened and 1- or 2-seeded, or strongly thickened, 1–12-
seeded ..................................................................................................... 3
3(2). Stipels present; ovules 10–12; legumes 7–15 × 4–5 cm, to 12-seeded,
lomentaceous .......................................................................... L. densiflorus
3. Stipels absent; ovules 2–6; legumes flattened and different from above
................................................................................................................ 4
4(3). Calyx and corolla without latex ducts; standard with ± distinct auriculae
or basal callosities, base truncate, claw glabrous ................ L. floribundus
4. Calyx and corolla with latex ducts; standard without auriculae or basal
callosities, base narrowed, claw pilose .................................................. 5
5(4). Lower surface of leaflets distinctly spreading-pilose, trichomes ca. 0.2 mm
long; margins of legume only slightly thickened ...................... L. martynii
5. Lower surface of leaflets appressed-pilose, appearing subglabrous, tri-
chomes 0.25–0.5 mm long; margins of legume distinctly thickened
........................................................................................................ L. urucu
6(1). Flowers arising directly from the spike and appearing verticillate, paired,
or solitary; peduncle of fruit not jointed ............................................... 7
6. Flowers paired on a common peduncle; peduncle of fruit jointed ............ 9
7(6). Flowering spikes 14–18 cm long ..................................................... L. fendleri
7. Flowering spikes ≤ 11 cm long ................................................................... 8
8(7). Leaflets obtuse ..................................................................... L. crucisrubierae
8. Leaflets acuminate ........................................................................ L. tubicalyx
9(6). Calyx denticulate ..................................................................................... 10
9. Calyx entire, or at most slightly undulate .............................................. 12
10(9). Spikes dense, flowers touching each other, often at least partly dark violet
to blackish when dry; leaflets 11–20 × 6.8–8.5 cm, without pellucid
dots; legumes to 7.5 × 2.3 cm .............................................. L. heptaphyllus
10. Spikes lax, flowers not touching each other and light-colored when dry;
leaflets 3.5–11 × 1.5–6.5 cm, with or without pellucid dots; legumes at
least 7.5 × 2.8 cm ................................................................................. 11
11(10). Leaflets rugose, the margin involute, the main veins of lower surface dis-
tinctly pilose; legumes 3.5–4 cm wide, to 10 mm thick, prominently and
abruptly thickened at the seeds, the seeds nearly spherical .................
........................................................................................... L. crassispermus
11. Leaflets flat, the margin flat, tomentose on both surfaces; legumes 2.8–3.8
cm wide, to 2 mm thick, not abruptly thickened at the seeds, the seeds
not spherical ........................................................................... L. hedyosmus
12(9). Leaves pellucid-punctate, glabrous; spikes 11–19 cm long; peduncles
7–10 mm long; pedicels 5–7 mm long; legumes 7–17 × 3–3.2 cm, glabrous
.................................................................................................. L. punctatus
12. Leaves not pellucid-punctate, glabrous to pubescent; spikes 9.5–15 cm
long; peduncles to 3 mm; pedicels to 3 mm long; legumes 4–11 × 1.3–2.5
cm, pubescent or glabrous ................................................................... 13
13(12). Vexillar margin of legume conspicuously winged ................ L. dipteroneurus
13. Vexillar margin of legume without wings ............................................... 14

14(13). Legumes (1–)3–7-seeded, 5.5–11 × 1.3–1.8 cm, bullate, dark brown, gla-
brous; leaves glabrous .......................................................... L. imatacensis
14. Legumes 1- or 2-seeded, 4–6.5 × 2–2.5 cm, woody, dark gray, pubescent;
leaves pubescent on lower surface, scabrid to pubescent on upper sur-
face .............................................................................................. L. sericeus
Lonchocarpus crassispermus Poppend.,
Novon 2: 53. 1992. —Jebe, Mahomo
chino.
Tree to 20 m tall. Semideciduous forests,
savanna edges, 200–300 m; Bolívar (15 km
southeast of El Callao, between La Encru-
cijada and El Pao, 7 km north of mouth of Río
Paragua, 17 km from Upata toward San
Félix). Endemic.
Lonchocarpus crucisrubierae Pittier,
Arb. Arbust. Venez. 6–8 [reprinted from
Bol. Minist. RR. EE. no. 8/9]: 100. 1927.
—Majomo rebalsero, Menudito, Tocorito.
Tree or shrub 2–8 m tall; trunk to 40 cm
diameter, mostly branched from the base, oc-
casionally with elongate vining branches;
flowers precocious. Semideciduous and sea-
sonally flooded forests, 50–200 m; Bolívar
(Ciudad Bolívar, Puerto Ordaz, lowlands of
Río Orinoco basin and tributaries). Barinas,
Guárico.
Lonchocarpus densiflorus Benth., Ann.
Nat. Hist. 3: 433. 1839.
Lonchocarpus glabrescens Benth., Hooker’s
J. Bot. Kew Gard. Misc. 2: 233. 1850.
—Derris glabrescens (Benth.) J.F.
Macbr., Field Mus. Nat. Hist., Bot. Ser.
13(3.1): 261. 1943.
Lonchocarpus boliviensis Pittier, Contr.
U.S. Natl. Herb. 20: 93, fig. 43. 1917.
Small shrub, flowering when 3 m tall, be-
coming a high-climbing liana; young
branches reddish brown, lenticellate. Ripar-
ian forests, 50–200 m; Delta Amacuro (Caño
Carisal), Bolívar (Río Orinoco), Amazonas
(Río Negro). Apure; Colombia, Peru, north-
ern Amazonian Brazil, Bolivia.
The 1-seeded segments of the peculiar
lomentaceous fruits are dispersed by water.
Lonchocarpus dipteroneurus Pittier,
Contr. U.S. Natl. Herb. 20: 90. 1917.
Lonchocarpus stenopteris Pittier, Arb.
Arbust. Venez. 6–8 [reprinted from Bol.
Minist. RR. EE. no. 8/9]: 102. 1927
Lonchocarpus 335
Small tree 3–10 m tall, often many-
stemmed or -branched from the base; twigs
gray to dark brown, lenticellate. Deciduous
forests, 100–300 m; Bolívar (Represa Guri,
Villa Lola). Aragua, Distrito Federal, Guá-
rico, Miranda.
The collections from Bolívar state are ei-
ther widely disjunct from the populations in
northern Venezuela, or else the species has
not been collected in intermediate areas.
Lonchocarpus fendleri Benth., J. Proc.
Linn. Soc., Bot. 4(suppl.): 94. 1860.
Lonchocarpus fendleri subsp. pubescens
Pittier, Contr. U.S. Natl. Herb. 20: 91. 1917.
Lonchocarpus sanctae-marthae Pittier,
Contr. U.S. Natl. Herb. 20: 92, fig. 42, pl.
1. 1917.
Lonchocarpus stenurus Pittier, Arb.
Arbust. Venez. 6–8 [reprinted from Bol.
Minist. RR. EE. no. 8/9]: 102. 1927.
Deciduous tree 6–25 m tall, often low-
branching, with dense, rounded to almost
flat crown, bark light gray; flowers appear-
ing before the leaves. Gallery forests, near
sea level to 100 m; Delta Amacuro (Caño
Arature). Frequent in northern Venezuela;
Panama, Colombia.
Lonchocarpus floribundus Benth., Ann.
Nat. Hist. 3: 432. 1839. —Derris
floribundus (Benth.) Ducke, Bol. Técn.
Inst. Agron. N. 18: 197. 1949.
Lonchocarpus nitidulus Benth., J. Proc.
Linn. Soc., Bot. 4(suppl.): 98. 1860.
Shrublet, flowering when 1 m tall, becom-
ing a large liana in closed forests. Locally fre-
quent in clearings, mostly on sandy soil,
100–200 m; Bolívar (Serranía Baraguán, Río
Parguaza), Amazonas (Río Negro, Río Ori-
noco and tributaries). Guyana, Suriname,
French Guiana, Amazonian Brazil.
Lonchocarpus floribundus is used as a
fish poison in Brazil.
Lonchocarpus hedyosmus Miq., Linnaea
18: 564. 1844. —Derris hedyosma (Miq.)
336 F ABACEAE
J.F. Macbr., Field Mus. Nat. Hist., Bot.
Ser. 13(3.1): 262. 1943. —Jebe, Majomo.
Lonchocarpus margaritensis Pittier, Contr.
U.S. Natl. Herb. 20: 91, fig. 41. 1917.
Lonchocarpus ernestii Harms, Repert.
Spec. Nov. Regni Veg. 17: 321. 1921,
“ernesti.” —Derris ernestii (Harms)
Ducke, Bol. Técn. Inst. Agron. N. 18:
196. 1949, “ernesti.”
Lonchocarpus paniculatus Ducke, Arch.
Jard. Bot. Rio de Janeiro 3: 161. 1922.
Tree 8–30 m tall; trunk to 1 m diameter,
often branched from the base; bark gray.
Semideciduous and evergreen lowland for-
est, forested slopes along savannas, 50–500
m; widespread in northern Bolívar, Ama-
zonas (San Fernando de Atabapo). Venezu-
elan Coastal Cordillera and Andes; Guyana,
Suriname, Brazil.
The hard and durable wood of Loncho-
carpus hedyosmus is used for construction.
Lonchocarpus heptaphyllus (Poir.) DC.,
Prodr. 2: 259. 1825. —Dalbergia hepta-
phylla Poir. in Lam., Encycl. suppl. 2:
445. 1811 [1812]. —Tocorito.
Dalbergia pentaphylla Poir. in Lam.,
Encycl. suppl. 2: 445. 1811 [1812].
—Lonchocarpus pentaphyllus (Poir.)
DC., Prodr. 2: 259. 1825.
Lonchocarpus latifolius H.B.K. ex DC.,
Prodr. 2: 260. 1825. —Derris latifolia
(H.B.K. ex DC.) Ducke, Bol. Técn. Inst.
Agron. N. 18: 195. 1949.
Lonchocarpus discolor Huber, Bol. Mus.
Paraense Hist. Nat. Ethnogr. 3: 421.
1902.
Tree 7–26 m tall; trunk to 38 cm or more
diameter, occasionally vining; hollow
branches ant-inhabited; bark reddish brown
to grayish. Evergreen lowland forests, 100–
300 m; Delta Amacuro (Serranía de Imataca),
Bolívar (Serranía de Imataca). Apure, Bari-
nas, Carabobo, Distrito Federal, Miranda,
Monagas, Sucre; Central America, West
Indies, Colombia, Peru, Brazil (Amapá, Pará).
Lonchocarpus imatacensis Poppend.,
Novon 2: 53. 1992. —Jebe, Majomo.
Tree to 26 m tall; trunk to 40 cm diameter.
Evergreen lowland forests, 50–400 m; Delta
Amacuro (Serranía de Imataca), Bolívar
(Altiplanicie de Nuria, Salto Pará, Serranía
de Imataca). Endemic. ŠFig. 291.
This species has been misidentified as
Lonchocarpus nitidus Benth., but can be dis-
tinguished by the bullate seed region of the
fruit. The wood is used for railroad ties.
Lonchocarpus martynii A.C. Sm., Amer. J.
Bot. 26: 577. 1937.
Scandent shrub or liana eventually reach-
ing large dimensions; young branches dark
brown, pubescent. Evergreen lowland to
lower montane forests, gallery forests, forest
borders, 200–900 m; Bolívar (Río Nichare ba-
sin, Santa Elena de Uairén). Guyana, adja-
cent Amazonian Brazil.
Lonchocarpus pictus Pittier, Arb. Arbust.
Venez. 2/3 [reprinted from Bol. Comerc.
Industr. Venez. no. 34]: 26. 1923. —Ma-
jomo, Tocorito, Tocorito blanco.
Slender or branched deciduous tree or
shrub 8–15 m tall; flowers present as or
rarely before the leaves expand. Semide-
ciduous to evergreen lowland forests, gallery
forests, 50–200 m; Delta Amacuro (near Río
El Toro), Bolívar (Ciudad Bolívar, Río
Suapure to La Paragua). Venezuelan Coastal
Cordillera; Guyana, Suriname, French Gui-
ana. ŠFig. 290.
Like Hibiscus tiliaceus L., the bark of
Lonchocarpus pictus is used locally against
erysipelas, a febrile disease producing deep
red color of the skin.
Lonchocarpus punctatus H.B.K., Nov.
Gen. Sp. (quarto ed.) 6: 383. 1823 [1824].
—Jebe.
Robinia violacea Jacq., Enum. Syst. Pl.
28. 1760, nom. dubium. —Lonchocarpus
violaceus (Jacq.) DC., Prodr. 2: 259.
1825.
Lonchocarpus benthamianus Pittier,
Contr. U.S. Natl. Herb. 20: 86. 1917.
Spreading, branched tree or shrub 1–15
m tall; bark grayish. Deciduous to semide-
cidous forests, 50–300 m; Bolívar (Río
Caroní, mouth of Río Paragua, Río Upata,
Río Yuruari). Venezuelan Coastal Cordil-
lera; West Indies, Colombia, Ecuador,
Peru, Brazil.
Lonchocarpus sericeus (Poir.) H.B.K. ex
DC., Prodr. 2: 260. 1825. —Robinia
sericea Poir. in Lam., Encycl. 6: 226.
1804. —Majomo.
 Lonchocarpus 337

Fig. 289. Lonchocarpus sericeus

Fig. 290. Lonchocarpus pictus

338 F ABACEAE
Fig. 291. Lonchocarpus imatacensis
Tree 12–25 m tall, the wood hard and du-
rable; leaf rachis, branchlets, and inflores-
cences ferruginous-pubescent; bark brown-
ish or grayish, lenticellate, yielding resinous
fluid when cut. Semideciduous to evergreen
lowland or lower montane forests, 50–400 m;
Delta Amacuro (Río Acure), Bolívar (near El
Dorado, near La Paragua). Mexico, Central
America, West Indies, Colombia, Ecuador,
Peru, Brazil, western Africa. ŠFig. 289.
Lonchocarpus tubicalyx Pittier ex Pop-
pend., Novon 2: 57. 1992.
Shrub or tree 4–5 m. Habitat not known,
near sea level to 100 m; Delta Amacuro
(Caño Mánamo south of Tucupita). Apure.
Lonchocarpus tubicalyx is currently
known from just two specimens with very
short pedicels, making the flowers appear
verticillate.
Lonchocarpus urucu Killip & A.C. Sm., J.
Wash. Acad. Sci. 20: 81, fig. 4, 1930.
—Derris urucu (Killip & A.C. Sm.) J.F.
Macbr., Field Mus. Nat. Hist., Bot. Ser.
13(3.1): 266. 1943.
Erect shrub becoming a huge woody vine,
often found in large clumps of several acres.
Well-drained, fertile soils in nonflooded for-
ests, ca. 100 m; Amazonas (La Esmeralda).
Amazonian Brazil.
Lonchocarpus urucu is used in Brazil as a
fish poison and a source of commercially ex-
ploited rotenone. It is very difficult to differ-
entiate from L. utilis, which is used in the
same manner. The leaf shape affords the
only, and often difficult, way to distinguish
the species.
Lonchocarpus utilis A.C. Sm., Amer. J.
Bot. 24: 580. 1937. —Derris utilis (A.C.
Sm.) Ducke, Bol. Técn. Inst. Agron. N.
18: 197. 1949. —Barbasco, Barbasco
blanco, Barbasco blanco caicareño, Bar-
basco bravo, Timbo.
Lonchocarpus nicou auct. non (Aubl.) DC.
1825: sensu Killip & A.C. Sm., J. Wash.
Acad. Sci. 20: 74. 1930. —Derris nicou
(Aubl.) J.F. Macbr., Field Mus. Nat.
Hist., Bot. Ser. 13(3.1): 263. 1943, non
Robinia nicou Aubl., Hist. Pl. Guiane
771. 1775.
Erect shrub becoming a woody vine. Sec-
ondary forests at sites of old cultivated ar-
eas, near sea level to 300(–1300) m; scat-
tered throughout Delta Amacuro, Bolívar,
and Amazonas. Elsewhere in Venezuela in
the adjacent Río Orinoco basin; widely culti-
vated throughout the Amazon basin, Guy-
ana, Suriname, French Guiana, and eastern
Brazil.
The rotenone-yielding roots of Loncho-
carpus utilis are used as a fish poison. The
flowers and fruits are unknown; it is propa-
gated vegetatively and spreads in cultivation
by means of its roots.
 Machaerium 339

37. MACHAERIUM Pers., Syn. Pl. 2: 276. 1807, nom. cons.

Drepanocarpus G. Mey., Prim. Fl. Esseq. 236. 1818.
by Velva E. Rudd
Lianas, trees, or shrubs, scandent with spine-like prehensile shoots; sap usu-
ally reddish. Leaves alternate, odd-pinnate, 3–many-foliolate; leaflets alternate, en-
tire; stipules usually indurated, spinescent or sometimes caducous; stipels lacking.
Inflorescence terminal or axillary, racemose, sometimes paniculate; bracts small,
caducous or stipule-like, spiny but smaller; bracteoles small, usually broadly ovate,
obtuse, paired at base of calyx. Flowers papilionaceous, ca. 4–18 mm long. Calyx
campanulate or briefly tubiform, 5-lobed to subtruncate; petals white to yellow,
pink, bluish, or purple. Stamens 10, monadelphous or diadelphous (5:5 or 9:1); an-
thers dorsifixed. Ovary 1- or 2-ovulate; style filiform; stigma minute, capitate, termi-
nal. Fruit indehiscent, stipitate, usually samaroid with a basal seminiferous body
and a sterile, terminal wing, or sometimes lunate, falcate, or subreniform with the
wing reduced or lacking. Seeds compressed, ovate, reniform, or orbicular.
Tropical and subtropical America, 1 species in coastal West Africa; ca. 130 spe-
cies, ca. 35 in Venezuela, 22 of these in the flora area.
Key to the Species of Machaerium
1. Leaflets with the secondary veins approximately parallel, usually well
defined, extending to the margin or nearly so ...................................... 2
1. Leaflets with the secondary veins branching, not always parallel, usually
not extending to the margin, sometimes weakly defined ..................... 9
2(1). Secondary veins ca. 3–25 mm apart; leaves 5–15-foliolate; leaflets 1.5–
11 cm wide; fruit winged ........................................................................ 3
2. Secondary veins ca. 1 mm apart; leaves 5–35-foliolate; leaflets 0.5–3 cm
wide; fruit winged or unwinged ............................................................ 5
3(2). Lower surface of leaflets with trichomes lax or curled; venation conspicu-
ous; fruit ferruginous-tomentulose, glabrescent with age; flowers 8–
17 mm long .............................................................................. M. quinatum
3. Lower surface of leaflets subglabrous or with trichomes appressed and
usually minute; secondary veins usually evident but lesser veins incon-
spicuous; fruit puberulent to tomentulose, usually subglabrous by ma-
turity; flowers 5–10 mm long ................................................................ 4
4(3). Leaflets predominantly ovate to elliptic, acute to acuminate, sometimes
obtuse; flowers 6–10 mm long on pedicels ca. 1 mm long; calyx 3–4 × 2–
2.5 mm; fruit with stipe 3–5(–10) mm long ...................... M. floribundum
4. Leaflets predominantly obovate to elliptic, obtuse; flowers 5–8 mm long,
essentially sessile; calyx 2.5–3 × 1.5–2 mm; fruit with stipe 2–3 mm
long .................................................................................. M. macrophyllum
5(2). Fruit with a well-developed terminal wing ............................................... 6
5. Fruit lunate or falcate, lacking a terminal wing ....................................... 7
6(5). Leaflets ca. 1–2 × 0.2–0.6 cm, linear to oblong or elliptic-oblong, the sur-
faces puberulent with lax trichomes, somewhat glabrescent with age;
flowers (7–)8–10 mm long, standard puberulent on the outer face; fruit
essentially straight ........................................................................ M. affine
340 F ABACEAE

6. Leaflets 1–5(–9) × (0.5–)1–2(–3.5) cm, elliptic to oblong, the upper surface

glabrous or nearly so, the lower surface puberulent with lax trichomes
or finely appressed-pubescent, glabrescent with age; flowers (8–)10–15
mm long, the standard glabrous or subglabrous on the outer face; fruit
usually bent at about a 90° angle ..................................... M. robiniifolium
7(5). Leaflets acute to obtuse or emarginate, aristate at the apex, the arista
(0.5–)1–5 mm long, rarely lacking, the secondary veins diverging from
the midvein at angles of about 90°; flowers 10–14 mm long; calyx 5–7 ×
4 mm .................................................................................... M. aristulatum
7. Leaflets obtuse or retuse, not apiculate, the secondary veins diverging
from the midvein at angles of about 45°; flowers (7–)8–13 mm long; ca-
lyx 3.5–5(–7) × 2–2.5 mm ....................................................................... 8
8(7). Flowers ca. 10–13 mm long; calyx ferruginous-pubescent; fruit falcate or
lunate, not curved into a circle, puberulent, glabrescent with age,
sessile, or stipe to ca. 5 mm long; leaves ca. 15–35-foliolate; lower sur-
face of leaflets puberulent with lax trichomes ............................... M. ferox
8. Flowers ca. 10 mm long; calyx glabrous to moderately pubescent, not fer-
ruginous; fruit usually curved into a circle, glabrous or subglabrous at
maturity, the stipe 5–10 mm long; leaves 5–13-foliolate; lower surface
of leaflets glabrous to moderately pubescent with minute appressed tri-
chomes ....................................................................................... M. lunatum
9(1). Leaflets ca. (0.2–)0.3–3(–4) × 0.1–1.1 cm, predominantly elliptic to linear
or oblong; fruit winged ......................................................................... 10
9. Leaflets ca. 2–20 × 1.5–8.5 cm, predominantly ovate to ovate-elliptic or
oblong; fruit with or without a terminal wing .................................... 12
10(9). Flowers (8–)10–12 mm long; calyx 3.5–5 × 2.5–3 mm; petals white to
purple, the standard pubescent on the outer face; leaflets elliptic to ob-
long, 5–30 × 3–11 mm ........................................................ M. altiscandens
10. Flowers 4–6(–7) mm long; calyx 1–2.5 × 1–1.5 mm; petals white to cream-
colored or yellowish, sometimes with a few purplish or reddish mark-
ings, the standard glabrous on the outer face; leaflets linear or linear-
oblong, 5–30(–40) × 1–10 mm .............................................................. 11
11(10). Leaves 37–90(–100)-foliolate; leaflets linear or linear-oblong, 5–13 × 1–
2 mm .............................................................................. M. multifoliolatum
11. Leaves ca. 35–51-foliolate; leaflets linear-oblong, 10–30(–40) × (2–)3–
5(–10) mm .......................................................................... M. myrianthum
12(9). Fruit lunate or subreniform, lacking a terminal wing ........................... 13
12. Fruit with a well-developed terminal wing ............................................. 15
13(12). Leaflets coriaceous, usually blackening on drying, sometimes pubescent
on lower surface along the midvein otherwise glabrous; fruit ferrugi-
nous- or fulvous-velutinous; flowers white to pinkish ........ M. inundatum
13. Leaflets subcoriaceous, usually not much darkening on drying, minutely
appressed-pubescent to subglabrous on lower surface; fruit fulvous-
tomentulose or puberulent, glabrescent; flowers dark red to purple .... 14
14(13). Fruit ca. 7–9 cm long, the stipe 5–7 mm long; pubescence generally whit-
ish or gray; trees ........................................................................ M. dubium
14. Fruit ca. 3.5 cm long, the stipe 1–3 mm long; pubescence generally brown-
ish or tawny; small trees, shrubs, or lianas ....................... M. leiophyllum
 Machaerium 341

15(12). Flowers 13–18 mm long; calyx 6–10 × 4–5 mm; fruit ferruginous-
tomentulose, sometimes setose, usually glabrous at maturity, the stipe
5–10 mm long ............................................................................... M. kegelii
15. Flowers 5–11 mm long; calyx 2–4 × 1–2.5 mm; fruit sericeous to puberu-
lent or glabrous, the stipe 2–15 mm long ............................................ 16
16(15). Lianas; flowers 8–11 mm long; calyx 3.5–4 × 2–3 mm ............................ 17
16. Lianas or trees; flowers ca. 5–8 mm long; calyx 2–3.5 × 1–2 mm .......... 18
17(16). Flowers 9–11 mm long; petals white but drying black, rusty-brown-
villous; leaves 11–17-foliolate; fruit sometimes puberulent at the base,
otherwise glabrous, the stipe 10–12 mm long ................... M. amazonense
17. Flowers 8–10 mm long; petals bluish; leaves 11- or 13-foliolate; fruit fulvous-
to feruginous-velutinous, the stipe 5–15 mm long ................. M. tovarense
18(16). Fruit fulvous-sericeous, glabrescent, not darkening on drying, the stipe 2–
5 mm long; flowers 5–7 mm long; calyx 2.5–3 × 2 mm, subsericeous
scarcely striate; petals purplish or white, not darkening on drying, the
standard sericeous on the outer face; upper surface of leaflets glabrous,
the lower surface sericeous or subsericeous to glabrous; usually lianas
.............................................................................................. M. madeirense
18. Fruit glabrous to puberulent, usually darkening on drying, the stipe 4–
15 mm long; flowers 5–8 mm long; calyx 2–3.5 × 1–2 mm; petals white
or greenish, usually darkening on drying; leaflets glabrous on both sur-
faces or puberulent to glabrous on upper surface, appressed-pubescent
to subsericeous on lower surface; trees or shrubs .............................. 19
19(18). Calyx ca. 2 × 2 mm, scarcely striate; fruit essentially glabrous at maturity,
the stipe (6–)10–15 mm long; leaves 11–17-foliolate; upper surface of
leaflets puberulent to glabrous, the lower surface appressed-pubescent
to subsericeous ..................................................................... M. acutifolium
19. Calyx ca. 2–3.5 × 1–2 mm, with prominent longitudinal striations; fruit
puberulent to glabrous at maturity, the stipe 4–10 mm long; leaves 3–
9-foliolate; leaflets glabrous on both surfaces ..................................... 20
20(19). Flowers ca. 5–6.5 mm long; calyx 2–2.5 × 1–1.5 mm; leaflets acuminate,
the acumen ca. 1.5–2 cm long; fruit with stipe 7–10 mm long ...............
............................................................................................ M. acuminatum
20. Flowers ca. 8 mm long; calyx 3–3.5 × 2 mm; leaflets acuminate, the acu-
men to ca. 1 cm long; fruit with stipe 5.5–7 mm long..... M. guaremalense

Machaerium acuminatum H.B.K., Nov. Machaerium acutifolium Vogel, Linnaea

Gen. Sp. (quarto ed.) 6: 391. 1823 [1824].
—Nissolia acuminata (H.B.K.) DC.,
Prodr. 2: 258. 1825. —Cumariche.
Tree to ca. 20 m tall. Deciduous or semiev-
ergreen forests, 100–1100 m; Amazonas
(near Puerto Ayacucho). Aragua, Guárico,
Lara, Miranda, Trujillo, Yaracuy. ŠFig. 293.
The only collection from the flora area
(Morillo 6741, VEN) is somewhat disjunct
from the rest of the range of Machaerium
acuminatum. It has slightly smaller fruit
and is tentatively assigned here.
11: 187. 1837.
Tree to 10 m tall. Venezuela, Brazil, Bo-
livia; 3 varieties, 1 in Venezuela, 1 in the
flora area.
M. acutifolium var. pseudacutifolium
(Pittier) Rudd, Phytologia 25: 399. 1973.
—Machaerium pseudacutifolium Pittier,
Bol. Soc. Venez. Ci. Nat. 7: 148. 1941.
—Tucurito.
Semideciduous forests, savannas, 50–300
m; Bolívar (Represa Guri, Río Asa, Río
342 F ABACEAE
Paragua). Falcón, Guárico; northern Brazil,
Bolivia.
Machaerium affine Benth., Comm. Legum.
Gen. 34. 1837. —Bainepá, Vainepá.
Machaerium rectipes Pittier, Bol. Soc.
Venez. Ci. Nat. 9: 121. 1944.
Tree to ca. 17 m tall. Seasonally flooded
savannas and gallery forests, 50–300 m;
Bolívar (near La Paragua). Guyana.
Machaerium altiscandens Ducke, Arch.
Jard. Bot. Rio de Janeiro 4: 75. 1925.
Liana or tree to ca. 25 m tall. Evergreen
lowland forests, 50–200 m; Bolívar (Río
Parguaza). Suriname, Amazonian Brazil.
The leaflets of this species show consider-
able variation in width.
Machaerium amazonense Hoehne, Arq.
Bot. Estado São Paulo, n.s. 1: 46, t. 56.
1939.
High-climbing liana. Evergreen lowland
to montane primary and secondary forests
and savannas, 100–900 m; Bolívar (near
Santa Elena de Uairén), Amazonas (slope of
Río Cataniapo). Brazil (Amazonas).
Machaerium aristulatum (Spruce ex
Benth.) Ducke, Arch. Jard. Bot. Rio de
Janeiro 4: 311. 1925. —Drepanocarpus
aristulatus Spruce ex Benth., J. Proc.
Linn. Soc., Bot. 4(suppl.): 69. 1860.
—Robasesina.
Tree, shrub, or liana, to ca. 10 m tall.
Moist savannas, river banks, 50–200 m,
Bolívar (near Ciudad Bolívar, San Rafael),
Amazonas (Isla Ratón, Río Orinoco). Apure,
Barinas; Colombia, Peru, northern Brazil,
Bolivia.
Machaerium dubium (H.B.K.) Rudd,
Phytologia 24: 122. 1972. —Drepa-
nocarpus dubius H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 390. 1823 [1824].
—Almendro.
Machaerium caicarense Pittier, Bol. Soc.
Venez. Ci. Nat. 9: 120. 1944.
Tree to ca. 15 m tall. Seasonally flooded
evergreen gallery forests, 50–100 m; Bolívar
(Río Orinoco at Caicara). Apure, Guárico.
Machaerium ferox (Mart. ex Benth.) Ducke,
Arch. Jard. Bot. Rio de Janeiro 4: 311.
1925. —Drepanocarpus ferox Mart. ex
Benth., Comm. Legum. Gen. 32. 1837.
—Bejuco de murciélago, Uña de gavilán.
Drepanocarpus ferox ß macrophyllum
Benth. in Mart., Fl. Bras. 15(1): 256.
1862.
Liana, shrub, or small tree, to ca. 20 m
tall. Flooded or nonflooded riparian forests,
near sea level to 200 m; Delta Amacuro
(Caño Guara near Isla Guara), Amazonas
(above La Esmeralda, Río Orinoco near
mouth of Río Atabapo, Río Pamoni). Colom-
bia (Amazonas), Guyana, Suriname, Brazil.
ŠFig. 294.
Machaerium floribundum Benth., J. Proc.
Linn. Soc., Bot. 4(suppl.): 68. 1860.
—Chaperno, Sangre de toro, Uña de
gavilán.
Drepanocarpus venezuelensis Pittier,
Contr. U.S. Natl. Herb. 20: 122, fig. 59.
1918. —Machaerium venezuelense
(Pittier) Hoehne, Fl. Brasilica 25(3): 53.
1941.
Machaerium decorticans Ducke, Arch.
Jard. Bot. Rio de Janeiro 3: 150. 1922.
Machaerium woodworthii Standl., Contr.
Arnold Arbor. 5: 81. 1933.
Machaerium rosescens Standl., Publ.
Carnegie Inst. Wash. 461: 24. 1935.
Drepanocarpus ?ovalifolium Pittier, Bol.
Soc. Venez. Ci. Nat. 7: 149. 1941.
—Machaerium longistipitatum Pittier,
Bol. Técn. Minist. Agric. 5: 119. 1944,
based on Drepanocarpus ovalifolium
Pittier, non Machaerium longistipitatum
Hoehne 1939.
Southern Mexico, Central America, Co-
lombia, Venezuela, Guyana, French Guiana,
Ecuador, Peru, northern Brazil, Bolivia; 3
varieties, all in Venezuela, 1 in the flora
area.
Variability in size and shape of leaflets
seems to be the chief reason for the several
synonyms.
M. floribundum var. floribundum
Tree, shrub, or liana to ca. 25 m tall. Moist
forests, near sea level to 200 m; Delta
Amacuro (Caño Araguao, Caño Daudacana),

Amazonas (between Cerro Yapacana and
Santa Bárbara, upper Río Atacavi, Río
Orinoco). Northern Venezuela; southern
Mexico, Central America, Colombia, Guy-
ana, French Guiana, Peru, northern Brazil.
ŠFig. 295.
Machaerium guaremalense Pittier, Arb.
Legum., part 3, Trab. Mus. Comercial
Venezuela 4 [reprinted from Bol. Minist.
R.R. E.E. no. 4–7]: 213. 1928.
Small tree or liana to ca. 5 m tall.
Semideciduous to evergreen lowland forests,
savannas, 50–300 m; Bolívar (island in Lago
Guri, Río Orinoco east of mouth of Río
Horeda and Cerro Gavilán). Anzoátegui,
Carabobo, Portuguesa.
This species is similar to and possibly
synonymous with Machaerium darienensis
Pittier, a species from Panama and is closely
related to M. striatum J. Johnst. from Colom-
bia and coastal Venezuela.
Machaerium inundatum (Mart. ex Benth.)
Ducke, Arch. Jard. Bot. Rio de Janeiro 4:
311. 1925. —Drepanocarpus inundatus
Mart. ex Benth., Comm. Legum. Gen.
32. 1837. —Jarizo, Urapo.
Liana or tree to 20 m tall. Flooded forests
along rivers, near sea level to 200 m; Delta
Amacuro (Río Acure, Serranía de Imataca),
northern Bolívar, Amazonas (widespread).
Apure, Monagas, Guárico; Colombia, Guy-
ana, Suriname, French Guiana, Peru, north-
ern Brazil. ŠFig. 296.
Machaerium kegelii Meisn., Linnaea 21:
257. 1848.
Machaerium bracteatum Benth., J. Proc.
Linn. Soc., Bot. 4(suppl.): 65. 1860.
Machaerium pachyphyllum Pittier, Contr.
U.S. Natl. Herb. 20: 469. 1922.
Machaerium steinbachianum, Hoehne,
Arq. Bot. Estado São Paulo, n.s. 1: 48.
1939.
Liana, shrub, or tree to ca. 20 m tall. Pri-
mary, semi-evergreen, and semideciduous
forests, 200–1300 m; Delta Amacuro
(southeast of Los Castillos de Guayana,
southeast of Piacoa), Bolívar (Caño Pablo
near Salto Pará, road between Tumeremo
and Bochinche), Amazonas (Río Mana-
Machaerium 343
piare). Anzoátegui, Barinas; Mexico, Cen-
tral America, Guyana, Suriname, French
Guiana, Ecuador, Peru, Brazil, Bolivia.
ŠFig. 298.
Machaerium leiophyllum (DC.) Benth.,
Comm. Legum. Gen. 36. 1837.
—Nissolia leiophylla DC., Prodr. 2: 258.
1825.
Colombia, Venezuela, Guyana, French
Guiana, Peru, Amazonian Brazil, Bolivia; 4
varieties, 2 in Venezuela, 1 of these in the
flora area.
M. leiophyllum var. latifolium (Benth.)
Rudd, Phytologia 22: 56. 1971. —Drepa-
nocarpus crista-castrensis var. latifolium
Benth. in Mart., Fl. Bras. 15(1): 258.
1862.
Drepanocarpus frondosus Mart. ex
Benth., Comm. Legum. Gen. 32. 1827.
—Machaerium frondosum (Mart. ex
Benth.) Ducke, Arch. Jard. Bot. Rio de
Janeiro 3: 151. 1922.
Liana, sometimes a tree or shrub. River
banks, flooded forests, 50–100 m; Bolívar
(Río Horeda, Río Parguaza), Amazonas
(mouth of Río Samariapo, mouth of Río
Sipapo). Apure; Guyana, French Guiana,
Amazonian Brazil. ŠFig. 297.
Without fruit this variety is virtually in-
distinguishable from var. leiophyllum, which
has much larger, less lunate fruit than var.
latifolium.
Machaerium lunatum (L. f.) Ducke, Arch.
Jard. Bot. Rio de Janeiro 4: 310. 1925.
—Pterocarpus lunatus L. f., Suppl. 317.
1781. —Drepanocarpus lunatus (L. f.) G.
Mey., Prim. Fl. Esseq. 238. 1818.
—Arepillo, Olvidanovia, Siete conchas.
Shrub or small tree to ca. 8 m tall, some-
times scandent. Brackish coastal marshes,
sandy soil, mangrove swamps, near sea level
to 50 m; Delta Amacuro (widespread).
Monagas, Sucre; Nicaragua, Costa Rica,
Panama, Hispaniola, Puerto Rico, Lesser
Antilles, coastal Colombia, Trinidad, Guy-
ana, Suriname, French Guiana, Brazil, west-
ern coast of Africa from Senegal to Angola.
ŠFig. 301.
344 F ABACEAE
Machaerium macrophyllum Benth.,
Comm. Legum. Gen. 35. 1837.
Colombia, Venezuela, Guyana, Suriname,
Peru, Brazil; 2 varieties, 1 in Venezuela.
M. macrophyllum var. macrophyllum
Liana, sometimes a tree, to ca. 18 m tall.
Evergreen forests, 100–800 m; Bolívar (Caño
Iguapo, Río Erebato, mouth of Río Nichare),
Amazonas (Cucurital near middle Caño
Yagua, Culebra, Mavaca, San Carlos de Río
Negro, Sierra de la Neblina, Simarawochi,
Yavita). Colombia (Amazonas), Guyana,
Suriname, Peru, Brazil (Amazonas).
Machaerium madeirense Pittier, Contr.
U.S. Natl. Herb. 20: 119. 1918.
Machaerium compressicaule Ducke, Arch.
Jard. Bot. Rio de Janeiro 4: 76. 1925.
Machaerium compressicaule var. mano-
ense Ducke, Arq. Inst. Biol. Veg. 2: 45.
1935. —Machaerium latifolium var.
manoense (Ducke) Hoehne, Fl. Brasilica
25(3): 65. 1941.
Liana, sometimes a tree to ca. 10 m tall.
Gallery forests, evergreen lowland forests,
100–500 m; Bolívar (west side of Amaruay-
tepui, southwest of El Dorado, Río Acanán,
Río Asa), Amazonas (below mouth of Caño
Yapacana, Culebra, El Porvenir, Río Orinoco
between Tamatama and Esmeralda). South-
eastern Colombia, Guyana, Suriname, Peru,
Amazonian Brazil, northern Bolivia. ŠFig.
302.
Variation in leaflet shape seems to be
chiefly responsible for the synonymy.
Machaerium multifoliolatum Ducke,
Bull. Mus. Hist. Nat. (Paris) sér. 2, 4:
734. 1932.
Liana to ca. 30 m long. Evergreen lowland
and lower montane forests, 100–1000 m;
Amazonas (slopes of Cerro Duida, San
Carlos de Río Negro). Colombia (Vaupés),
Brazil (Amazonas), Bolivia. ŠFig. 299.
Machaerium myrianthum Spruce ex.
Benth., J. Proc. Linn. Soc., Bot.
4(suppl.): 59. 1860.
Liana or scandent tree to ca. 10 m tall.
Lowland to lower montane forests, 200–800
m; Bolívar (Altiplanicie de Nuria, Cerro
Jaua, northwest of El Manteco, Santa Elena
de Uairén, Sierra de Lema). Colombia
(Vaupés), Guyana, Suriname, northern Bra-
zil. ŠFig. 300.
Machaerium quinatum (Aubl.) Sandwith,
Bull. Misc. Inform. Kew 1931: 359. 1931.
—Nissolia quinata Aubl., Hist. Pl.
Guiane 743, t. 4. 1775.
Nissolia ferruginea Willd., Sp. Pl. 3(2):
900. 1802. —Machaerium ferrugineum
(Willd.) Pers., Syn. Pl. 2: 276. 1807.
Tree, shrub, or liana to ca. 16 m tall. Low-
land to upland riparian forests, savannas,
usually on sandy soil. Colombia, Venezuela,
Guyana, Suriname, French Guiana, Brazil; 2
varieties, both in the flora area.
Key to the Varieties of M. quinatum
1. Flowers ca. 8–12 mm long; calyx 2.5–3 × 2–
3 mm; bracteoles 1–2 × 1.5–2.5 mm
................................. var. parviflorum
1. Flowers ca. 12–17 mm long; calyx 4–6 ×
3–4 mm; bracteoles 2–3 × 3–4 mm
...................................... var. quinatum
M. quinatum var. parviflorum (Benth.)
Rudd, Phytologia 24: 121. 1972.
—Machaerium ferrugineum ß parvi-
florum Benth. in Mart., Fl. Bras. 15(1):
253. 1862. —Acoi-yerí-yek (Arekuna),
Chaparillo, Maripa-yen-ya-pupen-yek
(Arekuna), Uña de murciélago.
Machaerium nervosum Vogel, Linnaea 11:
186. 1837.
100–1200 m; Bolívar (Arabopó, Roraima-
tepui, base of Sororopán-tepui), Amazonas
(San Carlos de Río Negro, Yavita). South-
eastern Colombia, French Guiana, Brazil.
ŠFig. 304.
M. quinatum var. quinatum
100–900 m; Bolívar (Río Cuyuní, Río
Parguaza, near Santa Elena de Uairén),
Amazonas (Simarawochi). Guyana, Suri-
name, French Guiana, Brazil (Amazonas).
ŠFig. 303.
Machaerium robiniifolium (DC.) Vogel,
Linnaea 11: 190. March 1837. —Nissola
robiniifolia DC., Prodr. 2: 258. 1825.
Machaerium sieberi Benth., Comm. Le-
gum. Gen. 34. June 1837.

Machaerium eggersii Hoehne, Arq. Bot.
Estado São Paulo, n.s. 1: 47, t. 58. 1939.
Tree to ca. 20 m tall, with stipular spines.
Semideciduous to evergreen lowland forests,
50–800 m; northern Bolívar. Widespread in
northern Venezuela; Colombia, Trinidad,
Guyana, Brazil (Rio Branco area). ŠFig. 292.
There has been confusion with Machae-
rium moritzianum Benth. and many speci-
mens so determined are actually M. robini-
ifolium.
Machaerium tovarense Pittier, Contr.
U.S. Natl. Herb. 20: 121. 1918.
Liana. Semideciduous forests on hills,
200–300 m; Bolívar (island in Lago Guri 40
Fig. 292. Machaerium robiniifolium
Machaerium 345
km south of Represa Guri). Aragua, Distrito
Federal, Falcón, Miranda, Zulia.
Machaerium sp. A
Tree ca. 25 m tall. Montane riparian for-
ests, 700–900 m; Amazonas (Sierra Parima
along Río Matacuni).
The single collection, Steyermark 107084
(US), previously annotated as Machaerium
acuminatum, bears flower buds but it cannot
be reliably identified. There appears to be
some relationship to M. acuminatum, M.
amazonense, and M. guaremalense. However,
it does not compare well with any of those spe-
cies and until better material is available it
is better to maintain it as Machaerium sp. A.
346 F ABACEAE

Fig. 293. Machaerium acuminatum

Fig. 294. Machaerium ferox

 Machaerium 347

Fig. 295. Machaerium floribundum var. floribundum

348 F ABACEAE

Fig. 296. Machaerium inundatum

Fig. 297. Machaerium leiophyllum var. latifolium

 Machaerium 349

Fig. 298. Machaerium kegelii

Fig. 299. Machaerium multifoliolatum

350 F ABACEAE

Fig. 300. Machaerium myrianthum

Fig. 301. Machaerium lunatum

 Machaerium 351

Fig. 302. Machaerium madeirense

352 F ABACEAE

Fig. 303. Machaerium

quinatum var.
quinatum

Fig. 304. Machaerium quinatum var. parviflorum

 Macroptilium 353

38. MACROPTILIUM (Benth.) Urb., Symb. Antill. 9: 457. 1928. —Phaseolus sect.
Macroptilium Benth. in Mart., Fl. Bras. 15(1): 189. 1859.
by Gerardo A. Aymard C.
Vines or sometimes erect or sprawling herbs, sometimes perennial from a thick
rootstock. Leaves pinnate, trifoliolate or rarely unifoliolate; stipules veined, not pro-
longed below the insertion; stipels ciliate to tomentose; leaflets glabrous or pubes-
cent, lacking hooked trichomes. Inflorescences with stiff, elongate peduncles, the
flowers mostly congested at the apex; bracteoles narrow, at least distally, veined, ca-
ducous; rachis with small swellings at the nodes, lacking extrafloral nectaries;
pedicels equaling or shorter than the calyx. Flowers purplish, violet, or white, the
interior often vivid. Calyx campanulate, the teeth free, the upper 2 teeth sometimes
reduced; standard orbicular, emarginate, with 2 small basal auricles, lacking medial
thickenings; wing petals longer than the standard and the keel petals, long-stipitate;
keel petals apically spiraled, basally adnate to the staminal tube. Style apically re-
curved and thickened, caducous. Fruit linear, turgid or compressed, nonseptate.
Seeds numerous, small, with a short hilum.
Pantropics, cultivated in Africa and Asia as forage; ca. 20 species, 6 species in
Venezuela, 4 of these in the flora area.

Key to the Species of Macroptilium

1. Leaves unifoliolate ............................................................... M. monophyllum

1. Leaves trifoliolate ....................................................................................... 2
2(1). Leaflets linear, 2–7 mm wide ........................................................... M. gracile
2. Leaflets ovate, elliptical, rhombic, oblong, or linear, 1–4 cm wide .......... 3
3(2). Stipels glabrous, ca. 3 mm long; stipules lanceolate, striate; calyx with lan-
ceolate teeth; fruit 7–12 cm long .......................................... M. lathyroides
3. Stipels long-ciliate, ca. 1 mm long; stipules lanceolate-subulate, veined;
calyx with deltoid teeth; fruit 3–6 cm long ............. M. longepedunculatum

Macroptilium gracile (Poepp. ex Benth.) Neotropics, cultivated as forage in Asia; 2

Urb, Symb. Antill. 9: 457. 1928. —Pha-
seolus gracilis Poepp. ex Benth., Comm.
Legum. Gen. 77. 1837.
Perennial herb or vine; flowers red, purple,
or pink. Savannas, edges of towns, 50–400 m;
Delta Amacuro (between Los Castillos and
Piacoa), Bolívar (Altiplanicie de Nuria,
Caicara, Ciudad Bolívar, La Paragua, La Ur-
bana, lower Río Caroní), Amazonas (Puerto
Ayacucho, Río Parucito). Anzoátegui, Apure,
Carabobo, Distrito Federal, Monagas, Por-
tuguesa; Mexico, Central America, Antilles,
Colombia, Guyana, Brazil (Amapá, Maran-
hão, Roraima), Argentina. ŠFig. 307.
Macroptilium lathyroides (L.) Urb., Symb.
Antill. 9: 457. 1928. —Phaseolus lathyr-
oides L., Sp. Pl. ed. 2, 2: 1018. 1762
[1763].
varieties, 1 in Venezuela.
M. lathyroides var. lathyroides
Erect herb; flowers purple-red to black.
Savannas, edges of towns, 50–200 m; Delta
Amacuro (Tucupita to La Horqueta road),
Bolívar (Ciudad Bolívar, El Palmar), Ama-
zonas (Puerto Ayacucho, Río Manapiare).
Barinas, Guárico, Miranda, Portuguesa;
Mexico, Central America, Antilles, Colombia,
Ecuador, Peru, Brazil, Bolivia, Paraguay, Ar-
gentina. ŠFig. 306.
Macroptilium longepedunculatum (Mart.
ex Benth.) Urb., Symb. Antill. 9: 457. 1928.
—Phaseolus longepedunculatus Mart. ex
Benth., Comm. Legum. Gen. 77. 1837.
Phaseolus campestris Mart. ex Benth.,
Comm. Legum. Gen. 77. 1837.
354 F ABACEAE

Fig. 306. Macroptilium

lathyroides var. lathyroides
Fig. 305. Macroptilium
monophyllum
Fig. 307. Macroptilium
gracile
 Monopteryx 355

Perennial vine; flowers red to violet. Dis- Macroptilium monophyllum (Benth.)

turbed places, Trachypogon savannas, 50–500
m; Bolívar (Altiplanicie de Nuria, Ciudad
Bolívar, middle Río Orinoco), Amazonas (La-
guna El Sillón 78 km northeast of Puerto
Ayacucho, lower Río Ventuari). Lara, Portu-
guesa; Mexico, Central America, Antilles, Co-
lombia, Guyana, Suriname, French Guiana,
Peru, Brazil (Amapá, Goiás, Mato Grosso),
Argentina.
Maréchal & Baudet, Bull. Jard. Bot. Belg.
47: 257. 1977. —Phaseolus monophyllus
Benth., Comm. Legum. Gen. 76. 1837.
Perennial herb or vine; flowers yellow to
orange. Savannas, 50–200 m; Bolívar (Mari-
pa, Río Ore in Río Parguaza basin), Amazonas
(Maypures, Puerto Ayacucho, Rincones de Cha-
chorro 30 km north of Puerto Ayacucho). Mo-
nagas; Colombia, Brazil, Paraguay. ŠFig. 305.

39. MONOPTERYX Spruce ex Benth. in Mart., Fl. Bras. 15(1): 307. 1862.
by Charles H. Stirton and Gerardo A. Aymard C.
Large trees to 50 m tall. Stipules small, elliptic-obovate, caducous. Leaves 3–
11-foliolate, odd-pinnate. Leaflets coriaceous, inserted opposite or alternate, small
and many or few and large. Inflorescences terminal panicles or racemose in upper
axils. Flowers pink, bisexual, zygomorphic, pedicellate, bracteolate, bracteate. Calyx
lobes united in 2 lips, the larger covering the flower in bud and the lower small with
3 very short teeth; petals sessile; standard obovate to suborbiculate; keel petals
subfalcate-oblong, fused along base and at apex, longer than the free wing petals.
Stamens subequal, kinked, free; anthers linear-oblong. Ovary stipitate, glabrous or
sparsely pilose, 1-ovulate; style short conical, arcuate, stigma laterally introrse.
Fruit flattened, arcuate, tapering at both ends, finely transversely ridged, with apex
acute.
Colombia, Venezuela, French Guiana, Brazil; 4 species, all in the flora area.

Key to the Species of Monopteryx

1. Rachis of the leaves canaliculate, ± winged at the base; petiolules 0.2–2

mm long; leaflets oblong, obovate, or obovate-elliptic, drying brownish
on the lower surface, apex rounded; fruit falcate ................................. 2
1. Rachis of the leaves not canaliculate, not subwinged at the base; peti-
olules > 4 mm long; leaflets ovate, elliptic, ovate-lanceolate, not drying
brownish on the lower surface, apex acuminate; fruit not falcate ....... 3
2(1). Leaves 3- or 5-foliolate; terminal leaflets 3–4 cm wide ..................... M. sp. A
2. Leaves 7–15-foliolate; terminal leaflets 1–2 cm wide ....................... M. inpae
3(1). Leaflets glaucous, densely minutely tomentulose and not reticulate on
the lower surface; calyx 9–12 mm long .............................. M. angustifolia
3. Leaflets glabrous, not glaucous, strongly reticulate on the lower surface;
calyx 6–9 mm long ........................................................................ M. uaucu

Monopteryx angustifolia Spruce ex woody, flat, dehiscent. Nonflooded evergreen

Benth. in Mart., Fl. Bras. 15(1): 307, t.
122. 1862. —Barbasco.
Tree 20–30 m tall, with buttressed trunks
and hard wood; leaves (5)7- or 9-foliolate,
ovate-lanceolate, coriaceous, the lower sur-
face densely minutely tomentulose, the up-
per surface nitid; petals dark pink; fruit
lowland forests, 50–200 m; Amazonas (Río
Atabapo, Yavita). Colombia, Brazil (Ama-
zonas: Rio Negro). ŠFig. 308.
Monopteryx inpae W.A. Rodrigues, Acta
Amazon. 5: 153, fig. 1. 1975.
Tree to 35 m tall; lower surface of leaflets
356 F ABACEAE

Fig. 308. Monopteryx angustifolia

drying brownish; fruits narrowly falcate,

with a winged border on both sides of le-
gume. Evergreen lowland forests on sandy
soil, 100–200 m; Amazonas (Río Mawa-
rinuma). French Guiana, Brazil (Amapá,
Amazonas).

Monopteryx uaucu Spruce ex Benth. in

Mart., Fl. Bras. 15(1): 308. 1862.
—Guaco.
Tree to 40 m tall, with buttressed trunks.
Evergreen lowland forests on ultisols, 50–
200 m; Amazonas (San Carlos de Río Negro
to Solano, Yavita to Maroa). Colombia, Brazil
(Amazonas).

Monopteryx sp. A
Tree ca. 15 m tall; fruits dark brown, fal-
cate, finely transversely ridged near matu-
rity. Lower montane forests, 200–300 m;
Bolívar (Minas de Manaima in middle Río
Paragua). Endemic.
The only specimen of this species, Stergios
10259 (MO, PORT), is in fruit only. It ap-
pears most closely related to Monopteryx
inpae.
 Mucuna 357

40. MUCUNA Adans., Fam. Pl. 2: 325. 1763, nom. cons.

Stizolobium P. Browne, Civ. Nat. Hist. Jamaica 290. 1756.
by Gerardo A. Aymard C.
Herbs to climbing woody vines. Leaves alternate, pinnately 3-foliolate; lateral
leaflets oblique; stipules and often stipels deciduous. Inflorescences axillary on leafy
shoots or on old branches, subumbellate, condensed-paniculate, or falsely racemose
through reduction of lateral branches, mostly pendent; bracts and bracteoles decidu-
ous, often subfoliaceus, enclosing the bud. Flowers showy, purple, reddish, orange,
greenish, yellow, or white. Calyx campanulate, 4- or 5-toothed, the upper teeth con-
nate and forming an entire or bifid lip, the lower 3 usually unequal, often with irri-
tating trichomes; standard rounded with median claw and pair of inflexed lateral
auricles at base, usually much shorter than other petals; wing and keel petals nar-
rowed into basal claw with small dorsal auricle, margins basally ciliate, the keel pet-
als partially connate along lower margin, usually horny, apically falcate and indu-
rate. Stamens 10, diadelphous, the filaments alternately thick and thin, long and
short, the vexillar stamen free; anthers barbate. Ovary tomentose, the short stipe
surrounded by a glandular disk; style slender, glabrous or pubescent; stigma small,
capitate, sometimes with a tuft of trichomes. Fruit ovoid, oblong, or linear, thick or
flattened, the margins often winged, undulate between the seeds and ± compressed
laterally between the seeds, the surface sometimes layered with parallel or irregu-
larly raised thin plates which may form elongated outgrowths or 1 or 2 transverse
ribs, usually densely covered with stiff irritating trichomes, usually dehiscent, occa-
sionally indehiscent. Seeds globose or oblong, the hilum narrow, encircling more
than 1/2 of the periphery.
Pantropics; 120 species, 5 in Venezuela, 3 of these in the flora area.

Key to the Species of Mucuna

1. Fruit 4–9 × 1–1.5 cm, without prominent transverse thin plates, but with
2 horizontal ribs on each valve; flowers purple with white tips ............
................................................................................................... M. pruriens
1. Fruit ≥ 10 × 2–7 cm, with prominent transverse thin plates; flowers red-
dish orange, yellow-green, or pale greenish white ............................... 2
2(1). Flowers 6–9 cm long, reddish orange; inflorescences 6–13(–15) cm long;
peduncle 1–1.2 cm long; anthers pilose .................................... M. rostrata
2. Flowers ≤ 5 cm long, pale greenish white or yellow-green; inflorescences >
15 cm long; peduncle > 15 cm long; anthers basally barbate ....... M. urens

Mucuna pruriens (L.) DC., Prodr. 2: 405.
1825. —Dolichos pruriens L., Herb.
Amb. 23. 1754. —Stizolobium pruriens
(L.) Medik., Vorles. Churpfälz. Phys.-
Öcon. Ges. 2: 399. 1787. —Caraota,
Nescafé, Pica Pica, Toddy.
Paleotropics, introduced in the Neotropics
for food and forage; 3 varieties, 2 in Venezu-
ela, 1 of these in the flora area.
M. pruriens var. utilis (Wall. ex Wight)
Baker ex Burck, Ann. Jard. Bot.
Buitenzorg 11: 187. 1893. —Mucuna
utilis Wall. ex Wight, Icon. Pl. Ind. Orient.
1: 280. 1840.
Stizolobium aterrimun Piper & Tracy,
U.S.D.A. Bur. Pl. Industr. Bull. 179: 18.
1910.
Stizolobium hassjo Piper & Tracy,
358 F ABACEAE
U.S.D.A. Bur. Pl. Industr. Bull. 179: 17.
1910. —Mucuna hassjoo (Piper & Tracy)
Mansf., Kulturpflanze 7: 204. 1959.
Vine, occasionally high-climbing; flowers
purple. Cultivated or escaped on disturbed
ground, 50–200 m; Bolívar (near El Dorado,
Tumeremo), Amazonas (Isla Ratón). Wide-
spread elsewhere in Venezuela; Mexico, Cen-
tral America, West Indies, Colombia, Suri-
name, Ecuador, Peru, Brazil, Bolivia. ŠFig.
309.
Mucuna rostrata Benth. in Mart., Fl. Bras.
15(1): 171. 1859. —Guaraguao, Ojo de
burro, Ojo de zamuro, Jijije.
High-climbing woody vine; flowers orange
to red. Evergreen lowland to lower montane
flooded or nonflooded forests, near sea level
to 500 m; Delta Amacuro (Caño Arature,
Caño Curiapo, Caño Guará, La Horqueta,
Punta Barima), Bolívar (base of Auyán-
tepui, Río Uruyen). Widespread elsewhere in
Venezuela; Guatemala, Belize, Honduras,
Panama, Colombia, French Guiana, Ecua-
dor, Peru, Brazil (Amazonas, Pará), Bolivia.
ŠFig. 310.
Fig. 309. Mucuna pruriens var. utilis
Mucuna urens (L.) Medik., Vorles.
Churpfälz. Phys.-Öcon. Ges. 2: 399.
1787. —Dolichos urens L., Syst. Nat. ed.
10, 2: 1162. 1759. —Bejuco de zamuro,
Calabazín, Jeamo, Ojo de zamuro, Pepa
de zamuro.
Dolichos altissmus Jacq., Enum. Syst. Pl.
27. 1760. —Stizolobium altissimum
(Jacq.) Pers., Syn. Pl. 299. 1807. —Mu-
cuna altissima (Jacq.) DC., Prod. 2: 405.
1825.
High-climbing vine; flowers pale greenish
white with dull purple. Humid or gallery for-
ests, 50–200 m; Delta Amacuro (Caño
Araguabisi between Caño Araguao and Caño
Güiniquina, Caño Joba-Suburu east of Caño
Sacupana), Bolívar (widespread), Amazonas
(Caño Yureba, near La Esmeralda, Río
Casiquiare, Río Mawarinuma, Río Padamo,
Río Parú). Widespread elsewhere in Venezu-
ela; Mexico, Guatemala, Costa Rica, Pana-
ma, West Indies, Suriname, Peru, Brazil, Ar-
gentina. ŠFig. 311.
 Mucuna 359

Fig. 310. Mucuna rostrata

Fig. 311. Mucuna urens

360 F ABACEAE

41. MUELLERA L. f., Suppl. Pl. 53, 329. 1781 [1782], nom. cons.
Coublandia Aubl., Hist. Pl. Guiane 937. 1775.
by Nidia L. Cuello A.
Shrubs or small trees, branches glabrous to sparsely pilose. Leaves alternate,
odd-pinnate; leaflets 5 or 7, subopposite, upper surfaces glabrous, lower surfaces
puberulent; stipules minute; stipels lacking. Inflorescences in axillary or lateral
racemes; bracts and bracteoles small, inconspicuous, caducous. Flowers in groups on
short peduncles along an axillary rachis, violet or white. Calyx campanulate, trun-
cate, shortly 5-dentate or with very short or obsolete teeth; petals clawed, glabrous;
standard broadly ovate or suborbicular, auricle callosities absent; wing petals fal-
cate-oblong, auriculate, slightly adherent to the keel petals; keel petals incurved,
obtuse, shorter than the wings. Stamens 10, monadelphous, connate into a tube
closed at both sides, vexillar stamen free at the base but connate from the middle
with the others into a closed tube; anthers versatile. Ovary subsessile; ovules nu-
merous; style filiform, incurved; stigma, small, terminal. Fruit thick, fleshy, hard,
indehiscent, subterete, constricted between the seeds and moniliform, or by abortion
1-seeded and subglobose. Seeds ovoid-subglobose, slightly compressed, with a lat-
eral hilum.
Mexico, Central America, Antilles, Colombia, Trinidad, Guyana, Suriname,
French Guiana, Peru, Brazil, Argentina; 2 species, 1 in Venezuela.

Fig. 312. Muellera frutescens

 Myrocarpus 361

Muellera frutescens (Aubl.) Standl., Tropi- level to 50 m; Delta Amacuro (Angosturita,

cal Woods 34: 41. 1933. —Coublandia
frutescens Aubl., Hist. Pl. Guiane 937, pl.
356. 1775.
Muellera moniliformis L. f., Suppl. Pl. 59,
329. 1781 [1782].
Tree 2–7 m tall; flowers purple. Flooded
and wet forests, mangrove swamps, near sea
Caño Araguao, Caño Bagre, Caño Joba-
Suburu east of Caño Sacupana, Pedernales).
Sucre; Mexico, Central America, Colombia,
Trinidad, Guyana, Suriname, French Gui-
ana, Peru, Brazil (Amapá, Maranhão, Pará).
ŠFig. 312.

42. MYROCARPUS Allemão, Pl. Novas. Brasil pl. 5. post 26 Oct. 1847.
by Gerardo A. Aymard C.
Large trees, with hard wood and resinous bark. Leaves odd-pinnate, (1–)3–10-
foliolate, alternate; leaflets chartaceous, alternate, with pellucid dots; stipules
small, stipels lacking. Inflorescences axillary or pseudoterminal, many-flowered spi-
cate racemes (almost catkin-like); bracts minute, deltoid. Flowers small,
subactinomorphic, pedicellate. Calyx turbinate-campanulate, with 5 short, subequal
lobes, the vexillar lobes sometimes connate; petals (3–)5, free, whitish, imbricate,
subequal, linear, clawed. Stamens 6, 8, or 10; filaments free, subequal, mostly ex-
ceeding the petals; anthers subcordiform, basifixed, dehiscent by lateral slits. Ovary
shortly stipitate, usually 3–5-ovulate; style short, inflexed; stigma minute, terminal.
Fruits elongate, laterally compressed, samaroid with marginal wings, resinous.
Seeds 1–5, oblong or compressed-fusiform, the hilum apical.
Venezuela, Brazil, Bolivia, Paraguay, Argentina; 4 species, 1 in Venezuela.
Myrocarpus venezuelensis Rudd, Phyto- Bolívar (mouth of Río Nichare), Amazonas
logia 23: 404. 1972. —Cereipo, Kama- (Río Oinoquito in upper Río Ofinoco basin).
kari (Yekwana), Nosamo de casa Apure (San Camilo). ŠFig. 313.
(Yekwana). This species has purple heartwood and is
Large tree to 20 m tall; outer bark corky, extremely resistant to termites; it is used lo-
with strong lenticels; fruits elongate, ca. 9 × cally in house construction (center posts) and
2 cm. Evergreen lowland forests, 50–400 m; in wood carving.

Fig. 313. Myrocarpus venezuelensis

362 F ABACEAE

43. MYROSPERMUM Jacq., Enum. Syst. Pl. 4: 20. 1760.

by Charles H. Stirton
Large shrubs or small trees. Stipules minute or lacking. Leaves 10–25-
foliolate, odd-pinnate, alternate, deciduous. Leaflets chartaceous, alternate to
subopposite, with pellucid dot and linear balsam glands. Inflorescences axillary or
terminal racemes. Flowers white, zygomorphic, bracteate, bracteolate. Calyx turbi-
nate-campanulate with 5 subequal lobes, valvate in bud; petals 5, unequal, free, gla-
brous, clawed; standard white or yellow with a nectar patch, elliptic to obovate; wing
petals smaller and narrower than keel petals. Stamens 10, free, persistent; anthers

Fig. 314. Myrospermum frutescens

 Myroxylon 363

equal, uniform, ovoid or ellipsoid, dorsifixed on elongated filaments. Ovary stipitate,

5–7-ovulate, pubescent or glabrous; style minute or distinct; stigma small, terminal,
capitate. Fruits stipitate, compressed, indehiscent, samaroid (apically 1-seeded) and
distinctly winged, or oblong (1–3-seeded) and scarcely winged. Seeds oblong-reni-
form, the hilum subapical, elliptic.
Mexico, Guatemala, Nicaragua, Costa Rica, Panama, Puerto Rico, Colombia,
Trinidad, Venezuela; 3 species, 1 in Venezuela.

Myrospermum frutescens Jacq., Enum. Mexico, Central America, Colombia. ŠFig.

Syst. Pl. 4: 20. 1760. —Cereipo.
Large shrub or small tree to 12 m tall;
leaves 10–21-foliolate, oblong to elliptic, sec-
ondary veins inconspicuous; fruits flat,
winged, indehiscent. Deciduous forests and
shrublands, granitic outcrops, 100–300 m;
Bolívar (50 km southwest of Caicara in
Serranía La Encaramada, islands in Lago
Guri, Puerto Ordaz). Anzoátegui, Falcón,
Guárico, Portuguesa, Yaracuy; southern
314.
Myrospermum frutescens is the emblem
species of Anzoátegui state. It is often con-
fused with Myrospermum secundum Klotzsch,
which is found in the West Indies, north-cen-
tral Venezuela, and Trinidad, but this is a
much larger fruited species. Myrospermum
frutescens has also been confused with the
genus Myroxylon, which has similar fruits
but quite different flowers.

44. MYROXYLON L. f., Suppl. Pl. 34, 233. 1781., nom. cons. —Myrospermum sect.
Myroxylon (L. f.) DC., Prodr. 2: 95. 1825.
Toluifera L., Sp. Pl. 384. 1753.
by Charles H. Stirton
Trees to 40 m tall. Stipules minute or lacking. Leaves 5–15-foliolate, odd-pin-
nate, alternate, deciduous. Leaflets chartaceous, alternate, lanceolate to elliptic,
acute to acuminate, with pellucid glandular dots and lines, margins undulate. Inflo-
rescences axillary or in terminal racemes. Flowers white, bisexual, zygomorphic;

Fig. 315. Myroxylon balsamum

364 F ABACEAE

bracteate and bracteolate, caducous. Calyx turbinate-campanulate, with 5 subequal

lobes, valvate in bud; petals 5, free, unequal, the standard much larger, the other 4 ±
equal. Stamens 10, free, persistent, exserted; anthers uniform oblong, acuminate,
sagittate, dorsifixed on short filaments. Ovary stipitate, 2-ovulate near apex,
smooth; style short, subulate; stigma terminal, minute. Fruits stipitate, compressed,
indehiscent, samaroid (apically 1-seeded), distinctly winged. Seeds reniform, resin-
ous, the hilum subapical, elliptic.
Mexico, Central America, Colombia, Venezuela, Guyana, Ecuador, Peru, Bra-
zil; 2 variable species, 1 in Venezuela.

Myroxylon balsamum (L.) Harms, Notizbl.
Königl. Bot. Gart. Berlin 5: 94. 1908.
—Toluifera balsamum L., Sp. Pl. 384.
1753. —Kamabadek (Pemón).
Large tree with buttressed trunks; leaves
5–15-foliolate, elliptic, one half usually
broader than the other, arcuate; young
branchlets with white lenticels. Cultivated
in the flora area, and possibly escaping.
Carabobo, Miranda, Yaracuy; Mexico, Cen-
tral America, Colombia, Suriname, Ecuador,
Peru, Bolivia. ŠFig. 315.
This species has been used for hundreds
of years as incense and as a medicinal plant.
It is still used in the perfume trade and as a
medicine in other countries. A sterile speci-
men from Río Paragua 13 km north of
Karún, A. Fernández 4618 (MO, PORT), ap-
pears to be this species.

45. ORMOSIA Jack., Trans. Linn. Soc. London 10: 358, t. 25. 1811. nom. cons.
by Charles H. Stirton and Gerardo A. Aymard C.
Shrubs to very tall trees. Clusters of pearl bodies usually present in axils of
leaflets, pulvinus, bracts, bracteoles or inflorescence branches, few to many. Leaves
1–21-foliolate, odd-pinnate, alternate, persistent, stipellate but early caducous; leaf-
lets chartaceous or coriaceous, alternate to opposite; stipules small or lacking, del-
toid to linear, caducous. Inflorescences axillary or terminal racemes. Flowers from
white, yellow, pink, mauve, or blackish purple, with variously colored nectar
patches, bisexual, zygomorphic, bracteate, bracteolate. Calyx campanulate, hy-
panthoid, with equal or subequal teeth or lobes, vexillar teeth fused for half their
length and with a convex sinus; petals 5, unequal, free, glabrous, clawed; standard
suborbicular or broadly ovate, emarginate, darker in color than other petals, broadly
ovate to obovate; wing petals oblique, obovate-oblong, longer and broader than keel
petals, petal sculpturing arching, transcostal; keel petals often imbricate along
lower margin. Stamens 10, free, subequal in 2 or 3 ranks; filaments all thin or vari-
ously thickened at the base, inserted at the apex of the hypanthium; anthers equal,
dorsifixed. Ovary sessile to stipitate, 2–7-ovulate, pubescent; style elongate, re-
curved; stigma introrse, bilobed. Fruits usually dehiscent, a few indehiscent, gla-
brous to densely velutinous, compressed to turgid, with or without septa, oblong to
elongate, apex usually acute. Seeds ellipsoid, globose, or lenticular, 1–6, monochro-
matic (red, scarlet, yellow, black) or dichromatic (red and black or orange and black),
usually polymorphic, the hilum terminal or subterminal, linear or elliptic.
Neotropics, southeast Asia to northeast Australia; 145 species, 23 in Venezuela,
18 of these in the flora area.
A collection (Gentry & Stein 46858, MO) from Río Mawarinuma, Amazonas
state, may represent a new species, but fruits and flowers are needed to determine
this with certainty.
 Ormosia 365

Key to the Species of Ormosia

1. Lower surface of leaflets sparsely minutely appressed-pubescent, glabres-

cent, or glabrous ..................................................................................... 2
1. Lower surface of leaflets densely sericeous, tomentose, or velutinous
.............................................................................................................. 10
2(1). Apex of leaflets conspicuously acuminate; secondary veins on lower sur-
face inconspicuous or scarcely raised; fruits black; seeds bright red
............................................................................................... O. grandiflora
2. Apex of leaflets acute, obtuse, shortly acuminate, emarginate, or trun-
cate; secondary veins on lower surface slightly to conspicuously raised;
fruits vermillion, dark brown, or chestnut brown; seeds dull red or red
and black ................................................................................................ 3
3(2). Apex of leaflets retuse or emarginate; fruits 3–4 × 1.5–2 cm, not woody
................................................................................................................ 4
3. Apex of leaflets acute, obtuse, or shortly acuminate; fruits > 5 × 3 cm,
woody ...................................................................................................... 7
4(3). Leaflets ovate, oblong, lanceolate, or ovate-lanceolate; fruits indehiscent
or very tardily dehiscent, broadly beaked, fulvous-tomentose ..............
................................................................................................. O. williamsii
4. Leaflets elliptic to obovate; fruits dehiscent, acutely beaked, glabrate
................................................................................................................ 5
5(4). Upper surface of leaflets smooth; valves of fruit ± woody or coriaceous, 1–
2 mm thick ................................................................................ O. costulata
5. Upper surface of leaflets rugose; fruit woody, 2–7 mm thick ................... 6
6(5). Leaflets with secondary veins 3–10 mm apart; inflorescences ferruginous-
tomentulose; beak of fruit straight ........................................... O. coccinea
6. Leaflets with secondary veins 10–25 mm apart; inflorescences fulvous-
tomentulose; beak of fruit arcuate ....................................... O. subsimplex
7(3). Leaflets elliptic-oblong, lower surface with slightly raised secondary
veins, upper surface with major veins flush with the surface; flowers 9–
10 mm long ............................................................................... O. paraensis
7. Leaflets ovate to ovate-oblong, elliptic, or obovate, lower surface with dis-
tinctly raised secondary veins, upper surface with major veins deeply
impressed on the surface; flowers 10–25 mm long ............................... 8
8(7). Leaflets with secondary veins 3–10 mm apart, midvein on lower surface
tomentulose; inflorescences ferruginous-tomentulose; beak of fruit
straight ....................................................................................... O. coccinea
8. Leaflets with secondary veins 10–25 mm apart, midvein on lower surface
puberulent or appressed-pubescent; inflorescences fulvous-tomen-
tulose; beak of fruit arcuate .................................................................. 9
9(8). Leaflets rigid-coriaceous, elliptic to obovate; bracts deltoid or linear-del-
toid, 2–3.5 mm long; calyx ferrugineos-tomentulose, 7–9 mm long
............................................................................................... O. subsimplex
9. Leaflets subcoriaceous to coriaceous, ovate to ovate-oblong; bracts linear,
3–10 mm long; calyx gray-pubescent, 10–15 mm long ........ O. macrocalyx
10(1). Lower surface of leaflets aureo-velutinous ............................................. 11
10. Lower surface of leaflets sericeous or tomentose .................................... 14
366 F ABACEAE

11(10). Secondary veins of leaflets terminating in a series of prominent arches

forming a distinct intramarginal vein ............................... O. macrophylla
11. Secondary veins of leaflets upturned and gradually diminishing apically
inside the margin, connected to superadjacent secondaries by a series
of cross veins without forming a distinct intramarginal vein ............ 12
12(11). Flowers 10–14 mm long, calyx 3–8 mm long; fruits obliquely elliptic or
oblong, sutures not ridged; seeds 1(2) ........................................ O. discolor
12. Flowers 15–25 mm long, calyx 10–15 mm long; fruits moniliform, sutures
distinctly ridged; seeds 2–6 ................................................................. 13
13(12). Lower surface of leaflets golden yellow, the secondary and tertiary vena-
tion prominent; fruits rugose, dehiscent along both sutures, the valves
recurved ....................................................................................... O. nobilis
13. Lower surface of leaflets silvery gray, the secondary and tertiary venation
weakly developed; fruits smooth, tardily dehiscent along lower suture,
the valves not recurved ....................................................... O. bolivarensis
14(10). Leaflets tomentose, secondary veins, almost parallel ............................ 15
14. Leaflets sericeous, secondary veins arcuate ........................................... 17
15(14). Leaflets elliptic or elliptic-oblong; fruits woody, 7–8 × 2.5–4 cm, to 25 mm
thick; seeds scarlet or scarlet and black ............................... O. lignivalvis
15. Leaflets elliptic-ovate, ovate, or obovate; fruits subwoody, 2–5 × 1–1.5 cm,
to 1.1 mm thick .................................................................................... 16
16(15). Young stems fulvo-velutinous; leaf base obtuse, secondary veins 18–
25; calyx externally and fruit velutinous.. ....................... O. maguireorum
16. Young stems cinerous- to fulvous-tomentulose; leaf base rounded to cor-
date, secondary veins 10–15; calyx externally fulvous- to ferruginous-
tomentulose; fruit velutinous essentially glabrous .............. O. amazonica
17(14). Fruits chartaceous, finely brown-velutinous, 15–25 × 11 mm, 1–2 mm
thick ..................................................................................... O. steyermarkii
17. Fruits woody, fulvous-velutinous, 25–40 × 15–25 mm, 2–5 mm thick ... 18
18(17). Leaflets ovate to elliptic, the apex acuminate to attenuate .............. O. sp. A
18. Leaflets oblong-elliptic or obovate, the apex acute to shortly acuminate
.............................................................................................................. 19
19(18). Trees to 30 m tall, pubescence tawny .......................................... O. coarctata
19. Shrubs to 3 m tall, pubescence ferruginous ....................................... O. sp. B

Ormosia amazonica Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 139. 1922.
Ormosia euneura Harms, Notizbl. Bot.
Gard. Berlin-Dahlem 9: 972. 1926.
Tree to 20 m tall; leaves 7–11-foliolate,
leaflets coriaceous, the lower surface finely
and tightly crisp-pubescent. Lowland wet
forests, 200–300 m; Amazonas (Río Mawa-
rinuma). Colombia, Ecuador, Peru, Brazil.
Ormosia bolivarensis (Rudd) Stirton,
comb. nov. —Ormosia nobilis var.
bolivarensis Rudd, Contr. U.S. Natl.
Herb. 32: 345. 1965. —Guanacoco, Wa-
naca-có (Yekwana).
Tree to 10–25 m tall; leaves 5–9-foliolate,
leaflets elliptic-oblong, coriaceous, the lower
surface minutely fulvous-sericeous. White-
sand and forested hills, 400–900 m; Bolívar
(Cerro Camarón, Cerro Ichún, Río Canara-
cuni, upper Río Caura, Río Curutu, Río Ica-
barú, Río Salto) Guyana, Brazil. ŠFig. 316.
Ormosia bolivarensis is distinguished
from O. nobilis by its smaller, pale orange to
orange-yellow seeds and distinctive fulvous-
sericeous lower surfaces of leaves.
Ormosia coarctata Jacks., Trans. Linn.
Soc. London 10: 363, t. 27. 1811.
—Pericoca.

Ormosia cuneata Ducke, Arch. Jard. Bot.
Rio de Janeiro 4: 64. 1925.
Tree to 30 m tall; leaves 5–11-foliolate,
leaflets obovate, coriaceous, the lower sur-
face densely pubescent with laxly crispate to
subpatent trichomes. Humid forests, 200–
300 m; Bolívar (53 km northeast of Los
Rosos, Reserva Forestal Imataca, Río
Botanamo, Río Cuyuní). Mérida; Trinidad,
Guyana, Suriname, French Guiana, Brazil,
Bolivia.
Ormosia coarctata is little collected in the
region. Flowering material is needed from
Venezuela.
Ormosia coccinea (Aubl.) Jacks., Trans.
Linn. Soc. London 10: 360, t. 25, 1811.
—Robinia coccinea Aubl., Hist. Pl.
Guiane 773. 1775. —Gateado, Macure,
Palo macure, Peonía, Peonío, Pionilla,
Pionina, Tento, Too’o (Yanomami), Ya’u
balé.
Large shrub or tree to 30 m tall; leaves 9
(–14)-foliolate; leaflets oblong, coriacious,
large. Forest margins and islands, shrubby
white-sand savannas, 50–200 m; Bolívar
(Piedra Marimare in middle Río Orinoco),
Amazonas (widespread). Apure, Monagas;
Colombia, Guyana, Suriname, French Gui-
ana, Ecuador, Peru, Brazil (Amazonas).
Ormosia costulata (Miq.) Kleinhoonte,
Recueil Trav. Bot. Néerl. 22: 392. 1925.
—Leptolobium costulatum Miq., Stirp.
Surinam. Select. 17. 1850 [1851].
—Peonía.
Tree to 13 m tall; leaves 1–7-foliolate;
leaflets obovate to elliptic, coriaceous, ter-
tiary veins scarcely visible. Sandy savanna
areas; 50–400 m; Bolívar (Río Asa, Río
Paragua). Guyana, Suriname, French Gui-
ana, Brazil (Amazonas). ŠFig. 318.
Ormosia discolor Spruce ex Benth. in
Mart., Fl. Bras. 15(1): 318. 1862.
—Peonía.
Ormosia micrantha Ducke, Arq. Inst. Biol.
Veg. 4: 21. 1938.
Tree to 19 m tall; leaves (3)5- or 7-
foliolate, ovate to oblong, coriaceous, lower
surface fulvous-sericeous; flowers small.
Nonflooded evergreen lowland forests, 100–
200 m; Amazonas (Río Casiquiare, Río Ne-
Ormosia 367
gro, Río Pasimoni, Río Yatúa). Amazon basin
of Colombia and Brazil.
Ormosia grandiflora (Tul.) Rudd, Contr.
U.S. Natl. Herb. 32: 301. 1965. —Dip-
lotropis grandiflora Tul., Arch. Mus.
Hist. Nat. 4: 109. 1844.
Tree or shrub to 10 m tall; leaves (1)3- or
5-foliolate; leaflets ovate, elliptic, or obovate,
subcoriaceous, glabrescent, shiny. Evergreen
lowland forests, ca. 300 m; Amazonas (Río
Sipapo). Costa Rica, Ecuador, Peru, Brazil,
Bolivia.
Ormosia lignivalvis Rudd, Contr. U.S.
Natl. Herb. 32: 331. 1965. —Chaparillo,
Kaguaitariyek (Arekuna), Pericosa,
Wanaekoko (Yekwana).
Tree to 50 m tall; leaves 5–11-foliolate;
leaflets elliptic or oblong, coriaceous, vena-
tion well developed. Forest margins, 200–
500 m; Bolívar (El Dorado, Río Asa, Río
Paragua at mouth of Río Karún, Río Tabaro,
Santa María de Erebato, Serranía Pia-Zoi,
east-southeast of Villa Lola). Guyana,
French Guiana, Peru, Brazil (Amazonas).
Ormosia macrocalyx Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 137. 1922.
Ormosia toledana Standl., Publ. Carnegie
Inst. Wash. 461: 64. 1935.
Ormosia chlorocalyx Ducke, Bol. Técn.
Inst. Agron. N., 2: 23. 1944.
Tree to 40 m tall; leaves 7–11-foliolate;
leaflets ovate to ovate-oblong, sparsely pu-
bescent to glabrescent. Evergreen lowland
forests, 100–200 m; Amazonas (Río Sipapo).
Barinas, Cojedes, Falcón, Portuguesa, Zulia;
Central Mexico, Central America, West
Indies, Colombia, Ecuador, Peru, Brazil.
Ormosia macrophylla Benth., Ann.
Wiener Mus. Naturgesch 2: 88. 1838.
—Kabaituri, Palo macure, Peonía,
Tento.
Tree to 10 m tall; leaves 5–9-foliolate;
leaflets broadly ovate or elliptic, tertiary
veins inconspicuous. Shrublands, white-
sand savanna edges, scrubby forests, 50–600
m; Amazonas (Caño Chimoni of Río Casi-
quiare, slopes of Cerro Aracamuni, base of
Cerro Yapapana, Río Barí, Río Guasacavi,
Río Pasimoni, Río Siapa, San Carlos de Río
368 F ABACEAE
Negro, Santa Rosa de Ucata). Colombia
(Amazonas, Vaupés), Ecuador, Brazil (Ama-
zonas). ŠFig. 317.
Branches of Ormosia macrophylla often
become hollowed out and are occupied by
ants.
Ormosia maguireorum Rudd, Contr. U.S.
Natl. Herb. 32: 351. 1965.
Tree 15–25 m tall; leaves 9-foliolate; leaf-
lets elliptic-ovate, coriaceous, venation pro-
minent, densely hairy. Montane forests,
1000–1200 m; Amazonas (Río Yatúa on Si-
erra de la Neblina). Endemic.
No collections with flowers have yet been
obtained of this species and are needed to
better characterize it.
Ormosia nobilis Tul., Arch. Mus. Hist. Nat.
4: 106. 1844.
Neotropics; 2 varieties, 1 in Venezuela.
O. nobilis var. nobilis
Shrub or tree to 10 m tall; leaves 7- or 9-
foliolate, elliptic to elliptic-oblong, coria-
ceous, finely velutinous, glabrescent. Savan-
nas, scrubby forests, 100–300 m; Bolívar (up-
per Río Caura), Amazonas (Caño Caname,
Río Aracamuni, Río Pasimoni, Río Siapa).
Colombia, Guyana, French Guiana, Suri-
name, Brazil (Amazonas, Maranhão, Pará),
Bolivia.
Ormosia paraensis Ducke, Arch. Jard. Bot.
Rio de Janeiro 4: 62. 1925. —Kapakun,
Metari-yek, Mureyenu-yek (Arekuna),
Mutare-yek, Peonía, Peonilla, Peonío,
Pionío.
Ormosia crassicarpa Pierce ex Pittier, Bol.
Soc. Venez. Ci. Nat. 10: 108. 1944.
Ormosia heterophylla Pires, Bol. Técn.
Inst. Agron. N. 38: 24. 1960.
Tree to 20 m tall; leaves (1–)7–15-
foliolate; leaflets oblong-elliptic, coriaceous,
secondary veins inconspicuous. Lowland and
upland forests, 50–1500 m; Bolívar (Gran
Sabana, Ptari-tepui, Río Asa, lower Río
Caura, Río Paragua, Río Turiba), Amazonas
(Caño Cucurital in Río Ventuari basin, 25 km
northeast of Puerto Ayacucho, Río Matacuni,
Serranía Batata 55 km southeast of Puerto
Ayacucho). Mérida, Zulia; Costa Rica, Pa-
nama, Colombia, Guyana, Suriname, French
Guiana, Ecuador, Peru, Brazil. ŠFig. 320.
Ormosia steyermarkii Rudd, Contr. U.S.
Natl. Herb. 32: 352. 1965. —Ormosia
microsperma Pittier, Bol. Soc. Venez. Ci.
Nat. 10: 109. 1945, non Baker 1868
[1867]. —Mutare-yek (Arekuna), Pro-
nilla.
Tree to 20 m tall; leaves 3–7-foliolate; leaf-
lets ovate, coriaceous, the lower surface densely
hairy, secondary veins prominent. Nonflooded
montane forests, 800–1600 m; Bolívar (Salto
de Pacairao near Kavanayén, Sororopán-
tepuí). Brazil (Amazonas, Roraima).
A collection in fruit from the Río Ayaiche,
Steyermark 89475, may be conspecific with
O. steyermarkii Rudd. or may represent a
new taxon.
The resinous wood of Ormosia steyer-
markii is said to be used to kindle fires.
Ormosia subsimplex Spruce ex Benth. in
Mart., Fl. Bras. 15: 316. 1862. —Or-
mosia coccinea var. subsimplex (Spruce
ex Benth.) Rudd, Contr. U.S. Natl. Herb.
32: 328. 1965. —Peonía.
Tree to 13 m tall; leaves 7–11-foliolate; leaf-
lets ovate, elliptic or oblong, coriaceous, sec-
ondary veins broadly spaced, tertiary veins
weak. Riparian forests in and around savanna
areas, 50–200 m; Bolívar (Río Parguaza),
Amazonas (lower Río Ventuari, San Fernando
de Atabapo, Solano). Apure; Panama, Brazil
(Amazonas: Rio Aracá). ŠFig. 319.
Ormosia subsimplex is related to O.
coccinea but has differently shaped fruits
and corollas, larger flowers, widely spaced
secondary veins and scarcely developed ter-
tiary venation.
Included here are Fariñas et al. 331
(VEN) from Amazonas state and Davidse &
González 12429 (MO) from Apure state.
These are trees to 8 m tall, with strongly
beaked fruits, and may represent a new spe-
cies.
Ormosia williamsii Rudd, Contr. U.S. Natl.
Herb. 32: 313. 1965. —Peonía rebalsera,
Peonío, Tento.
Tree to 15 m tall; leaves 5–9-foliolate;
leaflets ovate or oblong, coriaceous, glabres-
cent. Riparian black-water forests, 100–200
m; Amazonas (Caño Magua, Piedra Cocuy,
Río Atabapo, Samariapo, near San Carlos de
Río Negro). Colombia, Brazil (Amazonas: Rio
Negro).
 Ormosia 369

Fig. 316. Ormosia bolivarensis

Fig. 317. Ormosia macrophylla

370 F ABACEAE

Fig. 318. Ormosia costulata

Fig. 319. Ormosia subsimplex

 Ormosia 371

Fig. 320. Ormosia paraensis

372 F ABACEAE

Ormosia sp. A per base of petiole flattened; leaflets to

Tree to 25 m tall; leaves 5- or 7-foliolate;
leaflets ovate to elliptic, subcoriaceous, the
lower surface densely tawny; flowers not
known. Montane forests, 1300–1400 m; Bo-
lívar (La Escalera, Sierra de Lema). Endemic.
Ormosia sp. B
Shrub to 3 m; leaves 7-foliolate, the up-
20 cm long, elliptic-oblong, coriaceous,
densely ferruginous-pubescent with cris-
pate trichomes, tertiary veins visible, the
upper surface dark green, nitid, and
rugulate, the lower surface tawny-brown.
Secondary forests, 1100–1300 m; Ama-
zonas (35 km northeast of Parima B). En-
demic.

46. PACHYRHIZUS Rich. ex DC., Prodr. 2: 402. 1825, nom. cons.

by Nidia L. Cuello A.
Twining annual, rarely semierect perennial herbs to somewhat woody vines.
Roots tuberous. Leaves alternate, pinnately 3-foliolate; stipules linear; petioles with
prominent pulvinus; leaflets entire, lobed, or deeply dissected, with linear, caducous
stipels. Inflorescences axillary pseudoracemes; bracts progressively of reduced size
from main axis to lateral axis; peduncle often longer than the petiole, the flowers
somewhat congested apically; bracteoles subulate, shorter than and subtending the
calyx; pedicels slender, slightly longer than the calyx. Flowers blue, floral prophylls
caducous and silky. Calyx campanulate-tubular, 4-lobed with upper lobe larger and
emarginate; standard suborbicular, with green basal spot, emarginate and with 2
basal incurved auricles enclosing the auricles of the wings; wing petals medially ad-
herent to keel petals; keel petals obliquely oblanceolate, recurved with a hump,
equaling the wings, blades connate, not auriculate. Stamens 10, diadelphous; an-
thers elliptical, dorsifixed. Ovary subsessile, with a crenulate disc round its base;

Fig. 321. Pachyrhizus erosus

 Phaseolus 373

ovules numerous; style incurved, sometimes forming a circle, ventrally ciliate, dor-
sally glabrous; distal part of stigma subglobose on the vertical surface, widening at
apex. Legume linear-oblong, internally septate between seeds, externally contracted
between seeds. Seeds flat and suborbicular to flat and square, or plump and reni-
form.
Neotropics; 5 species, 2 in Venezuela, 1 of these in the flora area.
The genus is widely cultivated for the edible tuberous roots.

Pachyrhizus erosus (L.) Urb., Symb.
Antill. 4: 311. 1905. —Dolichos erosus L.,
Sp. Pl. 726. 1753. —O’ mochahcho
(Panare).
Herbaceous climbing or trailing vine.
Semideciduous forests bordering savannas,
50–100 m; Bolívar (El Corozal, lower Río
Parguaza). Aragua, Distrito Federal, Portu-
guesa; Mexico, Central America, West
Indies, French Guiana, Brazil, Paraguay, Ar-
gentina, introduced and naturalized in the
Paleotropics. ŠFig. 321.

47. PHASEOLUS L., Sp. Pl. 723. 1753; Gen. Pl. ed. 5, 323. 1754.
by Gerardo A. Aymard C.
Vines, herbs, or subshrubs. Leaves trifoliolate; stipules acute, veined, often pu-
bescent, not prolonged below the insertion; stipels often oblong, thin, glabrous; leaf-
lets mostly entire, pubescent or glabrate but minute, hooked trichomes present. In-
florescences axillary, the flowers congested in fascicles along the rachis, the nodes
not swollen, not glandular; rachis eglandular; bracteoles ovate or greatly reduced and
much shorter than the calyx, veined, puberulent, persistent at least until anthesis;
bracts ovate or lanceolate, veined, puberulent; pedicels mostly longer than the calyx.
Flowers blue, purple, violet, yellow, or white. Calyx mostly 2-lipped, the upper teeth
partly united; standard symmetrical, orbicular, basally appendaged; wing pet-

Fig. 322. Phaseolus lunatus

374 F ABACEAE

als partly spiralled, apically hooded; keel petals in 2 or 3 spirals. Vexillar stamen
free, the free part of the others elongate; anthers nearly uniform. Ovary 1–many-
ovulate; style apically thickened, curved in 1.5–2.5 spirals, hairy inside, distally ca-
ducous. Fruit linear-oblong, straight or slightly curved, not septate, compressed or
turgid, sometimes beaked. Seeds 1–many, oblong to reniform, the hilum short and
central.
Neotropics, widely cultivated in temperate and tropical countries; ca. 50 spe-
cies, 4 in Venezuela, 2 of these in the flora area.

Key to the Species of Phaseolus

1. Stipels 1–1.5 mm long; peduncle 2–3 cm long; standard pubescent out-

side; fruits curved ........................................................................ P. lunatus
1. Stipels 2–4 mm long; peduncle < 1 cm long; standard sparsely pilose to
glabrate on the midvein outside; fruits straight ........................ P. vulgaris

Phaseolus lunatus L., Sp. Pl. 724. 1753.
Climbing or trailing vine or erect herb;
flowers yellow or purplish. Escaped from
cultivation, 100–200; Amazonas (Río Ca-
taniapo). Mexico, Central America, Colom-
bia, Ecuador, Peru, Brazil, Bolivia. ŠFig.
322.
Phaseolus vulgaris L., Sp. Pl. 723. 1753.
Neotropics, but cultivated in almost every
country around the world; 2 varieties, 1 in
Venezuela.
P. vulgaris var. vulgaris
Climbing or trailing vine or erect herb;
flowers white or purplish. Northern Bolívar
(cultivated). Widely cultivated elsewhere in
Venezuela, but grows wild in Lara, Mérida,
Portuguesa, Táchira, Trujillo; widely culti-
vated as the source of black beans in almost
every temperate and tropical country.
This species was domesticated in the New
World probably some 8000–10,000 years ago
from a wild ancestral form distributed in the
highlands between northern Mexico and
northern Argentina.

48. PISCIDIA L., Syst. Nat. ed. 2, 1151, 1155, 1376. 1759, nom. cons.
by Gerardo A. Aymard C.
Trees or shrubs, unarmed. Leaves alternate, odd-pinnate, 5–27-foliolate;
stipules (bud scales) obliquely ovate, semiorbicular, or reniform, early caducous;
stipels absent; leaflets opposite. Inflorescences axillary or lateral, usually racemose,
sometimes spicate. Flowers white with pink to purplish markings; bracts at base of
pedicels, ovate, elliptic, or lanceolate, early caducous; bracteoles paired at base of
calyx, ovate, oblong, or linear, caducous. Calyx campanulate with 5 short lobes; stan-
dard suborbicular, usually pubescent on the outer face, but glabrous in one species;
wing petals falcate, oblong, commonly a little longer than the standard, adherent to
the keel; keel petals connate at the base. Stamens 10, monadelphous but the vexillar
filament free at the base; anthers oblong, dorsifixed. Ovary sessile, many-ovulate;
style glabrous above; stigma minutely penicillate. Fruit indehiscent, 1–10-seeded,
compressed, with 4 longitudinal wings. Seeds reniform, tan to reddish or dark
brown, laterally compressed, the hilum lateral, elliptic to suborbicular.
Neotropics; ca. 7 species, 1 in Venezuela.

Piscidia carthaginensis Jacq., Enum. ricensis Pittier, Arb. Legum., part 3,

Syst. Pl. 27. 1760. Trab. Mus. Comercial Venezuela 4 [re-
Piscidia guaricensis (Pittier) Pittier, Mesa printed from Bol. Minist. R.R. E.E. no.
Guanipa 49. 1942. —Lonchocarpus gua- 4–7.] 4: 231. 1928.
 Platymiscium 375

Fig. 323. Piscidia carthaginensis

Small tree; calyx pink; petals white and gas, Nueva Esparta, Portuguesa; Mexico,
pink. Dry to gallery forests, 50–100 m; Bo- Central America, Antilles, Colombia, Ecua-
lívar (Las Bonitas on the middle Río Ori- dor, Peru. ŠFig. 323.
noco). Anzoátegui, Guárico, Miranda, Mona-

49. PLATYMISCIUM Vogel, Linnaea 11: 198. 1837.

by Nereida Xena de Enrich
Trees; bark dark, rough, fissured. Leaves opposite or verticillate, compound,
odd-pinnate, with 5 or 7(9) opposite or subopposite leaflets; stipules coriaceous,
ovate-orbicular or lanceolate, caducous or subpersistent; stipels absent. Inflores-
cence axillary, paniculate, glabrous or variously pubescent. Flowers small, yellow,
numerous; hypanthium small or almost absent. Calyx campanulate, dark to pale
green, glabrous or pubescent, teeth small and unequal, the vexillar ones almost to-
tally connate; corolla yellow; standard ovate-orbicular, glabrous, yellow, with ± con-
spicuous brown nectar guides, without auricles, with a short claw; wing petals ob-
long-elliptic, glabrous, free, auriculate; keel petals slightly connate at the apex, up-
per margin ciliate. Stamens 10, frequently the vexillar one free or partially connate
376 F ABACEAE

to the staminal column; anthers monomorphic, ovate-hastate, versatile with longi-

tudinal dehiscence, extrorse. Pistil glabrous, stipe elongate; ovary with 1 ovule; style
curved, filiform; stigma terminal, inconspicuous. Fruit indehiscent, flat, stipitate,
membranous, oblong-elliptic or narrowly elliptic, with seed in central chamber. Seed
large, reniform, flattened, brown; embryo curved, the cotyledons thin, foliaceous,
the radicle curved, inflexed.
Neotropics; ca. 12 species, 3 in Venezuela, 2 of these in the flora area.
This genus is frequently cultivated as a shade and ornamental tree. The wood
is sold under the name “Panama redwood” in the lumber industry.

Key to the Species of Platymiscium

1. Inflorescences glabrous or scarcely pubescent; flowers pale yellow, 10–14

mm long; calyx pale green, generally glabrous; leaflets 5 or 7, ovate-el-
liptic, ± abruptly acuminate at apex ....................................... P. pinnatum
1. Inflorescences densely pubescent; flowers yellow-orange, 15–17 mm long;
calyx externally dark, golden-pubescent; leaflets 5, ovate-oblong, ob-
tuse at apex ................................................................................ P. trinitatis

Fig. 324. Platymiscium pinnatum

 Platypodium 377

Platymiscium pinnatum (Jacq.) Dugand, Platymiscium trinitatis Benth., J. Proc.

Contr. Hist. Nat. Colomb. 1: 7. 1938
—Amerimnon pinnatum Jacq., Select.
Stirp. Amer. Hist. 200. 1763. —Roble,
Roble blanco, Roble colorado.
Platymiscium polystachyum Benth. in
Seem., Bot. Voy. Herald 111, t. 21. 1853.
Platymiscium polystachyum var. fendleri
Benth., J. Proc. Linn. Soc., Bot. 4: 83.
1860.
Tree to 20 m tall. Deciduous forests,
woody savannas, 100–500 m; Bolívar (island
in Danto Manchado sector of Lago Guri, 30
km south of El Manteco, near El Palmar, La
Paragua, Puerto Ordaz, Río Icabarú, north-
east of Upata). Anzoátegui, Apure, Barinas,
Carabobo, Cojedes, Guárico, Lara, Monagas,
Portuguesa, Zulia; Central America, Colom-
bia, Brazil. ŠFig. 324.
Linn. Soc., Bot. 4: 82. 1860. —Roble,
Roble blanco, Roble colorado.
Platymiscium duckei Huber, Bol. Mus.
Paraense Hist. Nat. 6: 83. 1909.
Platymiscium nigrum Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 157. 1922.
—Platymiscium duckei var. nigrum
(Ducke) Ducke, Arch. Jard. Bot. Rio de
Janeiro 4: 87. 1925.
Tree to 20 m tall. Riparian, montane, and
premontane semideciduous or deciduous for-
ests, 100–500 m; Bolívar (northern decidu-
ous zone, Cerro Cotorra between La Paragua
and mouth of Río Paragua, Temblador on
lower Río Caura, northwest of Tumeremo).
Anzoátegui, Apure, Aragua, Barinas, Falcón,
Lara, Mérida, Miranda, Sucre, Yaracuy; Tri-
nidad, Brazil (Pará).

50. PLATYPODIUM Vogel, Linnaea 11: 420. 1837.

by Gerardo A. Aymard C.
Trees 20–30 m tall; branches pilose-pubescent. Leaves odd- or even-pinnate,
alternate; rachis pilose-pubescent; leaflets 1.5–2.5 × 2–7 cm, in 5–18 pairs, alternate
or irregularly opposite, upper surface glabrous, lower surface puberulent to pubes-
cent, apex emarginate; stipules linear, deciduous; stipels absent. Inflorescences
racemose in the upper leaf axils; bracts and bracteoles small, caducous. Flowers
showy, yellow or yellow-orange, pedunculate; bracts 2, acute, deciduous. Calyx tur-
binate at the base, glabrous or laxly pilose, lobes pubescent within, the upper 2 teeth
broader, connate high up; standard glabrous, obovate, emarginate, not appendaged;
wing petals obliquely obovate or oblong; keel petals oblong or obovate, obtuse, con-
nate at the apex (dorsally). Stamens 10, divided equally among 2 fascicles or some-
times the vexillar stamen free; anthers versatile. Ovary long-stipitate, glabrous or
laxly pilose; ovules 3–5; style filiform; stigma small, suborbicular terminal. Fruit
stipitate, 4–8 mm long, samaroid, indehiscent, woody at the apex, narrowed at the
base into an obliquely oblong venose wing. Seeds 1(2), oblong to reniform.
Panama, Colombia, Venezuela, Brazil, Bolivia, Paraguay; 2 species, 1 in Ven-
ezuela.

Platypodium elegans Vogel, Linnaea 11:
421. 1837.
Panama, Colombia, Venezuela, Brazil,
Bolivia, Paraguay; 2 subspecies, both in Ven-
ezuela, 1 in the flora area.
P. elegans subsp. maxonianum (Pittier)
H.C. Lima, comb. nov. —Platypodium
maxonianum Pittier, Contr. U.S. Natl.
Herb. 18: 234. 1917. —Canalete ama-
rillo.
Tree 20–30 m tall; flowers yellow. Semi-
deciduous to wet forests, 100–300 m; Bolívar
(Túriba). Barinas, Falcón, Portuguesa, Tru-
jillo, Zulia; Panama, Colombia, Brazil, Bo-
livia, Paraguay. ŠFig. 325.
378 F ABACEAE

Fig. 325. Platypodium elegans

subsp. maxonianum

51. POECILANTHE Benth., J. Proc. Linn. Soc., Bot. 4(suppl.): 80. 1860.
by Nidia L. Cuello A.
Small trees or shrubs. Leaves odd-pinnate, sometimes unifoliolate; leaflets al-
ternate or opposite; stipels minute or absent; stipules caducous or inconspicuous.
Flowers in short axillary or lateral racemes or panicles; bracts and bractlets small.
Calyx turbinate at the base, the upper 2 lobes connate into one ± 2-toothed lip; petals
clawed; standard orbiculate, not appendaged; wing petals falcate-oblong or obovate;
keel petals incurved, subrostrate, connate at the back. Stamens all connate into a
sheath split above, sometimes nearly diadelphous; anthers alternately longer and
basifixed or shorter and versatile. Ovary subsessile or shortly stipitate; ovules sev-
eral; style filiform, glabrous, incurved; stigma small, terminal. Fruit flat-com-
 Poecilanthe 379

pressed, woody-coriaceous, 2-valved. Seed obovate, hard and shining, compressed,

with a basal hilum, estrophiolate.
Colombia, Venezuela, Guyana, Suriname, French Guiana, Ecuador, Peru, Bra-
zil, Bolivia, Paraguay, Argentina; ca. 10 species, 2 in Venezuela, both in the flora
area.

Key to the Species of Poecilanthe

1. Pedicels 0.5–1 mm long; flowers 8–10 mm long, purple or red; fruit 4–6.5
cm long .................................................................................... P. amazonica
1. Pedicels 3–4 mm long; flowers 12–15 mm long, brown; fruit 10–13 cm long
................................................................................................ P. hostmannii

Poecilanthe amazonica (Ducke) Ducke,

Bull. Mus. Hist. Nat. (Paris) sér. 2, 4:
734. 1932. —Cyclolobium amazonicum
Ducke, Arch. Jard. Bot. Rio de Janeiro 3:
146. 1922. —Arepito.
Small tree or shrub; flowers purple or red.
Evergreen lowland and flooded forests, 100–
200 m; Amazonas (Caño Morocoto below San
Fernando de Atabapo, Río Sipapo, San
Carlos de Río Negro). Brazil (Amapá,
Amazonas). ŠFig. 326.

Poecilanthe hostmannii (Benth.) Amshoff,

Meded. Bot. Mus. Herb. Rijks Univ.
Utrecht 52: 61. 1939. —Cyclolobium
hostmannii Benth., J. Proc. Linn. Soc.,
Bot. 4: 52. 1860.
Tree; flowers deep maroon. Evergreen
lowland forests, 100–200 m; Amazonas (base
of Cerro Yapacana, Río Baría). Suriname,
French Guiana, Brazil (Amapá, Amazonas).

Fig. 326. Poecilanthe

amazonica
380 F ABACEAE

52. POIRETIA Vent., Mém. Cl. Sci. Math. Inst. Natl. France 1807(1): 4. July 1807,
nom. cons.
by Nidia L. Cuello A.
Herbs or suffrutescents, erect or scandent, 1–4 m tall. Leaves alternate, 4-
foliolate; leaflets chartaceous to coriaceous, linear to ovate, obovate, or suborbicular,
glandular-punctate on lower surface, glands sometimes visible on upper surface, op-
posite; stipules lanceolate, usually glandular, caducous; stipels present at base of
lower pair of leaflets, linear to lanceolate or ovate, usually glandular, caducous. In-
florescences axillary or terminal, racemose, sometimes paniculate or spicate, few- or
many-flowered; bracts stipule-like; bracteoles lacking. Flowers small to medium-
sized, 4–20 mm long. Calyx glandular-punctate, campanulate with 5 subequal lobes
shorter than the tube, the vexillar lobes smaller than the others and somewhat con-
nate; petals yellow, glandular-punctate; standard reflexed, glabrous or nearly so.
Stamens 10, monadelphous; anthers dimorphic, alternately oblong and basifixed or
elliptic and dorsifixed. Ovary sessile, glabrous or pubescent; ovules 1–8; stigma ter-
minal, minutely capitate. Fruit sessile, oblong, laterally compressed, with 1–8 ar-
ticles, glandular-punctate, sometimes verruculose. Seeds brown, reniform, com-
pressed, hilum present, elliptic.
Mexico, Central America, Antilles, Colombia, Venezuela, Ecuador, Peru, Brazil
(most diverse in east-southeast part), Bolivia; ca. 6 species, 1 in Venezuela.

Poiretia punctata (Willd.) Desv., J. Bot.

Agric. 1: 122, pl. 5, fig. 17. 1813. —Gly-
cine punctata Willd., Sp. Pl. 3(2): 1066.
1802.
Herbaceous vine 1–3 m long; flowers
bright yellow. Open areas, forest edges, road-
sides, 100–200 m; Bolívar (La Paragua). Ara-
gua, Carabobo, Distrito Federal, Miranda;
Mexico, Central America, Antilles, Colombia,
Ecuador, Peru, eastern and southeastern
Brazil, Bolivia. ŠFig. 327.

Fig. 327. Poiretia punctata

 Pterocarpus 381

53. PTEROCARPUS Jacq., Select. Stirp. Amer. Hist. 283. 1763, nom. cons.
by Nereida Xena de Enrich, Celia Gil, and Paul E. Berry
Trees; bark smooth or slightly to strongly fissured, exuding a red resin; crown
usually umbrella-shaped; young branches terete, glabrous to densely pubescent, fre-
quently with lenticels. Leaves alternate, compound, odd-pinnate, nonstipulate;
stipels linear or narrowly triangular, caducous, rarely persistent; petiole and peti-
olules terete, 2–5(–6) cm long, glabrous or pubescent; leaflets (3)5–11(13), alternate
or subopposite, occasionally opposite, varying in shape and size along the rachis, gla-
brous or densely pubescent, the base rounded, obtuse, or sometimes acute, the apex
acuminate or emarginate and sometimes mucronate, margins entire. Inflorescences
axillary, paniculate or racemose, glabrescent or densely pubescent; bracts and
bracteoles small, linear or triangular, early caducous or sometimes persistent, gla-
brous or pubescent. Flowers small, 11–20 mm long, numerous, zygomorphic, pedicel-
late or sessile; hypanthium turbinate, frequently basally incurved, 1–4 mm long.
Calyx 5-dentate, teeth small, unequal, the vexillar ones almost joined, apex rounded
or acute; corolla yellow-orange, nectar guide usually whitish or violet at the middle
of the standard; standard 10–19 mm long, orbicular-ovate or obovate-elliptic, gla-
brous, basally attenuate and ending in a small or medium-sized claw, apex slightly
emarginate; wing petals 10–18 mm long, oblong-elliptic or elliptic-obovate, free, gla-
brous, auriculate, basally clawed; keel petals 10–15 mm long, glabrous, auriculate,
connate at the middle and basal parts, basally clawed. Stamens 10, the vexillar one
frequently joined up to 1/4 of the staminal column or shortly joined basally; anthers
versatile, isomorphic, elliptic, longitudinally dehiscent. Ovary 1-locular, stipitate to
sessile, glabrous or pubescent; ovules 2–7; style incurved, filiform, glabrous; stigma
terminal, inconspicuous. Fruit flattened, indehiscent, winged, orbicular or subor-
bicular, sometimes asymmetric, stipitate or sessile, glabrous or glabrescent, mem-
branous or coriaceous; seed chamber hard, central, distal, or lateral, sometimes par-
titioned. Seeds 1–3, reniform; testa smooth or slightly rugose, the hilum conspicu-
ous.
Pantropics; ca. 20 species, 5 in Venezuela, all of these in the flora area.
The wood from species in this genus is used for construction, musical instru-
ments, crafts, and toothpicks. The resin is called “sangre de drago” and is widely
used in popular medicine.

Key to the Species of Pterocarpus

1. Fruit with a prominent, membranous or chartaceous wing; seed chamber

in central or proximal position; inflorescence densely pubescent, a
simple raceme or else with 1–3 short branches at the base of the main
axis; calyx pubescent on outside ........................................................... 2
1. Fruit with a coriaceous or much-reduced wing; seed chamber in a central
or more often a lateral position; inflorescence glabrous or pubescent, a
raceme or a much-branched panicle; calyx glabrous or pubescent on
outside .................................................................................................... 3
2(1). Fruit stipitate, the stipe 8–25 mm long, the wing with a small outgrowth
distally; seeds 1–3, the testa smooth, pale brown or brown-pink; pedicel
382 F ABACEAE

(6–)7–13(–17) mm long; leaflets (7)9–11(13), apex emarginate, mucr-

onate, or briefly acuminate-mucronate (usually the mucro caducous)
.............................................................................................. P. acapulcensis
2. Fruit estipitate, the wing without small outgrowth distally; seed solitary,
the testa rugose, pale brown to dark brown; pedicel (2.5–)4–9(–12) mm
long; leaflets (3)5–9(11), apex short or long obtuse-acuminate ..... P. rohrii
3(1). Fruit stipitate, the stipe 2.5–6 mm long; inflorescence a much- and usu-
ally laxly branched axillary panicle; calyx glabrous on the outside; fruit
glabrous, the seed-bearing part variously thickened, usually distinctly
veined ....................................................................................... P. officinalis
3. Fruit estipitate; inflorescence a simple, axillary raceme, or if branched,
only 1–3 short and few-flowered branches near the base; calyx densely
brownish pubescent on the outside; fruit usually minutely brownish
tan-pubescent, or glabrescent, the seed-bearing part generally not dis-
tinctly veined ......................................................................................... 4
4(3). Fruit smooth to rugulose, the seed-bearing part slightly or rather evenly
thickened, gradually grading into the wing, glabrescent and usually
golden tan; inflorescence racemose, sometimes spindle-shaped; stems
generally hollow and ant-inhabited ...................................... P. amazonum
4. Fruit wrinkled or deeply rugose, the seed-bearing part rather abruptly
thickened from the small surrounding wing, usually yellowish to rusty
brown with a fine pubescence or else glabrescent; inflorescence race-
mose or with 1–3 branches at the base; stems solid, generally not ant-
inhabited ............................................................................ P. santalinoides

Pterocarpus acapulcensis Rose, Contr. Evergreen tree 5–15 m tall; branchlets

U.S. Natl. Herb. 5: 143. 1897. —Drago,
Sangre de drago, Sangredrago.
Pterocarpus podocarpus S.F. Blake, Contr.
U.S. Natl. Herb. 20: 524. 1924.
Pterocarpus vernalis Pittier, Arb.
Arbust. Venez. 6–8 [reprinted from
Bol. Minist. RR. EE. no. 8/9]: 100.
1927.
Deciduous tree 5–25(–30) m tall; fruits
persisting for several months. Deciduous,
semideciduous, and gallery forests, 50–500
m; northern Bolívar. Anzoátegui, Aragua,
Barinas, Carabobo, Cojedes, Distrito Fed-
eral, Lara, Miranda, Portuguesa, Táchira,
Zulia; Mexico, Central America, Colombia.
ŠFig. 329.
Pterocarpus amazonum (Benth.) Amshoff,
Meded. Bot. Mus. Herb. Rijks Univ.
Utrecht 52: 56. 1939. —Phellocarpus
amazonum Mart. ex. Benth., Comm. Le-
gum. Gen. 42. 1837.
Pterocarpus ulei Harms, Verh. Bot.
Vereins Prov. Brandenburg 48: 171.
1907.
thickened, hollow and ant-inhabited. Ripar-
ian and swamp forests, 50–300 m; Delta
Amacuro (Caño Jota-Sabuca near Caño
Mariusa), Bolívar (Isla Anacoco, Río Nichare
basin, Serranía de Imataca), Amazonas (Río
Atabapo). Apure; Colombia, Guyana, Ecua-
dor, Peru, Brazil, Bolivia.
Pterocarpus officinalis Jacq., Select.
Stirp. Amer. Hist. 283, t. 183, fig. 92.
1763. —Cacú, Drago, Iburu (Warao),
Lagunero, Mucanana, Mucunana, Mu-
cutena, Sangredrago, Sangrito.
Pterocarpus draco L., Sp. Pl. ed. 2, 1662.
1763.
Tree 5–30 m tall; lower trunk with sinu-
ous buttresses to 3 m tall and 5 m wide.
Swamp and flooded riparian forests, occa-
sionally nonflooded lowland forests, near sea
level to 300(–600) m; widespread in lower
Delta Amacuro, Bolívar (Altiplanicie de Nu-
ria, Serranía de Imataca, east of Túriba).
Anzoátegui, Monagas, Sucre; Mexico, Cen-
tral America, West Indies, Colombia, Tri-
nidad, Ecuador, Brazil. ŠFig. 328.

Pterocarpus officinalis is common along
water courses in Delta Amacuro state and is
easily recognizable by the characteristic but-
tress roots. The light wood is locally used in
artisanry to make carved wooden animals.
Pterocarpus rohrii Vahl, Symb. Bot. 2: 79.
1791. —Sangredrago, Sangrito alado.
Pterocarpus rupestris Pittier, Arb. Arbust.
Venez. 6–8 [reprinted from Bol. Minist.
RR. EE. no. 8/9]: 23. 1927.
Pterocarpus magnicarpa Schery, Field-
iana, Bot. 28: 261. 1952.
Pterocarpus rohrii var. rubiginosus
Schery, Fieldiana, Bot. 28: 261. 1952.
Tree 6–30 m tall; trunk narrowly but-
tressed. Semideciduous to evergreen lowland
forests, often along watercourses, 50–200 m;
Delta Amacuro (Río Toro area along Bolívar
border), Bolívar (middle Río Caura, Río
Paragua, Río Toro area), Amazonas (La
Esmeralda, Río Cunucunuma, around San
Carlos de Río Negro). Anzoátegui, Apure,
Barinas, Carabobo, Falcón, Lara, Miranda,
Yaracuy, Zulia; Mexico, Central America, Co-
lombia, Trinidad, Guyana, Suriname, French
Guiana, Ecuador, Peru, Brazil, Bolivia. ŠFig.
330.
Pterocarpus 383
Pterocarpus santalinoides L’Hér. ex DC.,
Prodr. 2: 419. 1825. —Cuai-eneru,
Guajenera, Para-para, Sangrito.
Pterocarpus amazonicus Huber, Bol. Mus.
Paraense Hist. Nat. 5: 402. 1909.
Pterocarpus grandis R.S. Cowan, Mem.
New York Bot. Gard. 10: 86. 1961.
Usually evergreen tree 5–15 m tall; trunk
buttressed. Swamp, riparian, and semide-
ciduous forests, near sea level to 300 m;
Delta Amacuro (widespread), Bolívar (Ser-
ranía de Imataca), Amazonas (lower Río Ci-
papo, Río Yatúa). Colombia, Trinidad, Guy-
ana, Suriname, French Guiana, Ecuador,
Peru, Brazil, Bolivia, Paraguay, Argentina,
western Africa.
The collections from Amazonas state (in-
cluding the type of Pterocarpus grandis) have
unusually large leaflets and fruits and are
questionably referred to this species.
Fig. 328.
Pterocarpus
officinalis
384 F ABACEAE

Fig. 329. Pterocarpus acapulcensis

Fig. 330. Pterocarpus rohrii

 Rhynchosia 385

54. RHYNCHOSIA Lour., Fl. Cochinch. 425, 460. 1790, nom. cons.
Dolicholus Medik., Vorles. Churpfälz. Phys.-Ökon. Ges. 2: 354. 1787, nom.
rejic.
Arcyphyllum Elliott, J. Acad. Nat. Sci. Philadelphia 1: 371. 1818.
by Renée H. Fortunato
Suffrutescent herbs, lianas, or subshrubs, erect, ascending, or twining to pros-
trate. Roots woody, turnip-shaped, spindle-shaped, or club-shaped. Stems terete to
angular, simple or branched. Leaves unifoliolate or pinnate-trifoliolate, subsessile to
long-petiolate; stipules free, persistent or deciduous; leaflets 3-veined, pubescent,
glabrous, glandular on both surfaces or only the lower one, glandular trichomes yel-
lowish, sometimes reddish brown when dried; stipels small, persistent (sometimes
early caducous) or lacking. Inflorescence racemose, axillary, simple or few-branched,
1–many-flowered; peduncles surpassing the main cauline leaves or not; bracts
small, persistent or deciduous; bracteoles lacking. Calyx campanulate, 5-lobed, the 2
lateral lobes subequal to 3 or more times the length of the tube, the 2 vexillar lobes
connate; corolla yellow, often striped or flushed with purple or reddish brown in the
standard veins; standard obovate to orbiculate, emarginate to rounded, sometimes
apiculate, biauriculate at the base, clawed, externally pubescent and glandular or
glabrous; wing petals narrowly oblong, uniauriculate, glandular-pubescent or gla-
brous at the apex, clawed; keel petals falcate, incurved and glandular-pubescent to
glabrous at apex, clawed. Stamens 10, diadelphous; anthers ellipsoid, dorsifixed,
uniform. Ovary sessile to subsessile, villous; ovules (1)2; style incurved, filiform, gla-
brous; stigma small. Legumes 2-seeded, ovoid, obovoid, ellipsoid, falcate, or dumb-
bell-shaped, deeply to shallowly constricted between seeds, usually compressed, pu-
bescent and glandular, dehiscent, beaked. Seeds subreniform to suborbicular,
brown, black, mottled, or red and black, lustrous; strophiole inconspicuous to promi-
nent, the hilum ovoid to linear; funicular attachment distal or central to subcentral
to the hilum.
Widely distributed in warm-temperate and tropical areas of both hemispheres,
but absent from Europe and northern and central Asia; ca. 190 species, 6 in Venezu-
ela, all of these in the flora area.

Key to the Species of Rhynchosia

1. Lateral calyx lobes > 2 times as long as the tube ..................................... 2

1. Lateral calyx lobes ± equal to the length of the tube ................................ 3
2(1). Standard pubescent and gland-dotted externally, 6–10 × 4–6 mm; stems
twining or creeping; legumes puberulous to glabrate, trichomes uni-
form in length .......................................................................... R. reticulata
2. Standard glabrous externally, 12–20 × 10–17 mm; stems erect to ascend-
ing; legumes viscid-hirsute and puberulous, trichomes of 2 lengths
........................................................................................... R. schomburgkii
3(1). Legumes deeply to shallowly constricted between the seeds (dumbbell-
shaped); seeds red and black; leaflets and standard with glands not
turning black when dried ...................................................................... 4
3. Legumes ellipsoid or falcate; seeds black, brown, or mottled; leaflets and
standard usually with glands turning black when dried ..................... 5
386 F ABACEAE

4(3). Legumes glabrous or puberulous, glabrescent and black with age; seeds
predominantly black, red only in the hilum area; stipels present;
pedicels 2–5 mm long ......................................................... R. melanocarpa
4. Legumes densely brown- or yellowish brown-puberulous, not glabrescent
with age; seeds with nearly equal areas of red and black almost equal;
stipels usually absent; pedicels 0.5–1.5 mm long .............. R. phaseoloides
5(3). Legumes ellipsoid; funicle attached distally to hilum; flowers 7.5–12 mm
long ................................................................................................. R. edulis
5. Legumes ± falcate; funicle attached centrally or nearly so to hilum; flow-
ers 3–6(–6.5) mm long ................................................................ R. minima

Rhynchosia edulis Griseb., Abh. Königl.
Ges. Wiss. Göttingen 19: 123. 1874.
—Eriosema edule (Griseb.) Burkhart,
Darwiniana 6: 261. 1943.
Rhynchosia melanosticta Griseb., Abh.
Königl. Ges. Wiss. Göttingen 19: 123.
1874.
Eriosema volubile Micheli, Mém. Soc.
Phys. Genève 28(7): 36. 1883.
Herbaceous or suffrutescent vine; stems
erect, suberect, prostrate, and twining; sti-
pules persistent; leaflets ovate, deltate, or
rhombic, glands on both surfaces; inflores-
cences elongate, usually equaling or exceed-
ing the leaves. Savannas, edges of forests in
open grassland, along riverbanks, 100–400
m; Bolívar (near Upata and Guasipati).
Aragua, Carabobo, Distrito Federal, Mérida,
Miranda, Portuguesa, Trujillo; southeastern
U.S.A., Central America, Colombia, Peru,
Brazil, Paraguay, Argentina.
Rhynchosia melanocarpa Grear, Mem.
New York Bot. Gard. 31(1): 43. 1978.
Rhynchosia phaseoloides var. erecta
Micheli, Mém. Soc. Phys. Genève 28(7):
33. 1883.
Twining, suffrutescent vine; stems few-
branched; stipules caducous; leaflets rhom-
bic, ovate, or deltate; stipels linear; inflores-
cences elongate, equaling or exceeding the
leaves. Thickets, margins of gallery forests
along rivers, disturbed areas, 400–500 m;
Bolívar (near Kamarata). Anzoátegui, Apure,
Guárico, Monagas; Peru, Brazil, Bolivia,
Paraguay, northeastern Argentina. ŠFig.
333.
Rhynchosia minima (L.) DC., Prodr. 2:
385. 1825. —Dolichos minimus L., Sp.
Pl. 726. 1753.
Glycine reflexa Nutt., Gen. N. Amer. Pl. 2:
115. 1818.
Suffrutescent to herbaceous vine; stems
twining or prostrate; stipules persistent;
leaflets rhombic-circular, ± rhombic, or
ovate; inflorescences elongate, equaling or
exceeding the leaves. Weed in disturbed and
cultivated land in different soil types, near
sea level; Delta Amacuro (Pedernales). Wide-
spread elsewhere in Venezuela; tropics and
subtropics worldwide. ŠFig. 334.
Rhynchosia phaseoloides (Sw.) DC.,
Prodr. 2: 385. 1825. —Glycine pha-
seoloides Sw., Prodr. 105. 1799. —Doli-
cholus phaseoloides (Sw.) DC. in Kuntze,
Revis. Gen. Pl. 3(2): 62. 1898.
Woody vine; stems angular, becoming ter-
ete; stipules caducous; racemes elongate,
usually equaling or exceeding the leaves,
branched. Margins of woods bordering sa-
vannas, clearings in evergreen forests, dis-
turbed areas, 50–1100 m; Delta Amacuro
(Río Orocoima), Bolívar (Altiplanicie de
Nuria, Río Grande, Río Suapure, west of
Santa Elena de Uairén). Distrito Federal,
Guárico, Miranda, Sucre, Yaracuy, Zulia;
southern Panama, Antilles, Colombia,
Guyana, Suriname, French Guiana, Peru,
Brazil, Bolivia. ŠFig. 331.
Rhynchosia reticulata (Sw.) DC., Prodr. 2:
385. 1825. —Glycine reticulata Sw.,
Prodr. 2: 105. 1788.
Dolicholus kuntzei Harms ex Kuntze,
Revis. Gen. Pl. 3(2): 61. 1898. —Rhyn-
chosia kuntzei Harms ex Kuntze, Revis.
Gen. Pl. 3(2): 61. 1898, as synonym.
—Rhynchosia reticulata var. kuntzei
(Harms ex Kuntze) Grear, Mem. New
York Bot. Gard. 31(1): 116. 1978.
 Rhynchosia 387

Fig. 331. Rhynchosia phaseoloides Fig. 332.

Rhynchosia
schomburgkii

Fig. 333. Rhynchosia melanocarpa Fig. 334. Rhynchosia minima

388 F ABACEAE

Suffrutescent herb or prostrate or twining Rhynchosia schomburgkii Benth. in

vine; stems 3-angled; stipules lanceolate to
widely ovate, base truncate or cordate, cadu-
cous, rarely some persistent; leaflets ovate,
rhombic, or round-ovate; racemes congested
to elongate, shorter than or exceeding the
leaves. Thickets, margin forests, along riv-
ers, grasslands, shrub savannas, 100–200 m;
Bolívar (El Cuchivero). Barinas, Carabobo,
Distrito Federal, Falcón, Monagas, Sucre,
Táchira, Yaracuy; Central America, West
Indies, Colombia, Guyana, Suriname,
French Guiana, Ecuador, Peru, Brazil,
Bolivia.
Mart., Fl. Bras. 15(1): 203. 1859.
Dolicholus pittieri Standl., Contr. U.S.
Natl. Herb. 17: 433. 1914.
Rhynchosia caliensis Harms, Notizbl. Bot.
Gard. Berlin-Dahlem 11: 783. 1933.
Subshrub to suffrutescent herb; stems
erect to twining at the tips; stipules cadu-
cous; leaflets discolorous; flowers 1–2.2 cm
long; calyx only exceeding the corolla by the
keel lobes. Sandy savannas, patches of for-
ests, along rivers, 200–1200 m; Bolívar
(widespread). Zulia; Colombia, Guyana,
northern Brazil. ŠFig. 332.

55. SESBANIA Scop., Intr. Hist. Nat. 308. 1777, nom. cons.
by Gerardo A. Aymard C.
Annual or perennial herbs, shrubs, or small trees. Leaves alternate with nu-
merous leaflets; leaflets entire, even-pinnate, obtuse at the apex, rounded at the
base, often glaucous, short-petiolate; stipules small, deciduous; stipels minute or
apparently absent. Inflorescences short and few-flowered axillary racemes; bracts
and bracteoles small, apparently deciduous. Flowers with the hypanthium broadly
campanulate, as broad as long or broader, the 5 teeth equal, often short, truncate to
triangular with the tip acute to acuminate; petals white, purplish yellow, red, or var-
iegated; standard longer than other petals, mostly orbicular or nearly so, reflexed,
short-clawed and usually appendaged at the base; wing petals free; keel petals
curved, joined below, with long claws. Stamens diadelphous, the vexillar stamen
free, geniculate near the base, the united stamens equal or 5 alternate ones some-
what longer. Ovary stipitate, linear; stigma small, capitate. Fruit linear, short-stipi-
tate, beaked, septate between the seeds within, the exocarp occasionally indented
between the seeds in some species, 2-valved, mostly dehiscent, occasionally indehis-
cent, the calyx often not persistent in fruit. Seeds many, oblong, slightly compressed,
smooth, brown.
Pantropics; ca. 50 species, 5 in Venezuela, 2 of these in the flora area.

Key to the Species of Sesbania

1. Pedicels 0.8–1 mm wide; calyx teeth 1–1.5 mm long; standard suborbicu-

lar, 1–1.5 cm long; wing petals oblong, 1.5 cm long; fruit 2–3.5 mm wide
...................................................................................................... S. emerus
1. Pedicels 1.5–2 mm wide; calyx teeth 3–4 mm long; standard ovate, 3 cm
long; wing petals oblong to obovate, to 2.5 cm long; fruit 4–6 mm wide
................................................................................................ S. exasperata

Sesbania emerus (Aubl.) Urb., Repert.
Spec. Nov. Regni Veg. 16: 149. 1919.
—Aeschynomene emerus Aubl., Hist. Pl.
Guiane 775. 1775. —Maguey.
Erect herb or shrub to 3 m tall; flowers
yellow with purple spots. Savannas, wet
places, near sea level to 200 m; Delta
Amacuro (Caño Araguao, Isla Cocuina,
Sacupana), Bolívar (El Manteco, Kilómetro
88, Represa Guri). Aragua, Falcón, Guárico,
 Sesbania 389

Fig. 335. Sesbania

exasperata
390 F ABACEAE

Lara, Miranda, Portuguesa, Zulia; Mexico,
Central America, West Indies, Colombia,
Guyana, Suriname, French Guiana, Ecua-
dor, Peru, Brazil.
Sesbania exasperata H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 534. 1823 [1824].
Erect herb or shrub to 3 m tall; flowers
yellow. Wet places, edges of rivers and
creeks, near sea level to 200 m; Delta
Amacuro (Caño Güiniquina), Amazonas (Río
Cataniapo). Apure, Barinas, Cojedes, Portu-
guesa, Zulia; Central America, West Indies,
Colombia, Guyana, Suriname, French Gui-
ana, Peru, Brazil, Bolivia, Paraguay, Argen-
tina. ŠFig. 335.

56. SOEMMERINGIA Mart., Soemmeringia Nov. Pl. Gen. 27. 1828.

by Nidia L. Cuello A.
Small, creeping herbs from a central tap root, sometimes prostrate or some-
what shrubby. Leaves alternate, odd-pinnate, sensitive; leaflets 7–9, bluish green,
oblanceolate, or obovate, shallowly emarginate at apex, glabrous; blades asymmetric
about the midvein, the secondary venation parallel and prominent on both surfaces,
extending to the leaflet margin and forming small teeth; stipules linear-lanceolate,
sharply acuminate, striate, papery, persistent; stipels absent. Flowers 1 or 2, axil-
lary; bracts small, subulate; bracteoles ovate to lanceolate, striate, persistent at ca-
lyx base. Calyx green, 2-lipped, upper lip 2-toothed, the teeth much longer than the
tube, the upper 2 broader and fused almost to their apex; corolla yellow; standard
subsessile, reticulate-venose, becoming brownish and scarious and persisting as a
delicately veined protective covering completely enclosing the green plicate fruit.
Stamens in 2 lateral bundles; anthers uniform. Ovary stipitate; ovules several; style
incurved. Fruit stipitate, joints plano-convex, reticulate.
Venezuela, Brazil, Bolivia; 1 species.

Soemmeringia semperflorens Mart., Paragua, lower Río Caroní). Anzoátegui,

Diss. de Soemmer 28. 1828. Apure, Cojedes, Guárico, Monagas, Portu-
Prostrate shrub or subprostrate herb; pet- guesa; Brazil (Amapá, Bahia, Ceará, Maran-
als light yellow. Open savannas, disturbed hão, Pará, Roraima), Bolivia. ŠFig. 336.
areas, 50–300 m; Bolívar (near Caicara, La

Fig. 336. Soemmeringia semperflorens

 Spirotropis 391

57. SPIROTROPIS Tul., Arch. Mus. Hist. Nat. 4: 113. 1844.

by Charles H. Stirton and Gerardo A. Aymard C.
Trees or lianas. Stipules spatulate-oblanceolate. Leaves odd-pinnate, 1–4-ju-
gate, opposite or subalternate, coriaceous, the upper surface glabrous, the lower sur-
face velvety, apex sharply acuminate. Inflorescence terminal, racemose-paniculate.
Flowers purple to purplish red. Calyx split into 1 or 2 parts at maturity, reflexed,
upper half 2-lipped, lower half 3-lipped; petals short-clawed; standard elliptic to or-
bicular, glabrous; wing petals equal to keel petals but shorter than standard. Fila-
ments free or scarcely united at their base. Ovary with 3–6 ovules, densely pubes-
cent; style filiform; stigma capitate. Fruit flattened, reticulate, hard, oblong, taper-
ing at both ends, apex acute, indehiscent.
Venezuela, Guyana, Suriname, French Guiana; 2 species, both in the flora
area.

Fig. 337. Spirotropis longifolia

392 F ABACEAE

Key to the Species of Spirotropis

1. Leaves and flowering axis finely puberulous, tawny or silvery; calyx split
into 2 halves, recurved, quite twisted; filaments fused near the base
.................................................................................................. S. longifolia
1. Leaves and flowering axis coarsely pubescent, ferruginous; calyx split
once into a broad, recurved, slightly twisted flap; filaments nearly free
.......................................................................................................... S. sp. A
Spirotropis longifolia (DC.) Baill., Hist. The Venezuelan collections could belong
Pl. 2: 364. 1870. —Swartzia longifolia to two undescribed species; more flowering
DC., Mém. Légum. 406. 1806. —Turä- material is needed to be certain.
yek (Arekuna).
Dipteryx phaeophylla Steyerm., Ann. Mis- Spirotropis sp. A
souri Bot. Gard. 71: 313. 1984. Small tree 8–10 m tall; petals purple. Gal-
Tree 4–15 m tall. Lowlands, montane for- lery forests, along river margins and islands,
ests, 100–1000 m; Bolívar (Río Carapo on 300–400 m; Bolívar (Río Canaracuni, Río
Cerro Guaquinima). Guyana, Suriname, Ichún). Endemic.
French Guiana. ŠFig. 337.

58. STYLOSANTHES Sw., Prodr. 108. 1788.

by Gerardo A. Aymard C.
Herbs, rarely subshrubs. Stems erect, ascending, or sprawling-procumbent,
diffusely branched, terete, finely striate, unarmed. Leaves alternate, pinnately 3-
foliolate; leaflets linear-lanceolate, elliptic, or oblong, apices obtuse to acute, occa-
sionally mucronate, the base attenuate, the costas prominent; petioles 1.5–15 mm
long; stipules amplexicaul, pubescent, 2-lobed, 3–11-veined; stipels absent. Inflores-
cences spicate, terminal or axillary, 1–several-flowered, each flower surrounded by a
series of bracts and bracteoles, the outermost bracts similar to stipules and giving
rise to 1–3 reduced leaflets, bracts subtended by a smaller outer bracteole, usually 3-
veined, ciliate along the margin. Flowers 5-merous. Calyx 4–15 mm long, tubular, 5-
lobed; petals yellow or yellow-orange with purple stripes. Stamens 10, monadelp-
hous; 5 versatile anthers alternating with 5 subbasifixed ones. Ovary subsessile,
placentation marginal; ovules 2(3); style long, filiform; stigma minute, terminal.
Fruit a loment, with 2 articles, distally fertile, proximally abortive or fertile, reticu-
late or muricate, beaked. Seeds ovate or lenticular, compressed, light brown to black,
lustrous.
U.S.A., Mexico, Neotropics (most diverse), introduced and naturalized in Ma-
laysia and Australia; ca. 50 species, 10 in Venezuela, 6 of these in the flora area.
The taxonomy of some of the taxa included here probably will not be resolved
until the genus has been revised.

Key to the Species of Stylosanthes

1. Leaflets linear to linear-lanceolate ............................................................ 2

1. Leaflets lanceolate to elliptic or oblong ..................................................... 3
2(1). Spike densely capitate; bracts hispid with golden trichomes; loment 1.5–
2.5 cm wide, with sessile glands, glabrous or with very short pubes-
cence (tuberculate) ................................................................. S. guianensis
2. Spike elongate; bracts ciliate, otherwise nearly glabrous; loment 0.8–
1.5 cm wide, minutely pubescent ........................................ S. angustifolia
 Stylosanthes 393

3(1). Branches densely pubescent to laxly pilose with short, viscid, sometimes
glandular black trichomes ........................................................... S. viscosa
3. Branches glabrous to puberulent or pilose, without glandular black tri-
chomes .................................................................................................... 4
4(3). Stems usually ascending or sometimes prostrate; beak of fruit 1–3.5 mm
long, strongly hooked ............................................................................. 5
4. Stems erect, rarely scandent and prostrate; beak of fruit 0.2–0.8 mm long,
erect or shortly hooked .......................................................................... 6
5(4). Leaflets glabrous to lax-pilose on both surfaces, apex obtuse to subacute;
flowers with 2 inner bracteoles .................................................. S. hamata
5. Leaflets short-bristly-ciliate to nearly glabrous on both surfaces, apex
acute; flowers with 1 inner bracteole ......................................... S. humilis
6(4). Spikes capitate to elongate, bracts 3–9-veined, glabrous to puberulent
with bristly golden trichomes, outer bracteole glabrous to pilose; stan-
dard suborbiculate ................................................................. S. guianensis
6. Spikes capituliform, bracts 11–17-veined, copiously soft-pubescent, outer
bracteole ciliate; standard obovate ............................................ S. capitata

Stylosanthes angustifolia Vogel, Linnaea

12: 63. 1838.
Erect herb, much-branched from the base;
flowers yellow. Wet savannas, 100–300 m;
Bolívar (Caicara, Cerro San Borja along
lower Río Suapure, near Ciudad Bolívar,
Maripa, lower Río Caroní, lower Río Par-
guaza); Apure, Aragua, Guárico, Sucre;
Guyana, Suriname, French Guiana, Brazil.

Stylosanthes capitata Vogel, Linnaea 12:

70. 1838.
Stems erect, branched, to 1 m tall; flowers yel-
low. Open places, savannas, 50–100 m; Bolívar
(near Ciudad Bolívar). Anzoátegui, Apure,
Guárico, Monagas, Zulia; Brazil, Bolivia.

Stylosanthes guianensis (Aubl.) Sw.,

Kongl. Vetensk. Acad. Nya Handl. 11:
296. 1789. —Trifolium guianense Aubl.,
Hist. Pl. Guiane 776, pl. 309. 1775.
Widespread in Neotropics; 7 varieties, 3
in Venezuela, 2 of these in the flora area.

Key to the Varieties of S. guianensis

1. Leaflets linear-lanceolate; inflorescences

usually terminal; spikes densely capi-
tate; loment with sessile glands (mi-
nutely tuberculate) or glabrous ..............
Fig. 338.
......................................... var. gracilis Stylosanthes
1. Leaflets elliptic to lanceolate; inflores- guianensis var.
cences terminal or sometimes axillary; gracilis
spikes loosely capitate; loment usually
glabrous or with very short pubescence
.................................... var. guianensis
394 F ABACEAE

S. guianensis var. gracilis (H.B.K.) Vogel,
Linnaea 12: 66. 1838. —Stylosanthes
gracilis H.B.K., Nov. Gen. Sp. (quarto
ed.) 6: 507, t. 596. 1823 [1824]. —Ca-
dillo.
Erect herb; flowers yellow. Savannas,
meadows, 100–1100 m; Bolívar (widespread),
Amazonas (Puerto Ayacucho, Sierra Parima).
Anzoátegui, Guárico, Monagas, Sucre, Tru-
jillo, Zulia; Panama, Colombia, Guyana,
Suriname, French Guiana, Peru, Brazil, Bo-
livia, Paraguay, Argentina. ŠFig. 338.
S. guianensis var. guianensis
Semierect to erect herb; flowers yellow.
Savannas, open places, 200–1100 m; Bolívar
(Ciudad Bolívar, Gran Sabana, lower Río
Caroní, Río Cuyuní, near Tumeremo), Ama-
zonas (near Puerto Ayacucho, Río Ocamo, Si-
erra Parima). Aragua, Barinas, Carabobo,
Cojedes, Distrito Federal, Mérida, Monagas,
Portuguesa, Sucre, Trujillo, Zulia; Mexico,
Central America, Trinidad, Guyana, Suri-
name, French Guiana, Peru, Brazil, Bolivia,
Paraguay.
Stylosanthes hamata (L.) Taub., Verh. Bot.
Vereins Prov. Brandenburg 32: 22. 1890.
—Hedysarum hamatum L., Syst. Nat.
ed. 10, 2: 1170. 1759.
Ascending herb, or sometimes prostrate,
often branched; flowers yellow. Savannas,
open places, 50–400 m; Bolívar (Ciudad Piar,
La Escalera, near Puerto Ordaz). Anzoá-
tegui, Aragua, Falcón, Guárico, Lara, Nueva
Esparta, Portuguesa, Sucre, Táchira, Yara-
cuy, Zulia; U.S.A. (Florida), West Indies,
Guatemala, Panama, Colombia, Brazil.
Stylosanthes humilis H.B.K., Nov. Gen.
Sp. (quarto ed.) 6: 506, t. 594. 1823
[1824]. —Cola de Zorro.
Herb, usually ascending, sometimes pros-
trate; flowers yellow. Open savannas with
Trachypogon, 50–200 m; Bolívar (Caicara,
near Ciudad Bolívar, Ciudad Piar, Maripa,
lower Río Caroní), Amazonas (near Puerto
Ayacucho). Apure, Barinas, Carabobo, Co-
jedes, Guárico, Mérida, Monagas, Nueva
Esparta, Portuguesa, Sucre, Trujillo, Zulia;
Mexico, Central America, West Indies, Co-
lombia, Brazil, Bolivia, introduced and natu-
ralized in Malaysia and Australia.
Stylosanthes viscosa (L.) Sw., Prod. 108.
1788. —Hedysarum hamatum var.
viscosum L., Pl. Jamaic. Pug. 20. 1759.
—Cola de zorro, Oregano canaotero,
Oregano montañero, Peludita.
Ascending and spreading or prostrate
herb; flowers yellow. Open savannas, 100–
900 m; Bolívar (Ciudad Bolívar, El Manteco,
El Pao, Gran Sabana, near Maripa, Río
Caroní, near Tumeremo, Upata). Anzoátegui,
Barinas, Falcón, Guárico, Monagas, Nueva
Esparta, Sucre, Trujillo, Zulia; U.S.A. (Texas,
introduced in Illinois), Mexico, Central
America, Antilles, Colombia, Guyana, Suri-
name, French Guiana, Peru, Brazil, Bolivia,
Paraguay, Argentina.

59. SWARTZIA Schreb., Gen. Pl. 2: 518. 1791, nom. cons.

Possira Aubl., Hist. Pl. Guiane 934. 1775.
Tounatea Aubl., Hist. Pl. Guiane 549. 1775.
Rittera Schreb., Gen. Pl. 1: 364. 1789.
Hoelzelia Neck., Elem. Bot. 3: 62. 1790.
Riveria H.B.K., Nov. Gen. Sp. (quarto ed.) 7: 266. 1825.
Fairchildia Britton & Rose, N. Amer. Fl. 23: 348. 1930.
Huertia Mutis ex Alva in Mutis, Diario de Observ. 1: 215. 1957.
by Nidia L. Cuello A. and Richard S. Cowan
Shrubs, trees, rarely lianas; stipules minute and needle-like to large and folia-
ceous. Leaves odd-pinnate, 1–many-foliolate; petiole and rachis winged to terete, of-
ten stipellate at insertion of leaflets. Inflorescence racemose to paniculate-racemose,
borne on trunk, branches, or branchlets, axillary, or supra-axillary, solitary to fas-
ciculate; bracts usually present at base of pedicels, bracteoles often present at base
of or on pedicels. Calyx entire in bud, opening in 2–5 regular or irregular segments;
 Swartzia 395

petal 1 or none, usually clawed. Stamens dimorphic (except for 2 species), larger ones
2–25, with larger filaments and anthers than the many smaller stamens. Gynoecium
1(–3)-pistillate; gynophore usually well developed; style usually present and either
terminal on ovary or lateral to its long axis, occasionally absent. Fruit 1–many-
seeded, oval, moniliform, or elliptic, thin- to thick-walled, coriaceous to woody. Seeds
generally ovoid with crustose or chartaceous testa, mostly arillate, the aril white,
yellow, or red.
Neotropics; ca. 200 species, ca. 55 species in Venezuela, 52 of these in the flora
area.

Key to the Species of Swartzia

1. Leaves with a single leaflet, appearing simple ......................................... 2

1. Leaves with 3 or more leaflets ................................................................. 10
2(1). Stipules conspicuously developed, at least 4 mm long, persistent ........... 3
2. Stipules not conspicuous or caducous ....................................................... 4
3(2). Lower leaflet surface and inflorescence pilose, primary veins of the leaf-
lets interconnected intramarginally; stipules foliaceous, lanceolate,
6.5–28 × 1.5–9 mm .................................................................... S. buntingii
3. Lower leaflet surface glabrous, primary veins of the leaflets not intercon-
nected in an intramarginal vein; stipules nonfoliaceous, 4–5 × 1 mm
................................................................................................... S. stipellata
4(2). Inflorescence axes and flower buds glabrous; blades of leaflets lanceolate
.......................................................................................................... S. sp. F
4. Inflorescence axes and flower buds minutely strigulose, malpighiaceous-
strigulose, sericeous, or densely pilose; blades of leaflets elliptic, oblong
elliptic, broadly elliptic, or ovate ........................................................... 5
5(4). Inflorescence axes and flower buds sparingly minutely strigulose to seri-
ceous, or malpighiaceous-strigulose, flower buds globose to elliptic in
outline .................................................................................................... 6
5. Inflorescence axes and flower buds densely pilose-velutinous, flower buds
oval in outline ........................................................................................ 8
6(5). Lower surface of leaflets golden-silky with appressed, contiguous and par-
allel trichomes at least along midrib ........................................ S. palustris
6. Lower surface of leaflets glabrous or sparingly micro-strigulose or glau-
cous ......................................................................................................... 7
7(6). Blades of leaflets coriaceous, the lower surface glaucous or yellowish, the
apex rounded and bluntly apiculate; calyx segment deciduous; style
1–2 mm long, not persistent in fruit .................................... S. vaupesiana
7. Blades of leaflets chartaceous, the lower surface not glaucous or yellow-
ish, the apex bluntly acute to acuminate; calyx segment persistent with
the fruits; style 4.5–6 mm long, persistent in fruit ................... S. conferta
8(5). Gynoecium densely villose marginally ......................................... S. maguirei
8. Gynoecium glabrous ................................................................................... 9
9(8). Leaflets narrowly oblong to elliptic-oblong, 8–18(–22) × 3.5–6(–9) cm,
margin of the leaflets usually narrowly revolute ................. S. floribunda
9. Leaflets broadly oblong (18–)24–34 × 9–14 cm, margin of leaflet plane
.......................................................................................................... S. sp. E
396 FABACEAE

10(1). Lateral leaflets alternate .......................................................... S. microcarpa

10. Lateral leaflets opposite ........................................................................... 11
11(10). Inflorescences axillary, simple racemes .................................................. 12
11. Inflorescences borne on trunk, branches, or branchlets, simple or panicled
racemes ................................................................................................ 21
12(11). Lateral leaflets ≥ 2 pairs .......................................................................... 13
12. Lateral leaflets 1 or 2 pairs ...................................................................... 16
13(12). Gynoecium and young fruits pubescent; stipules linear-lanceolate or folia-
ceous, 5–15 mm long ............................................................................ 14
13. Gynoecium and young fruits glabrous or partly sericeous; stipules trian-
gular or linear-attenuate, not foliaceous, < 5 mm .............................. 15
14(13). Leaflets 3 or 4 pairs, the blades 4–12 × 2.5–5.5 cm; base of leaflets rounded;
gynoecium 2-pistillate, stipules deciduous on old leaves and present on
new shoots, linear, pubescent, 5–10.5 mm long ......................... S. dipetala
14. Leaflets 2 or 3 pairs, the blades 10.5–21 × 4.5–9.5 cm; base of leaflets acute
to obtuse; gynoecium 1-pistillate; stipules persistent on old
leaves, foliaceous, 10–15 mm long .................................................. S. sp. C
15(13). Leaflets 2 or 3 pairs, often even-pinnate; petioles, leaf rachis, and inflor-
escense axes minutely strigulose to glabrescent; pedicels 12–23 mm
long, minutely strigulose; ovary laterally sericeous ............... S. parvifolia
15. Leaflets 4 or 5 pairs, odd-pinnate; petioles, leaf rachis and inflorescence
axes densely brownish-sericeous; pedicels stout, 8–12 mm long,
densely silky-pilose, clavately thickened upward; ovary glabrous
..................................................................................................... S. oedipus
16(12). Leaf rachis with a distinct narrow wing .................................. S. arborescens
16. Leaf rachis lacking a wing ....................................................................... 17
17(16). Inflorescence axes densely tomentose ..................................................... 18
17. Inflorescense axes glabrous or strigulose ................................................ 20
18(17). Lower surface of leaflets densely subappressed-pilose over entire surface
................................................................................................... S. roraimae
18. Lower surface of leaflets glabrescent, or strigose on midrib .................. 19
19(18). Flower buds 6–10 mm wide, densely brownish silky-pilosulous, pedicels
thick 2–4 mm wide, 12–25 mm long; lower surface of leaflets gland-
dotted, thinly strigose on midrib .............................................. S. aymardii
19. Flower buds 4–5 mm wide, golden-sericeous, pedicels slender < 1 mm
wide, 7–10 mm long; lower surface of leaflets glabrous over entire
surface, not gland-dotted .................................................... S. brachyrachis
20(17). Leaflets 1–3, oblong-elliptic, obtuse to rounded at base, lower surface of
leaflets glaucous or yellowish, secondary veins not prominent on upper
surface ................................................................................... S. vaupesiana
20. Leaflets always 3, obovate-elliptic, angustate at base, lower surface not
glaucous, secondary veins prominent on upper surface ................. S. sp. A
21(11). Style terminal, i.e., continuing long axis of ovary; fruit moniliform, rarely
elliptic-oblong ....................................................................................... 22
21. Style lateral, i.e., perpendicular to long axis of ovary or appearing so by
abrupt curvature of ovary apex; fruit variously shaped, but never
moniliform ............................................................................................ 39
22(21). Gynoecium densely pubescent or sericeous on all surfaces ................... 23
 Swartzia 397

22. Gynoecium sparingly strigulose, only partly pubescent or glabrous ...... 32

23(22). Rachis marginate to winged ..................................................................... 24
23. Rachis terete, canaliculate, or lacking stipels ......................................... 26
24(23). Leaflets rigid, bullulate by the impressed venation on the upper surface,
the veins strongly salient on the lower surface; ovary tomentose, gyno-
phore glabrous ..................................................................... S. pachyphylla
24. Leaflets chartaceous to thinly coriaceous, not at all bullulate; ovary seri-
ceous; the gynophore pubescent .......................................................... 25
25(24). Mature buds verruculose; petal surface obscured dorsally by the dense
pubescence; fruit ± oblong, ca. 4 cm wide, obliquely costate .... S. laxiflora
25. Mature buds costulate longitudinally or smooth; petal sericeous on the
back, at least on the veins; fruit moniliform, ca. 2 cm broad, ± verrucose
but not obliquely costate ....................................................... S. grandifolia
26(23). Young branchlets strongly sulcate, the stipules very broad, conspicuous
.............................................................................................................. 27
26. Young branchlets not sulcate, the stipules narrower and not conspicuous
.............................................................................................................. 28
27(26). Stipules acute at apex; rachis of leaves terete, stipellate at each pair of
leaflets, densely light brown to golden-tomentose ................... S. laxiflora
27. Stipules rounded at apex; rachis of leaves canaliculate on upper surface,
not stipellate at each pair of leaflets, densely dark brown-pannose
................................................................................................... S. panacoco
28(26). Lower surface of leaflets densely puberulent microscopically and buff-col-
ored ....................................................................................................... 29
28. Lower surface of leaflets glabrous or minutely and sparingly strigulose-
glabrescent ........................................................................................... 30
29(28). Lower surface of leaflets with 2 types of trichomes, peltate, short-stalked
ones and much longer, subulate, simple ones ........................ S. jenmannii
29. Lower surface of leaflets uniformly micro-puberulent, trichomes peltate,
short-stalked ...................................................................... S. schomburgkii
30(28). Rachis of leaves slightly canaliculate on upper surface; anther con-
nectives apiculate, at least on the larger stamens; fruit moniliform,
each section 1–2 × 1.5 cm .................................................. S. cardiosperma
30. Rachis of leaves terete, marginate or shortly winged; anther connectives
obtuse, not apiculate; fruit elliptic to oblong in outline, 4.5–9 × 2–
2.5 cm, if moniliform, each section 2.5–4.5 × 1.5–2 cm ...................... 31
31(30). Pedicels > 15 mm long; leaflets 1- or 2-jugate; filaments of larger stamens
17–22 mm long ............................................................................ S. pinnata
31. Pedicels mostly 7–10 mm long; leaflets 3- or 4-jugate; filaments of larger
stamens 5–9 mm long ............................................................ S. leptopetala
32(22). Lateral leaflets 2 or 3 pairs ...................................................................... 33
32. Lateral leaflets 4–8 pairs ......................................................................... 34
33(32). Leaflets obovate to obovate-elliptic; stipules nonfoliaceous .......... S. leiogyne
33. Leaflets elliptic; stipules foliaceous ......................................... S. macrocarpa
34(32). Young branchlets strongly ribbed; lateral leaflets 5–8 pairs .................. 35
34. Young branchlets not strongly ribbed; lateral leaflets 4 or 5 pairs ........ 36
35(34). Leaflets 6–8 pairs, blades narrowly lanceolate or linear lanceolate, 2.5–
6 × 0.8–1.8 cm; ovary laterally and ventrally densely pale sericeous, dor-
398 FABACEAE

sally glabrescent .......................................................................... S. cowanii

35. Leaflets 5 or 6 pairs, blades oval to elliptic to narrowly elliptic to oblong-
elliptic 4–18.5 × 2.5–8 cm; ovary glabrous over entire surface ..... S. sprucei
36(34). Leaflets glabrous on lower surface .......................................................... 37
36. Leaflets strigulose or strigose to pilose, not glabrescent on lower surface
.............................................................................................................. 38
37(36). Leaflets ± minutely warty; fruits woody, markedly white-lenticellate
................................................................................................. S. polyphylla
37. Leaflets not at all warty; fruits neither woody nor white-lenticellate
.................................................................................................. S. cuspidata
38(36). Blades of the lowermost pair of leaflets usually oval 7.7–10(–15) × 3.5–
6 cm, the blades of the other pairs elliptic to elliptic-oblong to oblong or
obovate-oblong, 13.5–23 × 4.5–9 cm; petal blade sinuate-dentate
.......................................................................................................... S. picta
38. Blade of all leaflets elliptic to obovate-elliptic, 3–9.5 × 1.5–2.5 cm; petal
blade not sinuate-dentate ......................................................... S. piarensis
39(21). Rachis of leaves distinctly winged ........................................................... 40
39. Rachis of leaves unwinged, terete, marginate or stipellate at each pair of
leaflets .................................................................................................. 43
40(39). Lateral leaflets 2 or 3 pairs ...................................................................... 41
40. Lateral leaflets 6–11 pairs ........................................................................ 42
41(40). Calyx densely pubescent on inner surfaces; apex of leaflets bluntly acumi-
nate; filaments of larger stamens glabrous or slightly strigulose
................................................................................................ S. guianensis
41. Calyx glabrous on inner surfaces; apex of leaflets abruptly acute to acumi-
nate; filaments white-villose .............................................. S. steyermarkii
42(40). Lateral leaflets 6 or 7 pairs; wing segment of rachis herbaceous, with a
deltate and sharply mucronate tip at the insertion of each pair of leaf-
let; valves of pods smooth and coriaceous .......................... S. alato-sericea
42. Lateral leaflets 10 or 11 pairs; wing segments coriaceous and revolute,
mostly with a rounded, slightly mucronate tip; valves of pods corru-
gated and woody .............................................................................. S. sp. B
43(39). Leaf rachis stipellate ................................................................................ 44
43. Leaf rachis lacking wings and stipels ..................................................... 46
44(43). Inner surfaces of calyx silky; filaments of larger stamens pubescent
....................................................................................................... S. sericea
44. Inner surfaces of calyx glabrous .............................................................. 45
45(44). Leaflets usually 2–4-jugate, elliptic to ovate ............................ S. tessmannii
45. Leaflets 6- or 7-jugate, narrowly oblong .............................. S. angustifoliola
46(43). Gynoecium glabrous ................................................................................. 47
46. Gynoecium pubescent .............................................................................. 50
47(46). Lateral leaflets 1 pair........................................................................... S. sp. D
47. Lateral leaflets 2 or more pairs ............................................................... 48
48(47). Leaflets 2 pairs, blades conspicuously reticulate-venose and elevated on
upper surface .......................................................................... S. wurdackii
48. Leaflets (2)3–5 pairs; veins of blades on upper surface plane or obscure or
slightly impressed ................................................................................ 49
49(48). Bracteoles lacking; blades of leaflets bluntly acute to rounded-obtuse at
apex ............................................................................................... S. pittieri
 Swartzia 399

49. Bracteoles present, foliaceous; blades of leaflets caudate-cuspidate at

apex, the cauda linear ................................................................ S. caudata
50(46). Principal lateral veins of leaflets joined submarginally into a ± conspicu-
ous vein; leaflets golden- or silvery-silky on lower surface ...... S. argentea
50. Principal lateral veins not forming a submarginal vein; leaflets not silky
on lower surface, strigose, strigulose, or glabrescent ......................... 51
51(50). Median leaflets 27 × 12 cm, strongly venose on both sides; petioles ca.
8 mm diameter at base ...................................................... S. benthamiana
51. Median leaflets smaller, venation less marked; petioles more slender
.............................................................................................................. 52
52(51). Mature buds warty; fruit with anastomosing, strongly salient, nearly
transverse ridges ............................................................... S. benthamiana
52. Mature buds not warty; fruit warty or smooth ....................................... 53
53(52). Buds oval, densely golden-silky, ca. 10 mm diameter; leaflets densely stri-
gulose on lower surface; inflorescence axes densely yellowish or golden
appressed-pubescent; valves of pods golden-sericeous in inmature pods,
papillose-verrucose to lenticellate in mature pods ........... S. cupavenensis
53. Buds globose or obnapiform, sparingly strigulose, 5–7 mm diameter; leaf-
lets usually glabrous; inflorescence axes thinly minutely silky-stri-
gulose or glabrescent, never yellowish or golden-pubescent, valves of
pods glabrous and smooth or glabrous and minutely warty but not ver-
rucose nor lenticellate in mature pods ................................................ 54
54(53). Ovary mostly glabrous except for few trichomes at base and along su-
tures; valves of mature pods smooth, at middle shallowly sulcate
lengthwise and the sutures developed into wings, 2 on the adaxial side
and 1 on the abaxial side ............................................................ S. triptera
54. Ovary densely pubescent in all surfaces; valves of mature pods minutely
warty, not sulcate lengthwise and the sutures unmodified ..... S. laevicarpa

Swartzia alato-sericea Barneby, Ann. Mis-
souri Bot. Gard. 78: 177. 1991.
Tree ca. 18 m tall. Lowland nonflooded for-
ests, 200–400 m; expected in Amazonas. Bra-
zil (Amazonas: known only from the south-
western base of Serra da Neblina).
Swartzia angustifoliola Schery, Fieldiana,
Bot. 28: 262. 1952. —Chamanare de
tierra firme.
Tree ca. 10 m tall. Nonflooded, riparian for-
ests, 100–200 m; Amazonas (Capibara, San
Carlos de Río Negro, Yavita to Maroa road).
Endemic. ŠFig. 339.
Swartzia arborescens (Aubl.) Pittier, J.
Wash. Acad. Sci. 11: 157. 1921. —Possira
arborescens Aubl., Hist. Pl. Guiane 934.
1775. —Guamo, Kajehai-detidi (Ye-
kwana), Sibara-koni.
Possira triphylla Sw., Prodr. 82. 1788.
Rittera dodecandra Vahl, Symb. Bot. 2:
60. 1791. —Swartzia dodecandra (Vahl)
Willd., Sp. Pl. 2: 1220. 1799.
Swartzia triphyllata Willd., Sp. Pl. 2:
1220. 1799.
Swartzia parviflora DC., Prod. 2: 423.
1825.
Swartzia bifida Steud., Flora 26: 757. 1843.
Swartzia rariflora Hoehne, Comiss. Lin-
has Telegr. Estratég. Mato Grosso Ama-
zonas, anexo 5, Bot. 12: 16. 1922.
Shrub or tree 3–10 m tall. Riparian for-
ests, secondary forests, 50–700 m; Bolívar
(mouth of Río Antavari, Río Nichare, Santa
María de Erebato), Amazonas (widespread).
Carabobo; Colombia, Guyana, Suriname,
French Guiana, Ecuador, Peru, Brazil, Bo-
livia. ŠFig. 342.
Swartzia argentea Spruce ex Benth. in
Mart., Fl. Bras. 15(2): 31. 1870.
Colombia, Venezuela, Brazil; 2 varieties,
1 in Venezuela.
400 FABACEAE
S. argentea var. argentea. —Chamanare
orillero, Jabón de raya, Macuca grande,
Palo de chamanare.
Tree 3–20 m tall; lower surface of leaves
silver-brown; petal yellow. Permanently
flooded forests along streams and rivers,
100–200 m; Amazonas (Caño San Antonio on
upper Río Orinoco, Río Cuao, Río Negro-Río
Casiquiare basin, Río Sipapo). Northwestern
Brazil (Amazonas: Rio Negro basin). ŠFig.
345.
The resin from the trunk is used to treat
sting-ray wounds.
Swartzia aymardii Barneby, Ann. Missouri
Bot. Gard. 78: 178. 1991.
Tree ca. 5–8 m tall; standard white.
Brushy savannas, gallery forests, 700–800
m; Bolívar (northern slope of Sierra Paca-
raima near source of Río Icabarú). Endemic.
Swartzia benthamiana Miq., Stirp. Suri-
nam. Select. 15. 1850 [1851].
Swartzia rosea Mart. ex Benth. in Mart.,
Fl. Bras. 15(2): 32. 1870.
Occasional in riparian forests, 100–400
m. Southeastern Colombia, Venezuela, Guy-
ana, Suriname, French Guiana, Peru, north-
western Brazil; 2 varieties, both in the flora
area.
Key to Varieties of S. benthamiana
1. Median leaflets smaller and venation less
marked than below; petioles < 8 mm di-
ameter at base; surface of fruits with
anastomosing, strongly salient, nearly
transverse ridges .. var. benthamiana
1. Median leaflets ca. 27 × 12 cm, strongly
venose on both sides, third degree
venules forming coarse reticulum; peti-
ole ca. 8 mm diameter at base; surface of
fruits golden-sericeous and verrucose,
without ridges .............. var. yatuensis
S. benthamiana var. benthamiana. —Cha-
manare.
Tree 6–15 m tall. Riparian forests, 50–300
m; Amazonas (Guayapo on lower Río Caura,
Río Autana, Río Guainía, Río Siapa, Río
Sipapo, Río Ventuari, San Carlos de Río Ne-
gro). Southeastern Colombia, Guyana, Suri-
name, French Guiana, northwestern Brazil.
S. benthamiana var. yatuensis R.S. Cowan,
Fl. Neotrop. Monogr. 1: 137. 1968.
Tree 10–30 m tall. Riparian forests, 100–
200 m; Amazonas (Río Baría, Río Mawari-
numa, Río Siapa, Río Yatúa). Endemic.
Swartzia brachyrachis Harms, Notizbl.
Königl. Bot. Gart. Berlin 6: 309. 1915.
Western Colombia, Venezuela, Guyana,
eastern Peru, Brazil; 5 varieties, 1 in Ven-
ezuela.
S. brachyrachis var. glabrata R.S. Cowan,
Fl. Neotrop. Monogr. 1: 192, figs. 39, 4a,
4b. 1968.
Tree or shrub 3–15 m tall; fruit orange.
Riparian forests, sometimes in dry secondary
forests, forested slopes, 50–700 m; Bolívar
(El Abismo, Río Samay). Guyana.
Swartzia buntingii R.S. Cowan, Fl. Neotrop.
Monogr. 1: 204. 1968.
Tree 8–20 m tall; flowers yellowish; fruits
green to orange. Riparian forests, 100–200
m; Amazonas (Río Cunucunuma, Río Mawa-
rinuma, Río Paciba, Río Yatúa). Endemic.
ŠFig. 341.
Swartzia cardiosperma Spruce ex Benth. in
Mart., Fl. Bras. 15(2): 33. 1870. —Cha-
manare, Guamo, Macuca montañero, Palo
de morrocoy.
Tree 6–20 m tall, cauliflorous; flowers yel-
low. Flooded and nonflooded rain forests,
100–300 m; Amazonas (Caño San Miguel,
Cerro Huachamacari, La Esmeralda, Río
Baría, Río Casiquiare, lower Río Guainía,
Río Mawarinuma, Río Negro, upper Río
Orinoco, Yavita to Maroa road). Southeast-
ern Colombia, eastern Peru, northwestern
Brazil.
Swartzia caudata R.S. Cowan, Fl. Neotrop.
Monogr. 1: 122. 1968. —Mudujai, Shi-ña-
teu.
Tree 4–25 m tall. Forests along rivers,
100–400 m; Amazonas (near Cerro Duida,
near Cerro Huachamacari, Ocamo, Río
Cunucunuma, upper Río Orinoco, Río
Padamo, Río Ventuari). Endemic.
Swartzia conferta Spruce ex Benth., Fl.
Bras. 15(2): 20. 1870.

Southeastern Colombia, Venezuela, north-
western Guyana, northwestern Brazil; 2 vari-
eties, 1 in Venezuela.
S. conferta var. conferta
Shrub or tree 1.5–15 m tall; flowers white;
fruit orange. Riparian and secondary forests,
50–700 m; Bolívar (Río Cuyuní), Amazonas
(Cerro Duida, Cerro Huachamacari, Río
Casiquiare-Río Negro basin, Río Cunucu-
numa). Southeastern Colombia, northwest-
ern Guyana, northwestern Brazil. ŠFig. 340.
Swartzia cowanii Steyerm., Brittonia 33:
33. 1981.
Tree 4–15 m tall, cauliflorous; flowers yel-
low. Near rivers on tepui summits, 700–800
m; Bolívar (Cerro Guaiquinima). Endemic.
ŠFig. 344.
Swartzia cupavenensis R.S. Cowan, Fl.
Neotrop. Monogr. 1: 139. 1968. —Dudu,
Guayaparu, Guayapu.
Tree 6–15 m tall; petal cream-white. For-
ests, 100–300 m; Amazonas (Caño Caname,
near the mouth of Caño Cupaven, Caño
Soromoni near La Esmeralda, Caño Yagua,
Isla Cangrejo near Cerro Duida, Río Cuao,
Río Guayapo, Río Orinoco, Río Pasimoni, Río
Sipapo, Río Ventuari, Río Yatúa, near San
Carlos Río Negro). Endemic.
Swartzia cuspidata Spruce ex Benth. in
Mart., Fl. Bras. 15(2): 36. 1870. —Cha-
manare.
Shrub or tree 3–18 m tall, cauliflorous;
flowers white. Riparian, roadside, and sec-
ondary forests, 100–700 m; Amazonas (Río
Casiquiare, Río Negro basin, Sierra de la
Neblina, Yavita to Maroa road). Eastern
Peru, Amazonian Brazil.
Swartzia dipetala Willd. ex J. Vogel,
Linnaea 11: 173. 1837. —Miriyon
(Panare), Panillo, Sowi (Yekwana).
Swartzia dicarpa Moric. ex Meisn., Pl.
Vasc. Gen., Commentarius 68. 1837.
Swartzia microstylis Benth., J. Bot.
(Hooker) 2: 89. 1840.
Tree 5–30 m tall; fruit orange; petals yel-
low; flowers fragrant. Riparian forests, semi-
deciduous forests, granitic slopes, savannas,
50–200 m; Bolívar (lower Río Caura, Río
Maniapure, Río Parguaza, lower Río Sua-
Swartzia 401
pure), Amazonas (near Galipero, Isla Ratón,
near Samariapo, San Juan de Manapiare).
Apure; Guyana, Brazil.
Swartzia floribunda Spruce ex Benth. in
Mart., Fl. Bras. 15(2): 21. 1870. —Anón,
Guaquito, Palo de golondrona.
Shrub or small tree 1.5–10 m tall; flowers
white. Lowland forests, bushy savannas, sec-
ondary forests, 50–200 m; Amazonas (Caño
Caname, Maroa, Río Atabapo, Río Atacavi,
Río Casiquiare, Río Guainía, Río Negro, Río
Temi, Yavita). Endemic.
Swartzia grandifolia Bong. ex Benth., J.
Bot. (Hooker) 2: 85. 1840. —Akayu-
wajunai, Lawadema-eje-de-deu, Yawa-
demo (Yekwana).
Tree 6–18 m tall, cauliflorous; flowers
creamy-white to yellow with purple veins. Low-
land forests along rivers on clay, sandy clay, or
rocky soils above level of seasonal flooding,
100–500 m; Bolívar (Río Caura, Río Suapure),
Amazonas (Río Casiquiare, Río Cunu-
cunuma, upper Río Orinoco, Río Ventuari).
Guyana, Suriname, French Guiana, north-
central and northwestern Brazil. ŠFig. 346.
Swartzia guianensis (Aubl.) Urb., Symb.
Antill. 5: 365. 1908. —Tounatea guia-
nensis Aubl., Hist. Pl. Guiane 550. 1775.
—Dawadema, Majako shodo (Yekwana).
Swartzia alata Willd., Sp. Pl. 2: 1220.
1799.
Tree 6–15 m tall. Understory of forests on
sandy or lateritic clay soil, riparian forests,
forested slopes, 100–400 m; Amazonas (Cer-
ro Huachamacari, Río Cunucunuma). Guy-
ana, Suriname, French Guiana.
The cortex of Swartzia guianensis is used
by Yekwana Amerindians to treat skin infec-
tions.
Swartzia jenmanii Sandwith, Bull. Misc. In-
form. Kew 1934: 361. 1934.
Large tree. Forested slopes, 800–1100 m;
Bolívar (Uei-tepui). North-central Guyana.
Swartzia laevicarpa Amshoff, Meded. Bot.
Mus. Herb. Rijks Univ. Utrecht 52: 37.
1939.
Tree 4–30 m tall; fruit gray-green. Wide-
spread in lowland, high forests along creeks
402 FABACEAE
and rivers, usually on sandy soils but also on
sandy clay, mostly in nonflooded locations,
also in scrub forest on summit of tepui, for-
ested slopes of tepui with to 20 m trees, 100–
1000 m; Bolívar (Amaruay-tepui), Amazonas
(Isla Maracá in Río Negro, Río Mawarinuma).
Southeastern Colombia, Guyana, Suriname,
northwestern Brazil.
Swartzia laxiflora Bong. ex Benth., J. Bot.
(Hooker) 2: 86. 1840. —Tounatea laxi-
flora (Bong. ex Benth.) Taub., Bot. Cen-
tralbl. 47: 391. 1891. —Tounatea laxi-
flora (Bong. ex Benth.) Kuntze, Revis.
Gen. Pl. 1: 211. 1891. —Chamanare.
Swartzia polycarpa Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 126. 1922.
Tree 4–15 m tall; flowers orange. Frequent
in nonflooded riparian lowland forests. 50–
100 m; Amazonas (Río Cuao, Río Guayapo,
Río Sipapo). Brazil (Amazonas).
Swartzia leiogyne (Sandwith) R.S. Cowan,
Fl. Neotrop. Monogr. 1: 46. 1968.
—Swartzia grandifolia var. leiogyne
Sandwith, Bull. Misc. Inform. Kew 1937:
103. 1937.
Tree 13–22 m tall. Lowland to lower mon-
tane forests, 50–400 m; expected to be found
in Amazonas. Guyana, Brazil (Roraima).
Swartzia leptopetala Benth., J. Bot.
(Hooker) 2: 87. 1840. —Carrasposo, Cha-
manare, Congrio, Yawademo de rebalse.
Swartzia discolor Poepp., Nov. Gen. Sp.
Pl. 3: 62. 1845.
Swartzia fugax Spruce ex Benth. in Mart.,
Fl. Bras. 15(2): 30. 1870.
Swartzia melanoxylon Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 123. 1922.
Swartzia rotundata R.S. Cowan, Mem.
New York Bot. Gard. 8: 116. 1953.
Tree 5–30 m tall; flowers yellow. Frequent
in riparian forests, 50–300 m; Bolívar (Río
Asa, Río Caroní, Río Caura, Río Erebato),
Amazonas (Cerro Yutajé, Maroa, Río Casi-
quiare, Río Cataniapo, Río Coro Coro, Río
Guainía, Río Matacuni, Río Ocamo, Río Ori-
noco). Anzoátegui, Apure, Miranda, Mona-
gas, Yaracuy; southeastern Colombia, Brazil.
ŠFig. 347.
Swartzia macrocarpa Spruce ex Benth. in
Mart., Fl. Bras. 15(2): 38. 1870.
Tree to 5 m tall. Riparian, seasonally or
permanently flooded forests, 100–200 m; ex-
pected to be found in Amazonas. Northwest-
ern Brazil (Amazonas: Rio Negro Basin).
ŠFig. 350.
Swartzia maguirei R.S. Cowan, Mem. New
York Bot. Gard. 8: 115. 1953. —Guaquito.
Tree 3–10 m tall; fruit orange. Evergreen
lowland forests, 100–300 m; Amazonas (Cer-
ro Sipapo, Culebra, El Gavilán near Puerto
Ayacucho, Río Cunucunuma, Río Coromoto,
Río Yureba, near San Carlos Río Negro). Bra-
zil (Amazonas).
Swartzia microcarpa Spruce ex Benth., Fl.
Bras. 15(2): 35. 1870. —Chamanare
negro.
Tree to 25 m tall. Flooded plains, 50–200
m; Amazonas (Río Guayapo, Río Pasimoni,
Río Sipapo). Colombia, Brazil.
Swartzia oedipus Barneby, Ann. Missouri
Bot. Gard. 78: 179, fig. 1. 1991.
Tree to 20 m tall; trunk to 20 cm diameter.
Nonflooded forests, 100–500 m; Amazonas
(Río Negro near Piedra Cocuy). Brazil.
Swartzia pachyphylla Harms, Notizbl.
Königl. Bot. Gart. Berlin 6: 310. 1915.
Tree 10–15 m tall. Gallery forests, ca.
1400 m; Bolívar (Roraima-tepui). Endemic.
Swartzia palustris Barneby, Ann. Missouri
Bot. Gard., 78: 179. 1991. —Wa-hú.
Slender tree 3–25 m tall, when crowded
sometimes producing long and lithe branches.
Swamp forests, rain forests on low hills, ri-
parian forests along black-water streams,
100–400 m; Amazonas (upper Río Baría, Río
Cunucunuma, southwestern base of Sierra de
la Neblina). Endemic.
Swartzia panacoco (Aubl.) R.S. Cowan, Fl.
Neotrop. Monogr. 1: 32. 1968. —Robinia
panacoco Aubl., Hist. Pl. Guiane 768.
1775.
Robinia tomentosa Willd., Sp. Pl. 3(2):
1134. 1802. —Swartzia tomentosa (Willd.)
DC., Prod. 2: 423. 1825.
Swartzia similis Benoist, Bull. Mus. Hist.
Nat. (Paris) 25: 297. 1919.
Tree. Upland and highland forests. Ven-
ezuela, Guyana, Suriname, French Guiana,

Brazil; 8 varieties, 2 in Venezuela, both in the
flora area.
Key to the Varieties of S. panacoco
1. Leaflets coriaceous, strigulose on lower
surface .......................... var. cardonae
1. Leaflets rigid, tomentose on lower surface
..................................... var. tepuiensis
S. panacoco var. cardonae (R.S. Cowan)
R.S. Cowan, Fl. Neotrop. Monogr. 1: 34.
1968. —Swartzia tomentosa var. car-
donae R.S. Cowan, Bol. Soc. Venez. Ci.
Nat. 13: 99. 1954. —Aka-yek, Chama-
nare, Guamo terciopelo, Kajari, Leche
cochino, Okoi-yek (Pemón), Tasajo,
Terciopelo.
Tree 10–30 m tall. Primary, riparian for-
ests on sandy nonflooded soils, 400–1200 m;
Bolívar (Río Canaracuni, Río Caura, Río
Erebato, Río Icabarú, upper Río Paragua,
Río Tonoró, Río Venamo), Amazonas (Río
Cunucunuma). Guyana, Brazil (Amazonas:
near junction of Rio Amazon and Rio Negro).
S. panacoco var. tepuiensis (Schery) R.S.
Cowan, Fl. Neotrop. Monogr. 1: 37. 1968.
—Swartzia tepuiensis Schery, Fieldiana,
Bot. 28: 265. 1952. —Ve-yek.
Tree 10–20 m tall. Slopes of sandstone
tepuis, dwarf open forests, 1600–2000 m;
Bolívar (Amaruay-tepui, Carrao-tepui, Gran
Sabana, Río Aícha near Los Caribes, Río Oris
in Río Paragua basin). Endemic. ŠFig. 352.
Swartzia parvifolia Schery, Fieldiana, Bot.
28: 263. 1952.
Tree 10–25 m tall. Summits of sandstone
tepuis, 700–1700 m; Bolívar (El Paují, near
Kavanayén, Macizo del Chimantá), Ama-
zonas (base of Cerro Cuao, Río Cuao). En-
demic. ŠFig. 349.
Swartzia piarensis R.S. Cowan, Fl. Neo-
trop. Monogr. 1: 45. 1968. —Uoki-yek.
Tree 8 m tall. Riparian forests, 500–800
m; Bolívar (Cerro Bolívar, Río Caroní near
mouth of Río Icabarú). Endemic.
Swartzia picta Spruce ex Benth. in Mart.,
Fl. Bras. 15(2): 25. 1870.
Tree 8–13 m tall. Riparian forests, for-
ested slopes of tepuis, 100–900 m. Venezuela,
Brazil; 2 varieties, both in the flora area.
Swartzia 403
Key to the Varieties of S. picta
1. Leaflets strigulose on lower surface, at-
tenuate basally, the apex shortly and
sharply acute; pedicels cylindric, 20–35
mm long; bracteoles arising below
pedicel apex ............. var. bolivarensis
1. Leaflets usually pilose on lower surface,
obtuse basally, the apex obtuse or shortly
and bluntly acute; pedicels clavate, 12–
24 mm long; bracteoles apical on pedicels
or on base of calyx ................ var. picta
S. picta var. bolivarensis R.S. Cowan, Fl.
Neotrop. Monogr. 1: 45. 1968.
400–900 m; Bolívar (quebrada Los Bra-
sileros near Icabarú, Río Chaberu, Río Sa-
may). North-central Brazil (Amazonas: near
Río Branco). ŠFig. 353.
S. picta var. picta. —Chamanare.
100–200 m; Bolivar (Río Acanán), Ama-
zonas (Río Negro, Río Guainía). Northwest-
ern Brazil. ŠFig. 354.
Swartzia pinnata (Vahl) Willd., Sp. Pl. 2:
1220. 1799. —Rittera pinnata Vahl,
Eclog. Amer. 2: 38. 1798.
Tree 7–15 m tall. Dry forests, 50–200 m;
Amazonas (Gavilán near Puerto Ayacucho,
Río Orinoco, Río Sipapo). Coastal Venezuela;
Trinidad.
Swartzia pittieri Schery, Fieldiana, Bot. 28:
263. 1952.
Tree 7–10 m tall. Granitic, rocky areas,
50–200 m; Bolívar (50 km southwest of
Caicara in Serranía La Encaramada), Ama-
zonas (Caño Chimoni tributary of Río Casi-
quiare, near Puerto Ayacucho, Río Orinoco).
Anzoátegui, Barinas, Guárico, Portuguesa;
West-central and southwestern Venezuela.
ŠFig. 351.
Swartzia polyphylla DC, Prod. 2: 424. 1825.
—Cajario, Candilón, Canjilón de agua,
Dau bagibagi (Waroa), Guaraba, Palo de
raya.
Swartzia acuminata Willd. ex J. Vogel,
Linnaea 11: 173. 1837. —Tounatea
acuminata (Willd. ex J. Vogel) Taub., Bot.
Centralbl. 47: 390. 1891.
Tounatea acuminata var. puberula Taub.,
Flora 75: 81. 1892. —Swartzia acu-
404 FABACEAE
minata var. puberula (Taub.) Glaz.,
Bull. Soc. Bot. France 53. Mem. 36:
155. 1906.
Swartzia acuminata var. tridynamia
Huber, Bol. Mus. Paraense Hist. Nat.
Ethnogr. 2: 506. 1898.
Swartzia platygyne Ducke, Arch. Jard.
Bot. Rio de Janeiro 3: 127. 1922.
Swartzia opacifolia J.F. Macbr., Field
Mus. Nat. Hist., Bot. Ser. 13(3.1): 226.
1943.
Swartzia urubuensis Ducke, Bol. Técn.
Inst. Agron. N. 2: 21 1944.
Tree 3–40 m tall; flowers white. Evergreen
forests, permanently flooded areas along riv-
ers, seasonally flooded river margins, near
sea level to 800 m; Delta Amacuro (Caño
Araguabisi between Caño Araguao and Caño
Güiniquina, Caño Güiniquina, Caño Joba-
Suburu east of Caño Sacupana, Caño Jobure,
near Curiapo, Misión San Francisco de
Guayo), Amazonas (Río Casiquiare, Río
Cuao, Río Mawarinuma, Río Negro, base of
Sierra de la Neblina). Colombia, Guyana,
French Guiana, eastern Peru, Amazonian
Brazil. ŠFig. 343.
The wood of Swartzia polyphylla is used for
paddles.
Swartzia roraimae Sandwith, Bull. Misc.
Inform. Kew 1934: 362. 1934.
Tree of unknown stature. Forested slope of
tepui, 1500–1600 m; Bolívar (southwestern
face of Roraima-tepui). Endemic.
Swartzia schomburgkii Benth. in Mart., Fl.
Bras. 15(2): 38. 1870. —Dudu, Kana-
guarajodi (Yekwana), Níspero, Oliverio,
Para-cutaco, Paretaura, Pore-taurai, Pre-
taurayese (Pemón), Sánamo, Shodocona
(Yanomami), Wvaraba (Yeral).
Swartzia schomburgkii var. guayanensis
R.S. Cowan, Fl. Neotrop. Monogr. 1: 90.
1968.
Tree 10–30 m tall; flowers white. Riparian
forests, 50–800 m; Bolívar (Cerro Camarón,
El Cácaro on upper Río Caura, between El
Dorado and Kilómetro 88, Río Erebato, Río
Ichún, Río Karún, Río Nichare, Río Para-
gua), Amazonas (Caño Ucata southeast of
Síquita, Río Atabapo, Río Negro-Río Cunu-
cunuma, upper Río Orinoco, Río Siapa near
base of Cerro Aracamuni, Río Ugueto, base of
Sierra de la Neblina). Southeastern Colom-
bia, Guyana, Suriname, northwestern Brazil.
Swartzia schomburgkii is treated here as
including var. guayanensis since the differences
in fruiting material are not consistent. Also,
some specimens from the southern part of
Amazonas state have a pubescent gynoecium,
which means that the var. guayanensis is not
restricted to Bolívar state as was thought.
The green wood of Swartzia schomburgkii
is cut into fine pieces and used to start fires.
Swartzia sericea J. Vogel, Linnaea 11: 176.
1837.
Southeastern Colombia, Venezuela, French
Guiana, Brazil (Amazonas); 2 varieties, 1 in
Venezuela.
S. sericea var. sericea. —Chamanare, Chi-
manare, Cupana, Cupana de chamanare,
Maca, Macuca, Tucana guarana.
Tree 4–15 m tall; fruits orange, with thick
white aril along seed. Riparian forests, 100–
600 m; Bolívar (El Dorado to Santa Elena),
Amazonas (Caño Cupueni, Pimichín, Río
Atabapo, Río Atacavi, Río Baría, Río Guainía,
Río Negro-Río Casiquiare, Río Siapa, Río
Sipapo, Río Temi, Río Ventuari, Río Yatúa,
San Juan). Southeastern Colombia, French
Guiana, Brazil (Amazonas: Rio Negro basin).
ŠFig. 355.
The seeds of this species are used for fish
bait.
Swartzia sprucei Benth. in Mart., Fl. Bras.
15(2): 37. 1870.
Tree 3–30 m tall. Guyana, Venezuela; 2
varieties, both in the flora area.
Key to the Varieties S. sprucei
1. Filaments of larger stamens sparingly vil-
lose; leaflets strigose but glabrescent
and not tessellate on lower surface; leaf
rachis distinctly stipellate at each pair
of leaflets .......................... var. sprucei
1. Filaments of larger stamens glabrous;
sparingly strigulose and usually minutely
tessellate on lower surface; leaf rachis
not at all stipellate ...... var. tessellata
S. sprucei var. sprucei
Tree ca. 10 m tall, cauliflorous. Riparian
forests, 50–300 m; Bolívar (Río Caura near
Salto Pará), Amazonas (Río Pasimoni). En-
demic.

S. sprucei var. tessellata R.S. Cowan, Fl.
Neotrop. Monogr. 1: 40. 1968.
Lower slope of ironstone hill, 100–500 m;
Bolívar (road between El Dorado and Kiló-
metro 88). Guyana.
Swartzia steyermarkii R.S. Cowan, Fl.
Neotrop. Monogr. 1: 160. 1968.
Tree 4–10 m tall. Lowland to montane for-
est, 100–1000 m; Bolívar (Río Ayaiche near
base of Sierra de Lema, Río Uroi). Endemic.
ŠFig. 358.
Swartzia stipellata R.S. Cowan, Fl. Neotrop.
Monogr. 1: 211. 1968.
Tree 5–7 m tall. Riparian forests, 100–200
m; Amazonas (Río Puruname). Brazil (Ama-
zon basin).
Swartzia tessmannii Harms, Notizbl. Bot.
Gart. Berlin-Dahlem 9: 971. 1926.
—Okiyeu (Pemón), Wadi (Yekwana).
Tree 8–24 m tall; flowers yellow; fruit or-
ange. Riparian forests, 200–500 m; Bolívar
(Cerro Cimarrón on upper Río Caura, Cerro
Sarisariñama, El Cácaro between Río Caura
and Río Paragua, El Paují, upper Río Caura,
Río Kurutú on Río Paragua, Río Uenan-
Ikabarú), Amazonas (upper Río Cunucu-
numa). Amazonian Peru and Brazil. ŠFig.
356.
Swartzia triptera Barneby, Ann. Missouri
Bot. Gard. 78: 181, fig. 2. 1991.
Tree 4–12 m tall; petal white; fruit green,
with sticky red exudate. Lowland nonflooded
forests, riparian forests, 100–200 m; Amazo-
nas (Cerro Marahuaka, Río Mawarinuma).
Endemic.
Swartzia vaupesiana R.S. Cowan, Fl.
Neotrop. Monogr. 1: 126. 1968. —Hoja del
sol.
Swartzia dolichopoda R.S. Cowan, Fl.
Neotrop. Monogr. 1: 130. 1968.
Swartzia vaupesiana var. glauca R.S.
Cowan, Fl. Neotrop. Monogr. 1: 127.
1968.
Tree 6–15 m tall; petals orange-yellow. Ri-
parian forests, 100–300 m; Amazonas (Caño
Cabeza Manteco tributary of Río Autana,
Caño Cupueini east of San Fernando de
Atabapo, Río Autana, upper Río Orinoco, Río
Sipapo, Río Yatúa, Sierra de la Neblina).
Southeastern Colombia. ŠFig. 357.
Swartzia 405
Swartzia vaupesiana is treated here as in-
cluding S. vaupesiana var. glauca and S.
dolichopoda, because of the great resem-
blance of all parts observed in the type speci-
mens of these three taxa. Variation in num-
ber of leaflets (1 or 3) seems to be common in
S. vaupesiana.
Swartzia wurdackii R.S. Cowan, Fl. Neo-
trop. Monogr. 1: 148. 1968.
Tree 15 m tall. Forests at base of escarp-
ments, 1300–1400 m; Bolívar (Macizo del
Chimantá [base of Amurí-tepui]). Endemic.
Swartzia sp. A. —Macho.
Tree 12–22 m tall, with reddish exudate;
fruits yellow to orange. Lower montane for-
ests, 200–300 m; Delta Amacuro (Río Grande).
Guyana, Suriname. ŠFig. 348.
This taxon is based on Blanco 164 (VEN), Za-
bala 126 (VEN), and Marcano-Berti 232 (MO).
Swartzia sp. B. —Chamanare.
Tree 15–20 m tall; fruits orange. Non-
flooded lower montane forests, 200–300 m;
Amazonas (Río Yatúa).
This taxon is based on Velazco 1420 (MO,
PORT).
Swartzia sp. C
Small tree; fruit orange, seed black with
white aril. Low open forest on granitic outcrop,
ca. 600 m; Amazonas (Cerro Aracamuni).
This taxon is based on Liesner & Carnevali
22271 (MO).
Swartzia sp. D
Tree 10 m tall; fruit bright orange, seed
black with white aril. Riparian forest, ca. 300
m; Amazonas (Río Mawarinuma).
This taxon is based on Davidse & Miller
27210 (MO).
Swartzia sp. E
Vining tree; flowers white. Nonflooded for-
ests, 500–600 m; Amazonas (Caño Colorado
and Serranía Batata in Cuao-Sipapo mas-
sif).
This taxon is based on Sanoja 3200 and
3371 (MO, PORT).
Swartzia sp. F
Small tree. Lowland riparian forest, 50–
100 m; Amazonas (Caño Monomi in Río
Casiquiare basin).
This taxon is based on Vareschi 7793 (VEN).
406 F ABACEAE

Fig. 339. Swartzia angustifoliola

Fig. 340. Swartzia conferta var. conferta Fig. 341. Swartzia buntingii
 Swartzia 407

Fig. 342. Swartzia arborescens

Fig. 343. Swartzia polyphylla

408 F ABACEAE

Fig. 344. Swartzia cowanii

Fig. 345. Swartzia argentea var. argentea

 Swartzia 409

Fig. 346. Swartzia grandifolia

410 F ABACEAE

Fig. 347. Swartzia leptopetala

Fig. 348. Swartzia sp. A

 Swartzia 411

Fig. 349. Swartzia parvifolia

Fig. 350. Swartzia macrocarpa

412 F ABACEAE

Fig. 351. Swartzia pittieri

Fig. 352. Swartzia panacoco var. tepuiensis

 Swartzia 413

Fig. 353. Swartzia

picta var.
bolivarensis
Fig. 354. Swartzia picta var. picta
414 F ABACEAE

Fig. 355. Swartzia sericea var. sericea

Fig. 356. Swartzia

tessmannii
 Swartzia 415

Fig. 357. Swartzia vaupesiana

Fig. 358. Swartzia steyermarkii

416 FABACEAE

60. TARALEA Aubl., Hist. Pl. Guiane 745. 1775.

by Haroldo Cavalcante de Lima and Gerardo A. Aymard C.
Trees or shrubs. Leaves alternate or opposite, odd-pinnate or even-pinnate;
leaflets alternate or opposite, pellucid-punctate or not; rachis cylindric or flattened,
tip often projected beyond the leaflets; stipules small, caducous; stipels none. Inflo-
rescence a terminal panicle. Calyx pellucid-punctate, bilabiate, upper lip with 2
large lobes, lower lip very shortly 3-dentate; petals violet; standard emarginate; keel
petals adnate on lower side. Stamens 10, connate in a sheath open on upper side.
Ovary short-stalked; ovule 1. Fruit elastically dehiscent, 2-valved, flat. Seed com-
pressed-ovoid; cotyledons fused, convolutely wrinkled; radicle central at base of a
deep, closed sinus.
Colombia, Venezuela, Guyana, Suriname, French Guiana, Amazonian Peru
and Brazil; 5 or 6 species, 4 in Venezuela, all in the flora area.

Key to the Species of Taralea

1. Leaflets thick-coriaceous, appressed-puberulent on lower surface and

midrib ...................................................................................... T. crassifolia
1. Leaflets coriaceous to thin-coriaceous, glabrous ....................................... 2
2(1). Leaflets with venation prominent and reticulate on both surfaces, apex
obtuse or retuse-acute; upper lip of calyx obovate-orbicular or orbicu-
lar, 7–11 mm wide .................................................................... T. reticulata
2. Leaflets with venation inconspicuous on upper surface, apex acute or
acuminate; upper lip of calyx oblong or oblong-lanceolate, 3–6 mm wide
................................................................................................................ 3
3(2). Upper lip of calyx coriaceous, oblong-lanceolate; ovary densely pilose;
fruit 5–7 × 3–4 cm; seed 2–2.5 × 1.5–2 cm .......................... T. oppositifolia
3. Upper lip of calyx membranous, oblong to round; ovary pilose only at the
margins; fruit 3–4 × 1.5–2 cm; seed 1–1.5 × 0.5–0.8 cm ............. T. cordata

Taralea cordata Ducke, Arch. Jard. Bot.
Rio de Janeiro 4: 71. 1925. —Dipteryx
cordata (Ducke) R.S. Cowan, Mem. New
York Bot. Gard. 10(1): 152. 1958.
Colombia, Venezuela, Guyana, Suriname,
French Guiana, Brazil; 2 varieties, both in
the flora area.
Key to the Varieties of T. cordata
1. Leaflets 4–6(7), ovate or orbicular, the
base cordate; rachis tip acute or obtuse
......................................... var. cordata
1. Leaflets 1–3(–5), ovate-lanceolate, the
base acute; rachis tip subulate ...............
........................................... var. rigida
T. cordata var. cordata
Shrub to slender tree 1–8 m tall. Wet low-
land and montane forests, 200–1400 m;
Amazonas (Cerro Coro Coro, Cerro Duida,
Cerro Sipapo). Colombia, Guyana, Suri-
name, French Guiana, Brazil. ŠFig. 359.
T. cordata var. rigida (Schery) H.C. Lima,
comb. & stat. nov. —Taralea rigida
Schery, Fieldiana, Bot. 28: 266. 1952.
—Dipteryx rigida (Schery) R.S. Cowan,
Mem. New York Bot. Gard. 10(1): 15.
1958.
Slender shrub 0.5–3 m tall, with elongate
stems. Savannas, 100–1500 m; Bolívar (Ica-
barú, Kavanayén), Amazonas (Cerro Cucuri-
to, Cerro Yapacana, Río Temi, San Fernando
de Atabapo). Brazil (Amazonas: Serra Aracá).
Taralea crassifolia (Benth.) Ducke, Rev.
Int. Bot. Appl. Agric. Trop. 14: 407. 1934.
—Dipteryx crassifolia Benth., J. Bot.
(Hooker) 2: 235. 1840.
 Taralea 417

Fig. 359. Taralea cordata var. cordata

Fig. 360. Taralea crassifolia

418 FABACEAE

Fig. 361. Taralea oppositifolia

Fig. 362. Taralea reticulata

 Tephrosia 419

Shrub or tree 2–20 m tall; flowers pur-
ple. Savannas on sandstone, evergreen for-
ests, 500–1100 m; Bolívar (Auyán-tepui,
Cerro Guacamaya, Cerro Guaiquinima,
Cerro Mahedi, El Paují, 26.5 km east of
Icabarú, Río Asa, upper Río Caroní, Río
Paragua, upper Río Supamo), Amazonas
(Caño Iguapo, Cerro Mahedi, Río Coro
Coro). Guyana, Suriname, French Guiana,
Brazil. ŠFig. 360.
Taralea oppositifolia Aubl., Hist. Pl.
Guiane 745. 1775. —Dipteryx opposi-
tifolia (Aubl.) Willd., Sp. Pl. 3(2): 910.
1802. —Coumarouna oppositifolia
(Aubl.) Taub., Bot. Centralbl. 47: 389.
1891. —Arepillo, Zapatero.
Dipteryx applanata Benth., J. Bot.
(Hooker) 2: 234. 1840.
Dipteryx oppositifolia var. parviflora
Benth., J. Proc. Linn. Soc., Bot.
4(suppl.): 126. 1860.
Tree 8–30 m tall; flowers purple. Forests
on black-water river banks, 100–400 m;
Amazonas (widespread). Colombia, Guyana,
Suriname, French Guiana, Peru, Brazil.
ŠFig. 361.
Taralea reticulata (Benth.) Ducke, Rev.
Int. Bot. Appl. Agric. Trop. 14: 407. 1934.
—Dipteryx reticulata Benth., J. Bot.
(Hooker) 2: 235. 1840.
Taralea steyermarkii Schery, Fieldiana,
Bot. 28: 268. 1952.
Tree 5–30 m tall. Savannas, shrubby for-
ests along savannas, 1000–1400 m; Bolívar
(Ikabarú, Kavanayén, upper Río Aponguao,
Río Karaurín). Guyana. ŠFig. 362.

61. TEPHROSIA Pers., Syn. Pl. 2: 328. 1807.

by Gerardo A. Aymard C.
Herbs or shrubs, erect or trailing; roots usually thick; pubescence usually
dense and matted. Leaves alternate, odd-pinnate; leaflets (1 or 2)3–41, almost al-
ways pubescent, at least on lower surface; rachis usually grooved on upper surface;
exstipellate but tufts of trichomes sometimes present in the axils; stipulate. Inflores-
cence terminal or axillary, racemose, elongate, the flowers in clusters of 2–6 or more
at the nodes, each cluster usually with a primary bract at the base and each pedicel
with a secondary bract at the base. Flowers red, purple, or white. Petals clawed;
standard hairy outside, often densely so; wing petals about as long as the standard
and usually basally adnate to the keel petals. Stamens usually diadelphous, the
vexillar stamen frequently fused to the stamen tube above, free at the base. Ovary
sessile, slender, usually hairy; style bearded above in many species, or glabrous.
Fruit flat, linear or oblong, straight or slighty upcurved, often obliquely contracted
distally, beaked, nonseptate or occasionally slightly septate, usually coiling in dehis-
cence. Seeds several to many, circular to oblong, flattened.
Temperate North America, tropics and subtropics (most diverse in Mexico, Af-
rica, and tropical Australia); ca. 400 species, 8 in Venezuela, 5 of these in the flora
area.
Many species produce rotenone and related compounds which are used as fish
poisons and insecticides. The vernacular name “barbasco” is used for some of the
fish-poisoning species throughout the Neotropics. The taxonomy of some of the taxa
included here probably will not be resolved until the genus has been revised.

Key to the Species of Tephrosia

1. Herbaceous plants, either erect annuals, prostrate to ascending herbs, or

climbing herbaceous vines ..................................................................... 2
1. Erect perennial herbs with woody base, subshrubs, or shrubs to 3.5 m tall
................................................................................................................ 3
420 FABACEAE

2(1). Stems, inflorescences, and leaflets on both surfaces brownish yellow-pu-

bescent; stipules 7–10 mm long; inflorescence 15–30 cm long; legume
12–15-seeded ................................................................................ T. adunca
2. Stems, inflorescences, and lower surfaces of leaflets silvery-strigose or
laxely pilose; stipules 3–5 mm long; inflorescence 5–13 cm long; legume
8–10-seeded .................................................................................. T. cinerea
3(1). Erect shrubs to 3.5 m tall; leaves 15–30 cm long; leaflets 15–40; calyx
5–8 mm long, densely pubescent with long yellow-brown trichomes
..................................................................................................... T. sinapou
3. Erect herbs or subshrubs to 1.5 m tall; leaves 3–10 cm long; leaflets 3–9;
calyx 2–4 mm long, shortly sericeous, pubescent, or laxely pilose ...... 4
4(3). Leaflets 5–9, oblong-obovate, appressed-pubescent on both surfaces or
glabrate; terminal leaflet 2.5–4 cm long; flowers purple or red-purple
......................................................................................................... T. senna
4. Leaflets 3, lanceolate-obovate, the lower surface with silvery-silky pubes-
cence; terminal leaflet 5–10 cm long; flowers yellow ........... T. sessiliflora

Tephrosia adunca Benth., Ann. Nat. Hist. 3:
431. 1839.
Decumbent herb, stems ascending to 50
cm tall; flowers rose-pink. Open savannas,
50–200 m; Bolívar (Maripa, lower Río
Caura). Anzoátegui, Apure, Aragua, Barinas,
Carabobo, Guárico, Monagas, Portuguesa,
Sucre; Colombia, Guyana, Suriname, French
Guiana, Brazil, Bolivia, Paraguay, Argen-
tina, Uruguay.
Galega cathartica Sessé & Moc., Fl. Mexic.
ed. 2, 175. 1894. —Tephrosia cathartica
(Sessé & Moc.) Urb., Symb. Antill. 4: 283.
1905.
Erect suffruticose herb, much-branched;
flowers red-purple. Open areas, edges of
towns, ca. 50 m; Bolívar (near Ciudad Bo-
lívar, El Pao). Falcón, Sucre, Zulia; Mexico,
Central America, West Indies, Colombia,
Brazil.

Tephrosia cinerea (L.) Pers., Syn. Pl. 2:
328. 1807. —Galega cinerea L., Syst.
Nat. ed. 10, 2: 1172. 1759.
Vicia littoralis Jacq., Enum. Syst. Pl. 27.
1760. —Tephrosia cinerea var. littoralis
(Jacq.) Benth. in Mart., Fl. Bras. 15(1):
49. 1859. —Tephrosia littoralis (Jacq.)
Pers., Syn. Pl. 2: 329. 1807.
Prostrate to erect herb or herbaceous vine;
flowers purplish blue. Open savannas, edges
of towns, 50–1100 m; Delta Amacuro (near
Tucupita), Bolívar (near Caicara, near
Ciudad Bolívar, Ciudad Piar, El Manteco, El
Miamo, Gran Sabana, Puerto Ordaz, Re-
presa Guri, lower Río Paragua, base of Ro-
raima-tepui). Amazonas (near Puerto Aya-
cucho). Widespread elsewhere in Venezuela;
Mexico, Central America, West Indies, Co-
lombia, Guyana, Suriname, French Guiana,
Ecuador, Peru, Brazil, Bolivia, Paraguay, Ar-
gentina, Uruguay. ŠFig. 363.
Tephrosia senna H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 458. 1823 [1824].
Tephrosia sessiliflora (Poir.) Hassl.,
Repert. Spec. Nov. Regni Veg. 16: 162.
1919. —Cytisus sessiliflorus Poir. in
Lam., Encycl. suppl. 2: 439. 1811 [1812].
—Generala.
Erect herb or shrub to 1.5 m tall; flowers
yellow. Trachypogon and Curatella savan-
nas, 50–400 m; Bolívar (near Caicara, near
Maripa, lower Río Caroní, Río Caura, Río
Paragua, lower Río Suapure). Amazonas
(Caño Guanay, Cerro Yutajé basin, near
Puerto Ayacucho, Río Parucito, near San
Juan de Manapiare). Apure, Aragua, Bari-
nas, Cojedes, Guárico, Portuguesa, Zulia;
West Indies, Guyana, Suriname, French
Guiana, Colombia, Brazil, Paraguay. ŠFig.
364.
Tephrosia sinapou (Buc’hoz) A. Chev.,
Compt. Rend. Hebd. Séances Acad. Sci.
180: 1522. 1925. —Galega sinapou
Buc’hoz, Hist. Univ. Règne Vég. pl. 994.
1775. —Barbasco, Barbasco caicareño,
Damai (Yekwana).
 Tephrosia 421

Fig. 363. Tephrosia cinerea Fig. 364. Tephrosia sessiliflora

422 FABACEAE

Galega toxicaria Sw., Prod. 108. 1788. —
Tephrosia toxicaria (Sw.) Pers., Syn. Pl.
2: 329. 1807.
Shrub or erect subshrub to 3.5 m tall;
flowers white to red. Edges of towns, open
areas, 50–300 m; Bolívar (lower Río Caroní,
lower Río Caura, Río Paragua, Santa María
de Erebato), Amazonas (upper Río Cunu-
cunuma, Río Padamo, San Carlos de Río Ne-
gro). Anzoátegui, Carabobo, Cojedes, Lara,
Sucre, Zulia; Mexico, Central America, West
Indies, Panama, Colombia, Guyana, Suri-
name, French Guiana, Ecuador, Peru, Brazil,
Bolivia.
Tephrosia sinapou is locally cultivated for
its roots, which are used as a fish poison.

62. TERAMNUS P. Browne, Civ. Nat. Hist. Jamaica 290. 1756.

by Nidia L. Cuello A.
Slender, twining herbs, usually densely pubescent. Leaves pinnate, trifoliolate;
stipules narrow, striate; stipels subulate; leaflets mostly broadly or narrowly ovate,
mucronate. Inflorescences axillary racemes or fascicles; peduncles slender, several-
flowered; bracteoles 2, subtending the calyx or just below the top of the pedicel;
pedicels short, pubescent. Flowers inconspicuous, purplish; bracts small; bracteoles
lanceolate to subulate, striate; hypanthium none. Calyx teeth 5, subequal, or the 2
upper ones connate; standard obovate, narrowed at base, not appendaged, emargin-
ate, glabrous; wing petals narrow, adherent to and longer than the keel petals; keel
petals almost straight, obtuse. Stamens 10, pseudomonadelphous; alternate anthers
abortive or small and sterile. Ovary sessile, linear, pubescent; style incurved, gla-
brous; stigma capitate, thick, not bearded. Fruit linear, mostly straight, 2-valved,

Fig. 365. Teramnus uncinatus

subsp. uncinatus
 Vatairea 423

septate between the seeds, beaked by the persistent inflexed and hooked tip. Seeds
numerous, septate, plump, shiny, lenticulate.
Pantropics, ca. 8 species; 3 in Venezuela, 1 of these in the flora area.

Teramnus uncinatus (L.) Sw., Prodr. 105.
1788. —Dolichos uncinatus L., Sp. Pl. ed.
2, 2: 1019. 1762 [1763].
Pantropics; 3 subspecies, 1 in Venezuela.
T. uncinatus subsp. uncinatus
Herb or vine; flowers purplish. Savannas,
disturbed places, ca. 100 m; Bolívar (lower
Río Caroní). Aragua, Barinas, Cojedes, Dis-
trito Federal, Falcón, Mérida, Miranda,
Monagas, Portuguesa, Sucre, Táchira, Zulia;
Mexico, Central America, West Indies,
Mexico, Colombia, Guyana, Suriname,
French Guiana, Ecuador, Peru, Brazil, Bo-
livia, Paraguay, cultivated in Central Africa
as forage. ŠFig. 365.

63. VATAIREA Aubl., Hist. Pl. Guiane 755. 1775.

by Haroldo Cavalcante de Lima and Gerardo A. Aymard C.
Trees. Leaves alternate, odd-pinnate, crowded at the end of the branches, de-
ciduous; stipules small, caducous; stipels minute. Inflorescence a terminal panicle.
Calyx campanulate or funnelform-turbinate, margin 5-dentate; petals purplish;
standard orbiculate to ovate, emarginate; keel petals briefly joined at the back. Sta-
mens 10, connate in a sheath open on upper side. Ovary with 1 or 2 ovules; style cy-
lindric or flattened. Fruit indehiscent, samaroid or nut-like, winged at the top,
transverse-veined or wing rudimentary, basal seed chambers with spongy-fibrous
mesocarp. Seed 1, with straight radicle.
Mexico, Guatemala, Belize, Honduras, Nicaragua, Panama, Colombia, Venezu-
ela, Guyana, Suriname, French Guiana, Ecuador, Peru, Brazil, Bolivia; 8 species, 2
in Venezuela, both in the flora area.

Key to the Species of Vatairea

1. Leaflets with venation inconspicuous or discretely reticulate on the lower

surface; calyx 1–2.5 cm long, campanulate, densely gray-appressed-
pubescent; gynoecium fusiform, style cylindric; fruit nut-like, orbicular
to ovate, with rudimentary wing ............................................ V. guianensis
1. Leaflets with venation prominent and reticulate on the lower surface; ca-
lyx 5–7 mm long, funnelform-turbinate, sparsely appressed-pubescent;
gynoecium knife-like with flattened style; fruit samaroid, knife-
shaped, with transverse-veined wing at the top ..................... V. paraensis

Vatairea guianensis Aubl., Hist. Pl. Guiane
755. 1775. —Guaboa, Mapadaro.
Ormosia pacimonensis Spruce ex Benth.,
J. Proc. Linn. Soc., Bot. 4(suppl.): 119.
1890.
Vatairea surinamensis Kleinhoonte, Re-
cueil Trav. Bot. Néerl. 22: 403. 1926.
Tree 8–25 m tall; flowers purple. Season-
ally flooded forests mostly along black-water
rivers, 50–200 m; Delta Amacuro (Caño
Atoiba north of Caño Araguao, Serranía de
Imataca), Amazonas (Caño San Miguel, Río
Baría, Río Casiquiare, Río Negro, Río Pasi-
moni, Río Sipapo, Río Temi). Colombia, Guy-
ana, Suriname, French Guiana, Peru, Brazil.
ŠFig. 366.
Vatairea paraensis Ducke, Arch. Jard. Bot.
Rio de Janeiro 5: 140. 1930. —Canelito
negro, Pilón.
Tree 30–40 m tall; flowers purple or blue-
violet. Evergreen lowland to basimontane
forests, 50–300 m; Delta Amacuro (Serranía
de Imataca), Bolívar (El Palmar, mouth of
Río Paragua, Sierra de Lema, Soledad). Suri-
name, Brazil. ŠFig. 367.
424 FABACEAE

Fig. 366. Vatairea guianensis

Fig. 367. Vatairea paraensis

 Vataireopsis 425

Fig. 368. Vataireopsis speciosa

426 F ABACEAE

64. VATAIREOPSIS Ducke, Notizbl. Bot. Gart. Berlin-Dahlem 11: 473. 1932.
by Haroldo Cavalcante de Lima and Gerardo A. Aymard C.
Trees. Leaves alternate, odd-pinnate, crowded at the end of the branches, de-
ciduous; stipules small, caducous; stipels minute. Inflorescence a terminal panicle.
Calyx funnelform-turbinate, 5-dentate; petals blue-violet; standard orbiculate; keel
petals mostly free or occasionally adnate on back. Stamens 9 or 10, connate in a
sheath, except open 1/3 on basal side. Ovary with 1(2) ovules; style flattened. Fruit a
samara, indehiscent, winged at top, transverse-veined; basal seed chambers with 2
small lateral wings. Seed 1; radicle curved.
South America; 4 species, 2 in Venezuela, both in the flora area.

Key to the Species of Vataireopsis

1. Leaflets membranaceous; rachis of inflorescences, bracteoles, and calyx

externally and densely cinereous-appressed-pubescent; standard 7–
11 mm long .................................................................................. V. speciosa
1. Leaflets subcoriaceous; rachis of inflorescences, bracteoles, and calyx fer-
ruginous-appressed-pubescent externally; standard 14–17 mm long
............................................................................................ V. surinamensis

Vataireopsis speciosa Ducke, Notizbl. Bot. Vataireopsis surinamensis H.C. Lima,

Gard. Berlin-Dahlem 11: 474. 1932.
—Tarawuina (Yanomami).
Tree 5–15 m tall; flowers purple. Ever-
green lowland forests, 200–300 m; Amazonas
(Río Matacuni). Guyana, Brazil, Bolivia.
ŠFig. 368.
Rodriguésia 32): 30. 1980. —So-wo
(Yanomami).
Deciduous tree 12–30 m tall; flowers
purple. Evergreen lowland and riparian for-
ests, 200–300 m; Amazonas (Culebra, Río
Cunucunuma). Suriname, Brazil.

65. VIGNA Savi, Nuovo Giorn. Lett. 8: 113. 1824 ser. 3, nom. cons.
by Gerardo A. Aymard C.
Vines or herbs, rarely shrubby, mostly with sturdy rootstocks, lacking hooked
trichomes. Leaves pinnate or subdigitate-trifoliolate, rarely 1-foliolate, mostly en-
tire; stipules sometimes produced below the insertion, sometimes bilobed; stipels
veined, mostly blunt; glabrate. Inflorescences axillary or terminal, pseudoracemose
or subcapitate; rachis contracted, the nodes glandular; bracts and bracteoles cadu-
cous; pedicels 1 or 2 per node, mostly shorter than the calyx. Flowers yellowish or
white, sometimes with some purple or violet. Calyx 2-lipped, the upper lip emargin-
ate, the lowest tooth of the lower lip longest; standard orbicular, auriculate, some-
times appendaged on the dorsal face; wing petals about equaling the standard and
keel petals; keel petals apically curved to the first spiral, often oblique. Vexillar sta-
men free; anthers 10, uniform. Style apically thickened and barbate on the inner
face, caducous, sometimes with a curved beak. Legume linear to oblong, turgid or
compressed, straight or curved, not septate. Seeds reniform or quadrate, the hilum
short or long, a well-developed aril sometimes present.
Pantropics, mostly Paleotropics; ca. 150 species, 14 in Venezuela, 9 of these in
the flora area.
The taxonomy of some of the taxa included here may not be resolved until the
genus has been revised.
 Vigna 427

Key to the Species of Vigna

1. Leaflets linear or linear-oblong, 3–9 mm wide ............................... V. linearis

1. Leaflets narrow-ovate, ovate, ovate-deltoid, or rhombic, > 10 mm wide
................................................................................................................ 2
2(1). Leaflets sometimes 3-lobed, densely whitish-tomentulose; calyx and
pedicels sericeous .................................................................... V. lasiocarpa
2. Leaflets unlobed, glabrous to pilose; calyx and pedicels glabrous to pilose,
but not sericeous .................................................................................... 3
3(2). Leaflets ovate-deltoid, deltoid, or rhombic ................................................ 4
3. Leaflets narrowly ovate, ovate, oblong, lanceolate, or lanceolate-oblong
................................................................................................................ 5
4(3). Flowers white, sometimes with dull purple; fruits 5–7 mm wide ..............
..................................................................................................... V. candida
4. Flowers purple; fruits 2–3 mm wide ...................................... V. peduncularis
5(3). Stipules with a spur at the base, truncate or auriculate (auricle to 15 mm
long) ........................................................................................................ 6
5. Stipules truncate or subauriculate (auricle to 1.5 mm long) at base ....... 7
6(5). Leaflets ovate; stipels glandular; inflorescences pseudoracemose .............
..................................................................................................... V. juruana
6. Leaflets oblong, lanceolate, or oblong-lanceolate; stipels eglandular; inflo-
rescences subumbellate ........................................................... V. longifolia
7(5). Petiolules glabrate; bracteoles and bracts 1–2 mm long ................. V. luteola
7. Petiolules hispidulous or tomentose; bracteoles and bracts 3–4 mm long
................................................................................................................ 8
8(7). Leaflets 5–8 cm wide; flowers white; styles coiled; pods 8–10 mm wide
................................................................................................. V. adenantha
8. Leaflets 2–3.5 cm wide; flowers blue-lavender; styles erect; pods 4–5 mm
.................................................................................................... V. vexillata

Vigna adenantha (G. Mey.) Maréchal,
Mascherpa & Stainier, Taxon 27: 202.
1978. —Phaseolus adenanthus G. Mey.,
Prim. Fl. Esseq. 239. 1818.
Phaseolus truxillensis H.B.K., Nov. Gen.
Sp. (quarto ed.) 6: 451. 1823 [1824].
Perennial vine; flowers purplish or
pink. Disturbed and wet places, near sea
level to 200 m; Delta Amacuro (southeast
of Piacoa), Bolívar (Ciudad Bolívar, Piedra
Marimare on the Río Orinoco, Tumeremo
to Bochinche road), Amazonas (Puerto
Ayacucho). Apure, Aragua, Cojedes, Dis-
trito Federal, Guárico, Miranda, Monagas,
Portuguesa, Sucre, Táchira, Zulia;
Panama, Trinidad, Colombia, Ecuador,
Peru, Brazil, Bolivia, Paraguay, Argen-
tina, cultivated in Central Africa and Asia.
ŠFig. 373.
Vigna candida (Vell.) Maréchal, Mascherpa
& Stainier, Taxon 27: 201. 1978.
—Phaseolus candidus Vell., Fl. Flumin.
311. 1825 [1829].
Phaseolus appendiculatus Benth., Comm.
Legum. Gen. 73. 1837.
Perennial vine; flowers white and purple.
Open places 100–1400 m; Bolívar (Jabillal
on Río Caura, Represa Guri), Amazonas
(Cerro Moriche, Cerro Yutajé, Cuao-Sipapo
massif). Central America, Colombia, Brazil,
Paraguay.
Vigna juruana (Harms) Verdc., Kew Bull.
24: 540. 1970. —Phaseolus juruanus
Harms, Notizbl. Bot. Gart. Berlin-
Dahlem 7: 506. 1921.
Slender vine; flowers yellow. Open and/or
wet places, 100–400 m; Delta Amacuro (Caño
Güiniquina, Curiapo, Río Toro), Bolívar (Isla
Anacoco, Río Botanamo, upper Río Paragua,
Salto Pará), Amazonas (Río Siapa). Panama,
Colombia, Guyana, Suriname, French Gui-
ana, Ecuador, Peru, Brazil, Central Africa.
428 F ABACEAE
Vigna lasiocarpa (Mart. ex Benth.) Verdc.,
Kew Bull. 24: 539. 1970. —Phaseolus
lasiocarpus Mart. ex Benth., Comm. Le-
gum. Gen. 76. 1837. —Frijol de sapo.
Phaseolus diversifolius Pittier, Bol. Técn.
Minist. Agric. 5: 56. 1944.
Herbaceous vine; flowers yellow. Savan-
nas, edges of towns, 50–1000 m; Bolívar
(Ciudad Piar, Gran Sabana, Maripa, Río
Pargueni), Amazonas (La Esmeralda, San
Carlos de Río Negro, San Juan de Mana-
piare, Santa Barbara del Orinoco, Yavita).
Apure, Barinas, Guárico, Portuguesa; Cen-
tral America, Colombia, Guyana, Suriname,
French Guiana, Brazil, Paraguay, Argentina.
Vigna linearis (H.B.K.) Maréchal, Mas-
cherpa & Stainier, Taxon 27: 202. 1978.
—Phaseolus linearis H.B.K., Nov. Gen.
Sp. (quarto ed.) 6: 445. 1823 [1824].
Neotropics; 2 varieties, 1 in Venezuela.
V. linearis var. linearis. —Barbasquillo,
Golondrina.
Perennial herb or vine; flowers red-
purple. Savannas, Mauritia palm swamps,
open places, 50–900 m; widespread in
Bolívar and Amazonas. Widespread else-
where in Venezuela; Mexico, Central Amer-
ica, Antilles, Guyana, Suriname, French
Guiana, Colombia, Ecuador, Peru, Brazil,
Bolivia, Paraguay, Argentina. ŠFig. 369.
Vigna longifolia (Benth.) Verdc., Kew Bull.
24: 541. 1970. —Phaseolus longifolius
Benth., Comm. Legum. Gen. 75. 1837.
Slender vine; flowers yellow. Wet places,
100–200 m; Delta Amacuro (Caño Acoimo),
Amazonas (Minicio). Guárico, Lara; Panama,
Antilles, Colombia, Ecuador, Peru, Brazil,
Bolivia, Paraguay, Argentina, Uruguay.
Vigna luteola (Jacq.) Benth. in Mart., Fl.
Bras. 15(1B): 194. 1859. —Dolichos
luteolus Jacq., Hort. Bot. Vindob. 1: 39.
1770.
Vine; flowers yellow. Wet places, sandy
beaches, secondary scrub, near sea level to
100 m; Delta Amacuro (Caño Araguao, Caño
Güiniquina, Río Acure). Widespread in
northern Venezuela; U.S.A. (Florida), Mexico,
Central America, Antilles, Guyana, Suri-
name, French Guiana, Colombia, Ecuador,
Peru, Brazil, Paraguay, Argentina, Uruguay,
Central Africa, Southest Asia. ŠFig. 370.
Vigna peduncularis (H.B.K.) Fawc. & Ren-
dle, Fl. Jamaica 4: 68. 1920. —Phaseolus
peduncularis H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 447. 1823 [1824].
Neotropics; 3 varieties, 1 in Venezuela.
V. peduncularis var. peduncularis
Slender vine; flowers lavender and white.
Savannas, disturbed open places, 50–900 m;
Bolívar (Caicara, Río Ore in Río Parguaza
basin), Amazonas (La Esmeralda, Puerto
Ayacucho, Río Ocamo, lower Río Ventuari,
San Pedro del Cataniapo, Simarawochi).
Anzoátegui, Apure, Barinas, Distrito Fed-
eral, Guárico, Portuguesa, Sucre, Zulia; Cen-
tral America, Colombia, Ecuador, Peru, Bra-
zil, Bolivia, Paraguay, Argentina, Uruguay.
ŠFig. 371.
Fig. 369. Vigna linearis var. linearis
 Vigna 429

Fig. 370. Vigna luteola Fig. 371. Vigna peduncularis var.

peduncularis

Fig. 372. Vigna vexillata var. vexillata Fig. 373. Vigna adenantha
430 F ABACEAE

Vigna vexillata (L.) A. Rich. in Sagra, Hist.
Phys. Cuba, Bot. Pl. Vasc. 11: 191. 1845.
—Phaseolus vexillatus L., Sp. Pl. 724.
1753.
Pantropics; 6 varieties, 1 in Venezuela.
V. vexillata var. vexillata
Slender vine; flowers blue or lavender
turning white or yellow. Open, wet places,
granitic outcrops, near sea level to 200 m;
Delta Amacuro (Capure, Pedernales, Río
Acure), Amazonas (Puerto Ayacucho, Tama-
tama). Anzoátegui, Aragua, Carabobo, Dis-
trito Federal, Falcón, Mérida, Miranda,
Táchira, Yaracuy, Zulia; Central America,
Antilles, Colombia, Guyana, Suriname,
French Guiana, Ecuador, Peru, Brazil, Bo-
livia. ŠFig. 372.

66. ZOLLERNIA Wied-Neuw. & Nees, Nova Acta Phys.-Med. Acad. Caes. Leop.-
Carol. Nat. Cur. 13(2), pref. 13, t. C & D. 1827.
by Gerardo A. Aymard C.
Trees or shrubs. Leaves simple; petiole developed as a rugose joint extending
into the base of the blade; margin serrate-undulate; stipules free, rigid, borne on the
enlarged node. Inflorescences terminal and axillary racemes, often forming a
panicle; bracts small; bracteoles minute on upper half of pedicel. Calyx closed in bud,
ovoid to lanceolate, acumimate, splitting in 2 valves to short conical or saucer-
shaped, deciduous hypanthium at anthesis; petals 5(6), imbricate, slighty unequal,
shorter than calyx, obtuse, clawed; standard broader than the other petals and exte-
rior to them. Stamens (9)10–12(–15), uniform; filaments short, linear-lanceolate,
sagittate at base; anthers many times longer than the filaments. Ovary subsessile or
stipitate; ovules numerous; style short and subulate; stigma small, obliquely termi-
nal. Fruit indehiscent, drupaceous, ellipsoid, laterally compressed, stipitate, apicu-
late; mesocarp fleshy; endocarp thin and tough; seeds 1–few.
Neotropics; ca. 10 species, 2 in Venezuela, both in the flora area.
Zollernia kanukuensis R.S. Cowan, known from the Kanuku mountains in
Guyana, might eventually be found in the Venezuelan Guayana.

Fig. 374. Zollernia paraënsis

 Zornia 431

Key to the Species of Zollernia

1. Inflorescence rachis, pedicels, and calyx ferruginous-pubescent; anthers

sparsely pilose, with trichomes 1–1.5 mm long ................... Z. grandifolia
1. Inflorescence rachis, pedicels, and calyx sericeous-canescent; anthers gla-
brous or with trichomes ca. 0.2 mm long ................................ Z. paraënsis

Zollernia grandifolia Schery, Fieldiana, Zollernia paraënsis Ducke, Bol. Mus.

Bot. 28: 270. 1952.
Tree to 20 m tall; leaf blades elliptic, mar-
gins serrate; petals pink, 5–6 mm long; an-
thers pilose. Evergreen lowland forests, ca.
200 m; Bolívar (Río Caura, Río Nichare). En-
demic.
Goeldi Paraense Hist. Nat. Ethnogr. 6:
81. 1910. —Masiri (Yekwana).
Zollernia ulei Harms, Notizbl. Königl.
Bot. Gart. Berlin 6: 309. 1915.
Tree to 20 m tall. Evergreen lowland for-
ests, 200–400 m; Amazonas (Río Ocamo, lower
Río Orinoquito). Guyana, Brazil. ŠFig. 374.

67. ZORNIA J.F. Gmel., Syst. Nat. 2: 1076. 1791.

Myriadenus Desv., J. Bot. Agric. 1: 121. 1813.
by Gerardo A. Aymard C.
Annual herbs or suffruticose perennials to 2 m tall. Stems diffusely branched,
prostrate and spreading to erect and ascending. Leaves alternate, digitately 2- or 4-
foliolate; leaflets often pellucid-punctate, exstipellate; stipules subfoliaceous,
paired, peltate, auriculate, 3–11-veined, often punctate. Inflorescence spicate, rarely
racemose, terminal or axillary, interrupted to congested, 1–many-flowered; bracts
paired, lateral, stipuliform, 3–15-veined, sometimes punctate, enclosing a sessile
flower; bracteoles absent. Calyx subhyaline, the tube short, 5-lobed, usually 5–10-
veined, ciliate, the 2 upper lobes connate nearly to the apex, the 2 lateral lobes much
shorter, the lowest oblong or lanceolate, shorter than the upper lip; corolla yellow,
orange, or rarely white, frequently purple-striated; standard suborbicular, clawed,
to 17 mm long; wing petals obliquely obovate or oblong, shorter than standard, usu-
ally clawed and auriculate; keel petals incurved, subrostrate, smaller than the wing
petals. Stamens 10, monadelphous; filaments connate into a closed tube, splitting
dorsally, persistent; anthers alternately longer, subbasifixed, versatile. Ovary
sessile; ovules 2–several, campylotropous; styles filiform basally; stigmas minute,
terminal. Fruit a loment of 2–15 articles, compressed, the upper suture nearly
straight, the lower deeply sinuate, articles glabrous to puberulent or pilose, smooth
or echinate, usually reticulate, indehiscent. Seeds black or dark brown, orbicular to
subreniform, estrophiolate (lacking an appendage to the hilum).
Tropical and subtropical regions worldwide (most diverse in the Neotropics);
ca. 80 species, 15 in Venezuela, 11 of these in the flora area.
The taxonomy of some of the taxa included here probably will not be resolved
until the genus has been revised.

Key to the Species of Zornia

1. Annual herbs, very slender, to 20 cm tall ..................................... Z. herbacea

1. Perennnial herbs, stout near base, 30–100 cm tall ................................... 2
2(1). Stems, petioles, petiolules, leaflets, bracts of inflorescences, and calyx
densely sericeous .......................................................................... Z. sericea
432 F ABACEAE

2. Stems, petioles, petiolules, leaflets, bracts of inflorescences, and calyx gla-

brous or strigose to villous, never sericeous ......................................... 3
3(2). Leaflets linear-filiform, 1–2 mm wide; loment with 3 segments ................
................................................................................................. Z. lasiocarpa
3. Leaflets wider, 3–12 mm wide; loment with 4–15 segments .................... 4
4(3). Leaves 4-foliolate ....................................................................................... 5
4. Leaves 2-foliolate ....................................................................................... 8
5(4). Bracts of inflorescences elliptic, apex obtuse; loment densely glandular
................................................................................................................ 6
5. Bracts of inflorescences lanceolate-elliptic, apex acute; loment eglandular
................................................................................................................ 7
6(5). Leaflets lanceolate to broadly lanceolate, subacute; inflorescences many-
flowered; calyx pilose; loment reticulate ............................. Z. guanipensis
6. Leaflets obovate, retuse; flowers solitary, axillary; calyx glabrous; loment
without reticulation ............................................................... Z. myriadena
7(5). Leaflets oblanceolate, strigose; stipules 7-veined; loment erect, with hairy
bristles ................................................................................... Z. brasiliensis
7. Leaflets ovate-lanceolate or lanceolate; stipules 5-veined; loment curved,
without hairy bristles ................................................................. Z. curvata
8(4). Leaflets epunctate; loment glandular with numerous hairy bristles (never
retrorse) ...................................................................................... Z. diphylla
8. Leaflets punctate or rarely epunctate; loment eglandular with numerous
retrorse bristles ...................................................................................... 9
9(8). Bracts of inflorescences oblong-lanceolate or broadly lanceolate; loment
without reticulation .................................................................... Z. gemella
9. Bracts of inflorescences linear, lanceolate to lance-ovate; loment with evi-
dent reticulation .................................................................................. 10
10(9). Petiole glabrous; inflorescences congested near tip; bracts of inflores-
cences linear, 1–2 mm wide, auricle to 2 mm; loment with retrorse
bristles 0.6–1.5 mm long ............................................................. Z. latifolia
10. Petiole glabrous to pilose; inflorescences crowded; bracts of inflorescences
lanceolate to lance-ovate, 4–8 mm wide, auricle 2–3.5 mm long; loment
with retrorse bristles 0.2–0.4 mm long ................................... Z. reticulata

Zornia brasiliensis Vogel, Linnaea 12: 62.
1838.
Erect perennial herb to 60 cm tall; flowers
yellow. Edges of towns, open areas, savan-
nas, 50–300 m; Bolívar (Caicara, Puerto
Ordaz, Río Aro, near Tumeremo). Monagas,
Nueva Esparta; Brazil.
Zornia curvata Mohlenbr., Webbia 16: 132.
1961.
Erect herb to 50 cm tall; flowers yellow.
Savannas, edges of towns, 50–400 m; Bolívar
(Ciudad Bolívar, upper Río Paramichi). Ama-
zonas (Galipero, Isla Ratón, near Puerto Aya-
cucho). Anzoátegui, Aragua, Distrito Federal,
Lara, Mérida, Miranda, Monagas, Táchira,
Trujillo; Mexico, Panama, Colombia, Ecuador,
Peru, Brazil, Bolivia, Uruguay, Argentina.
Zornia diphylla (L.) Pers., Syn. Pl. 2: 318.
1807. —Hedysarum diphyllum L., Sp.
Pl. 747. 1753.
Perennial prostrate to erect herb; flowers
yellow. Edges of towns, savannas, open ar-
eas, 50–200 m; Bolívar (near Caicara),
Amazonas (Río Sipapo, San Carlos de Río
Negro, San Pedro del Orinoco). Mérida,
Trujillo, Zulia; Colombia, Guyana, Brazil,
Paraguay, Argentina, Zaire, Sri Lanka, Viet-
nam.
Zornia gemella (Willd.) Vogel, Linnaea 12:
61. 1838. —Hedysarum gemellum Willd.,
Sp. Pl. 5(1): 1178. 1810.
Erect herb to 60 cm tall; flowers yellow.
Savannas, open areas, 50–400 m; Bolívar
(Altiplanicie de Nuria, near Tumeremo).

Anzoátegui, Monagas; U.S.A. (Texas), Mex-
ico, Costa Rica, West Indies, Colombia,
Guyana, Suriname, French Guiana, Brazil,
Paraguay, Argentina, Uruguay.
Zornia guanipensis Pittier, Bol. Soc.
Venez. Ci. Nat. 6: 194. 1940.
Zornia tenuifolia var. latifolia Benth. in
Mart., Fl. Bras. 15(1): 81. 1859.
Zornia marajoara Huber, Bol. Mus.
Paraense Hist. Nat. 5: 150. 1908, nom.
nud.
Erect herb to 70 cm tall; flowers yellow.
Savannas, Mauritia palm swamps, 50–200
m; Bolívar (Tumeremo). Anzoátegui, Mona-
gas; Suriname, Brazil (Maranhão).
Zornia herbacea Pittier, Bol. Soc. Venez.
Ci. Nat. 6: 192. 1940.
Annual herb to 20 cm tall; flowers yellow.
Savannas, ca. 200 m; Bolívar (northeast of
Upata). Anzoátegui, Zulia.
Zornia lasiocarpa A.R. Molina, Ceiba 1:
257. 1951.
Erect herb 20–40 cm tall; flowers yellow.
Open areas, edges of towns, 50–200 m;
Bolívar (Ciudad Bolívar, Los Pijiguaos).
Mexico, Honduras, Brazil.
Zornia latifolia Sm. in Rees, Cycl. 39:
Zornia no. 4. 1819.
Herb to subshrub, 30–70 cm tall; flowers
yellow. Pantropics; 2 varieties, 1 in Venezu-
ela.
Z. latifolia var. latifolia. —Pega-pega.
Zornia pubescens H.B.K., Nov. Gen. Sp.
(quarto ed.) 6: 515. 1823 [1824].
Zornia surinamensis Miq., Ann. Mag. Nat.
Hist. 11: 14. 1843.
Herb to subshrub 30–70 cm tall; flowers
yellow. Savannas, edges of towns, 50–1200
m; widespread in Bolívar and Amazonas.
Apure, Cojedes, Monagas, Portuguesa, Tru-
Zornia 433
jillo; Antilles, Colombia, Guyana, Suriname,
French Guiana, Ecuador, Peru, Brazil, Bo-
livia, Paraguay, Argentina, Uruguay, West
Africa.
Zornia myriadena Benth. in Mart., Fl.
Bras. 15(1): 85. 1859.
Erect herb to 60 cm tall; flowers yellow.
Savannas, ca. 100 m; Bolívar (lower Río
Caroní). Cuba, Jamaica, Brazil (Bahia,
Minas Gerais, Pernambuco).
Zornia reticulata Sm. in Rees, Cycl. 39:
Zornia no. 2. 1819.
Erect herb to subshrub 30–70 cm tall;
flowers yellow. Savannas, open areas, 50–
500 m; Bolívar (Ciudad Piar, Las Trincheras,
Urimán), Amazonas (Caño San Miguel, Isla
Ratón, Puerto Ayacucho, Salto Yureba in
lower Río Ventuari basin, Yavita). Aragua,
Barinas, Distrito Federal, Falcón, Guárico,
Monagas, Portuguesa, Zulia; U.S.A. (Ari-
zona, Texas), Mexico. Central America,
Antilles, Colombia, Guyana, Ecuador, Brazil,
Bolivia, Paraguay, Argentina.
Zornia sericea Moric., Pl. Nouv. Amér. 126.
1844. —Crineja.
Erect herb to 50 cm tall; flowers yellow.
Curatella-Trachypogon savannas, edges of
semideciduous forests, 100–200 m; Bolívar
(Caicara, lower Río Caura near Maripa, be-
tween Upata and Villa Lola), Amazonas (San
Fernando de Atabapo). Apure, Mérida; Brazil
(Bahia, Maranhão, Mato Grosso, Minas
Gerais, Pará, Piaui). ŠFig. 375.
Fig. 375. Zornia sericea