Vol. 2 Cover. Fruits & Seeds

Agricultural Research Genera in the Subfamily Service Technical Bulletin Faboideae (Fabaceae) Number 1890 Volume II

Fruits of A, Centrolobium paraense E.L.R. Tulasne. B, Laburnum anagyroides F.K. Medikus. C, Adesmia boronoides J.D. Hooker. D, Hippocrepis comosa, C. Linnaeus. E, Campylotropis macrocarpa (A.A. von Bunge) A. Rehder. F, Mucuna urens (C. Linnaeus) F.K. Medikus. G, Phaseolus polystachios (C. Linnaeus) N.L. Britton, E.E. Stern, & F. Poggenburg. H, Medicago orbicularis (C. Linnaeus) B. Bartalini. I, Riedeliella graciliflora H.A.T. Harms. J, Medicago arabica (C. Linnaeus) W. Hudson.

Kirkbride is a research botanist, U.S. Department of Agriculture, Agricultural Research Service, Systematic Botany and Mycology Laboratory, BARC West Room 304, Building 011A, Beltsville, MD, 20705-2350 (email = [email protected]). Gunn is a botanist (retired) from Brevard, NC (email = [email protected]). Weitzman is a botanist with the Smithsonian Institution, Department of Botany, Washington, DC. Contents

Procedures ...... 3 Fruit morphology ...... 4 Fruit ...... 4 Loment ...... 8 Epicarp ...... 9 Mesocarp ...... 12 Endocarp ...... 13 Seeds ...... 14 Funiculus ...... 14 Aril ...... 15 Seed morphology ...... 16 Seed ...... 18 Testa ...... 20 Raphe ...... 22 Hilum ...... 24 Lens ...... 26 Endosperm ...... 28 Cotyledons ...... 28 Embryonic axis ...... 31 Radicle ...... 31 Plumule ...... 32 Seed key to three subfamilies of Fabaceae ...... 33 Seed keys to genera of subfamily Faboideae ...... 33 Master key to 16 seed keys and one genus ...... 33 Seed key 1: Aril present; fleshy. Cotyledons entire over radicle...... 33 Seed key 2: Aril present; fleshy. Cotyledons notched at radicle...... 35 Seed key 3: Aril present; fleshy. Cotyledons split at radicle...... 36 Seed key 4: Aril present; dry. Cotyledons entire over radicle. Hilum raised or flush...... 38 Seed key 5: Aril present; dry. Cotyledons entire over radicle. Hilum recessed...... 39 Seed key 6: Aril present; dry. Cotyledons notched at radicle. Endosperm present...... 43 Seed key 7: Aril present; dry. Cotyledons notched at radicle. Endosperm absent...... 48 Seed key 8: Aril present; dry. Cotyledons split over radicle. Hilum raised or flush...... 52 Seed key 9: Aril present; dry. Cotyledons split over radicle. Hilum recessed...... 54 Seed key 10: Aril absent. Cotyledons entire over radicle. Endosperm absent...... 58 Seed key 11: Aril absent. Cotyledons entire over radicle. Hilum raised or flush. Endosperm present...... 60 Seed key 12: Aril absent. Cotyledons entire over radicle. Hilum recessed. Endosperm present...... 63 Seed key 13: Aril absent. One (1) cotyledon scooped out to accommodate plicate radicle and other cotyledon entire...... 67 Seed key 14: Aril absent. Cotyledons notched at radicle...... 67 Seed key 15: Aril absent. Cotyledons split over radicle. Hilum raised or flush...... 71 Seed key 16: Aril absent. Cotyledons split over radicle. Hilum recessed...... 73

Synopses of fruit and seed characters Swartzieae (1.01-1.15)...... 78 Sophoreae (2.01-2.46) ...... 118 Dipteryxeae (3.01-3.03)...... 220 Dalbergieae (4.01-4.17) ...... 226 Abreae (5.01) ...... 270 Amorpheae (6.01-6.08) ...... 274 Millettieae (A-X) ...... 292 Robinieae (8.01-8.12) ...... 388 Indigofereae (9.01-9.07) ...... 424 Phaseoleae (10.01-10.83) ...... 434

Synopses of fruit and seed characters Desmodieae (11.01-11.25) ...... 636 Psoraleeae (12.01-12.09) ...... 696 Loteae (13.01-13.17) ...... 716 Aeschynomeneae (14.01-14.26) ...... 754 Adesmieae (15.01) ...... 810 Galegeae (16.01-16.22) ...... 814 Carmichaelieae (17.01-17.05) ...... 870 Hedysareae (18.01-18.07) ...... 882 Fabeae (19.01-19.05) ...... 894 Cicereae (20.01) ...... 910 Trifolieae (21.01-21.06) ...... 914 Brongniartieae (22.01-22.02) ...... 934 Bossiaeeae (23.01-23.10) ...... 938 Mirbelieae (24.01-24.26) ...... 960 Podalyrieae (25.01-25.09) ...... 1008 Hypocalypteae (26.01) ...... 1028 Crotalarieae (27.01-27.11)...... 1030 Euchresteae (28.01) ...... 1056 Thermopsideae (29.01-29.06) ...... 1058 Genisteae (30.01-30.25) ...... 1072 Character list ...... 1130 Literature cited...... 1160 Scientific name index ...... 1185 Desmodieae (11.01Ð11.25) smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon Genus: Brya P. Browne lobes; without hilar sinus; with umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a Phylogenetic Number: 11.01. bloom; colored; monochrome; reddish brown; glabrous; smooth; chartaceous. Fracture lines absent. Rim absent. Tribe: Desmodieae. Wings absent. Raphe from lens to base of seed and terminating; not bifurcating; darker than testa; dark Subtribe: Bryinae. reddish brown; raised. Hilum fully concealed; concealed by funicular remnant; without faboid split; Species Studied—Species in Genus: 4 spp.—4 spp. punctiform or larger than punctiform; 0.2Ð0.5 mm long; with straight outline; oblong; between cotyledon and Fruit a legume or loment (or loment segment); 0.8Ð2.5 radicle lobe; flush; within halo. Hilum halo color darker (upper length is estimated) ϫ 0.6Ð0.9 ϫ 0.07Ð0.08 cm; than testa. Lens not discernible. Endosperm absent. with deciduous calyx; without orifice formed by Cotyledons smooth; both outer faces convex; both the curving of fruit or fruit segments; straight or curved (or same thickness; both more or less of equal length; not slightly curved); not plicate; not twisted; asymmetrical; folded; margin entire 180 degrees from base of radicle; moniliform (with more or less elongated isthmuses); similar at apex; partially concealing radicle; notched at when asymmetrical with 1 straight and 1 curved suture radicle; with lobes; with lobes not touching; without or both sutures parallelly curved; widest near middle or basal groin formed by lobes; with the interface division D-shaped; not inflated; flattened; without beak; rounded terminating at base of radicle; without margins re- at apex; apex oblique with longitudinal axis of fruit; cessed; reddish tan; inner face flat; glabrous around short tapered at base; base aligned with longitudinal base of radicle. Embryonic axis deflexed; oblique to axis of fruit; with the apex and base uniform in texture; length of seed; without a joint evident between the membranous; seed chambers externally visible; with the radicle and the cotyledons. Radicle linear; deflexed and raised seed chambers not torulose. Fruit margin not parallel to cotyledon length; centered between cotyle- constricted; without sulcus; plain. Fruit substipitate or dons; less than 1/2 length of cotyledons. Plumule nonstipitate. Fruit indehiscent. Replum invisible. moderately developed; glabrous. Loment indehiscent. Loment segments (articles) inconspicuous; 8Ð15 mm long; widest across seed area; Distribution: Cuba (1 sp.) extending to Jamaica and another with all essentially similar in shape; D-shaped or species to Hispaniola. curved. Epicarp dull; monochrome; brown (dark reddish to greenish); pubescent and indurate; with 2 or Notes: Ohashi et al. (1981) started their treatment of the 3 types of pubescence; puberulent; with pubescence Desmodieae with these thoughts: “A sensible classifica- gray (short and plain tipped) or golden (long and gland tion of Desmodieae is prejudiced by the traditional tipped); with golden hooked and gray plain hairs; with over-weighting of fruit characters.” [sic] “The fruit pubescence uniformly distributed; with simple and normally consists of indehiscent jointed articles, but glandular hairs; pliable; with hair bases plain; glandu- fruits that open have arisen at least seven times...” They lar; with glandular hairs; without spines; not smooth; supplemented their text with a fruit-seed plate. They with elevated features; reticulately veined; not tubercu- placed Brya and Cranocarpus (11.02), the only two late; not exfoliating; without cracks. Mesocarp absent. New World endemic genera, in the new subtribe Endocarp more or less glossy; monochrome; tan; Bryinae, “characterized most notably by glochidiate fibrous (to somewhat smooth in seed chamber); septate; hairs.” Bailey et al. (1997), using the chloroplast rpl2 with septa thicker than paper, firm; with septa intron and ORF184, suggested that Brya, Cranocarpus, eglandular; chartaceous; not exfoliating; remaining Phylacium (11.22), and Neocollettia (11.26) are not fused to epicarp; entire. Seeds 1 or 2; length parallel members of Desmodieae and that they probably belong with fruit length; neither overlapping nor touching; in 1 in Aeschynomeneae (14). Lewis (1988) provided a series. Funiculus measured; up to 1 mm long; of 1 plate showing the fruit and seed of B. ebenus. length only; flattened; curved. Aril absent. Brya: B. ebenus (C. Linnaeus) A.-P. de Candolle (AÐE). A, Seed 3.5Ð5 ϫ 2Ð3.5 ϫ 0.5Ð0.6 mm; not overgrown; not Fruits (ϫ 3); B, seeds (ϫ 5.4); CÐD, testa (ϫ 50, angular; asymmetrical; reniform; flattened; with surface ϫ 1000); E, embryos (ϫ 5).

637 Genus: Cranocarpus G. Bentham bloom; colored; monochrome; reddish brown; glabrous; smooth; chartaceous. Pseudopleurogram absent. Phylogenetic Number: 11.02. Fracture lines absent. Rim absent. Wing(s) absent. Raphe not visible. Hilum fully concealed; concealed by Tribe: Desmodieae. aril remnant; with faboid split (assumed); larger than punctiform; ca. 0.4 mm long; with straight outline; Subtribe: Bryinae. linear; flush; not within corona, halo, or rim. Lens discernible; ca. 0.3 mm long; with margins straight; Species Studied—Species in Genus: 2 spp.—3 spp. linear; not in groove of raphe; adjacent to hilum; ca. 0.2 mm from hilum; flush; dissimilar color from testa; Fruit a loment (or loment segment or a 1-seeded loment); 1Ð black; not within corona, halo, or rim. 2 ϫ 0.5Ð0.8 cm; with persistent calyx; with calyx shorter than fruit; without orifice formed by curving of Distribution: Brazil. fruit or fruit segments; straight; not plicate; not twisted; asymmetrical; irregular or reniform; when asymmetri- Notes: Bailey et al. (1997), using the chloroplast rpl2 intron cal with both sutures unequally curved or 1 straight and and ORF184, suggested that Cranocarpus is not a 1 curved; narrowing in several places, resembling member of Desmodieae and that it belongs in Desmodium (11.09) fruit; not inflated; flattened; Aeschynomeneae. Harley (1978) monographed without beak; rounded at apex; apex right-angled or Cranocarpus (with two species), and Fernandes and more with longitudinal axis of fruit; short tapered at Bezerra (1979) named a new species. According to base; base aligned with longitudinal axis of fruit; with Harley (1978), C. mezii P.H.W. Taubert has one- or the apex and base uniform in texture; membranous; two-seeded fruits that are loments, and C. martii has seed chambers externally visible. Fruit margin con- one-seeded fruits by abortion of the lower ovule. The stricted or not constricted; constricted only on 1 margin; fruits of C. martii are homologous to those of C. mezii, without sulcus; plain. Fruit wings absent. Fruit and therefore are one-seeded loments. We had only one nonstipitate. Fruit indehiscent. Replum invisible. misshapen seed and one fruit for study. Loment indehiscent. Loment segments (articles) inconspicuous; 8Ð13 mm long; widest across seed area; with all essentially similar in shape; D-shaped (to galeiform). Epicarp dull; monochrome; reddish brown; pubescent and indurate; with 1 type of pubescence; villous; with pubescence gray; with pubescence uniformly distributed; with simple and glandular hairs (the simple hairs longer than the glandular ones); stiff and pliable; with hair bases plain; with glandular hairs; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only. Aril absent.

Seed ca. 5.5 mm long; ca. 4 mm wide; ca. 1 mm thick; not overgrown; not angular; asymmetrical; reniform; flattened; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces; without medial ridge on each face. Cuticle not exfoliating; not inflated; not wrinkled. Testa without pieces of adhering epicarp; not adhering to endocarp; free from endocarp; dull; not modified by a Cranocarpus: C. martii G. Bentham (AÐD). A, Fruit (ϫ 3.7); B, seed (ϫ 8.4); CÐD, testa (ϫ 50, ϫ 1000).

639 Genus: Arthroclianthus H.E. Baillon Seed 7Ð9 ϫ 4Ð4.5 ϫ 0.4Ð0.5 mm; not overgrown; not angular; asymmetrical; reniform (sub-); flattened; with Phylogenetic Number: 11.03. surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Tribe: Desmodieae. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; Subtribe: Desmodiinae. smooth; chartaceous. Fracture lines absent. Rim absent. Wings absent. Raphe from hilum to lens (near base of Species Studied—Species in Genus: 13 spp.—ca. 10 spp. seed); not bifurcating; darker than testa; dark reddish brown; raised. Hilum fully concealed; concealed by Fruit a loment (or loment segment); 8Ð14 ϫ 0.5Ð1.2 ϫ 0.1Ð funicular remnant; without faboid split; punctiform; 1.15 cm; with persistent calyx; with calyx shorter than marginal according to radicle tip; flush; within rim. fruit; without orifice formed by curving of fruit or fruit Hilum rim color darker than testa. Lens discernible; less segments; straight or curved (slightly); not plicate; not than 0.5 mm in length; with margins curved; circular; twisted; asymmetrical or symmetrical; moniliform not in groove of raphe; adjacent to hilum; 2.5 mm from (some with elongated isthmuses) or linear; when hilum (length of raphe); mounded; similar color as asymmetrical with 1 straight and 1 curved suture or testa; darker than testa; dark reddish brown; not within both sutures parallelly curved; narrowing in several corona, halo, or rim. Endosperm absent. Cotyledons places, resembling Desmodium (11.09) fruit; not smooth; both outer faces convex; both the same inflated; flattened; with beak; straight; with solid beak thickness; both more or less of equal length; not folded; the same color and texture as fruit; rounded at apex; margin entire 180 degrees from base of radicle; similar apex aligned with longitudinal axis of fruit; short at apex; not concealing radicle; entire over radicle; tapered at base; base aligned with longitudinal axis of without lobes; with the interface division terminating at fruit; with the apex and base uniform in texture; base of radicle; without margins recessed; yellow; inner coriaceous; seed chambers externally visible; with the face flat; glabrous around base of radicle. Embryonic raised seed chambers not torulose. Fruit margin axis deflexed; oblique to length of seed; without a joint constricted along both margins; without sulcus; plain. evident between the radicle and the cotyledons. Radicle Fruit wings absent. Fruit stipitate; with the stipe 14Ð40 linear; deflexed and parallel to cotyledon length; mm long. Fruit indehiscent. Replum invisible. Loment centered between cotyledons; less than 1/2 length of indehiscent. Loment segments (articles) inconspicuous; cotyledons. Plumule moderately developed; glabrous. 14Ð35 mm long; widest across seed area; with all essentially similar in shape; D-shaped or oblong. Distribution: New Caledonia. Epicarp dull; monochrome; greenish brown to brown; pubescent and indurate or glabrous; with 1 type of Notes: Hochreutiner (1907) monographed the genus, and pubescence; pilose; with pubescence gray; with Ohashi et al. (1981) noted that Arthroclianthus is pubescence uniformly distributed; with simple hairs; scarcely distinct from Nephrodesmus (11.04). We are pliable; with hair bases plain; eglandular; without unable to ascertain which of about ten species Ohashi et spines; not smooth; with elevated features; reticulately al. recognized. veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface uniformly veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; greenish tan; smooth; septate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 4Ð9; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; triangular; straight. Aril absent. Arthroclianthus: A. pseudo-malaccensis E. Vieillard ex A.T. Brongnairt & J.A.A. Gris (CÐE), A. spp. (AÐB). A, Fruit segments and articles (ϫ 1); B, seeds (ϫ 6.8); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

641 Genus: Nephrodesmus A.K. Schindler Caledonia (Royal Botanic Gardens, Kew) and N. albus collected by B. Balansa (2808), New Caledonia (The Phylogenetic Number: 11.04. Natural History Museum (London)). Another sheet of M. balansa (Royal Botanic Gardens, Kew) was studied Tribe: Desmodieae. from a colored photograph.

Fruit a loment (or loment segment); up to 15 ϫ 0.7Ð0.85 ϫ 0.1Ð0.4 (assumed) cm; with persistent or deciduous calyx; with calyx shorter than fruit; without orifice formed by curving of fruit or fruit segments; straight or curved (slightly); not plicate; not twisted; asymmetrical or symmetrical; moniliform; when asymmetrical with both sutures parallelly curved; not inflated; flattened; without or with beak; straight; with solid beak the same color and texture as fruit; rounded at apex; apex aligned with longitudinal axis of fruit; rounded at base; base oblique with longitudinal axis of fruit; with the apex and base uniform in texture; membranous or chartaceous (both assumed); seed chambers externally visible. Fruit margin constricted along both margins; without sulcus; plain. Fruit wings absent. Fruit stipitate; with the stipe 10Ð15 mm long. Replum invisible. Loment indehiscent. Loment segments (articles) inconspicuous; 15Ð20 mm long; widest across seed area; with all essentially similar in shape; oblong. Epicarp dull; monochrome; greenish brown or green; pubescent and indurate; with hairs appressed; with 1 type of pubescence; with pubescence reddish brown; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; longitudinally veined relative to fruit length (assumed); not tuberculate; not exfoliating; without cracks. Mesocarp present or absent (assumed to be like Desmodium (11.09). Endocarp septate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 8Ð9; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only.

Notes: Ohashi et al. (1981) noted that Nephrodesmus is scarcely distinct from Arthroclianthus (11.03). Neither seeds nor fruits were available for study. The only Nephrodesmus: N. albus A.K. Schindler (A). A, Fruits from materials available for study were a xeroxed herbarium a colored photograph of a specimen collected by M. sheet labeled N. sericeus (B.P.G. Hochreutiner) A.K. Balansa 2808 New Caledonia and accessioned at Kew Schindler collected by S.-R. Lenormand (2941), New (ϫ 0.6).

643 Genus: Trifidacanthus E.D. Merrill Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; Phylogenetic Number: 11.05. smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with Tribe: Desmodieae. faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; Subtribe: Desmodiinae. 0.4 mm long; with curved outline; elliptic; marginal according to radicle tip or between cotyledon and Species Studied—Species in Genus: 1 sp.—1 sp. radicle lobe; recessed; not within corona, halo, or rim. Lens discernible; less than 0.5 mm in length or equal to Fruit a loment (or loment segment); 2.8Ð6 ϫ 0.5Ð0.7 ϫ or greater than 0.5 mm in length; 0.5 mm long; with 0.055Ð0.057 cm; with persistent calyx; with calyx margins straight; linear; not in groove of raphe; shorter than fruit; without orifice formed by curving of confluent with hilum; recessed; dissimilar color from fruit or fruit segments; straight or curved (slightly); not testa; darker than testa; black; not within corona, halo, plicate; not twisted; asymmetrical; moniliform; when or rim. Cotyledons smooth; both outer faces convex; asymmetrical with 1 straight and 1 curved suture or both the same thickness; both more or less of equal both sutures parallelly curved; narrowing in several length; not folded; margin entire 180 degrees from base places, resembling Desmodium (9.09) fruit; not inflated; of radicle; similar at apex; not concealing radicle; entire flattened; with beak; straight or declined; with solid over radicle; without lobes; with the interface division beak the same color and texture as fruit; rounded at terminating at base of radicle; without margins re- apex; apex aligned with longitudinal axis of fruit; short cessed; inner face flat. Embryonic axis deflexed; tapered at base; base aligned with longitudinal axis of oblique to length of seed; without a joint evident fruit; with the apex and base uniform in texture; between the radicle and the cotyledons. Radicle linear; membranous; seed chambers externally visible; with the lobe tip straight; oblique to cotyledons; centered raised seed chambers not torulose. Fruit margin between cotyledons; less than 1/2 length of cotyledons. constricted on 1 margin and slightly constricted on the Plumule rudimentary. other margin; without sulcus; plain. Fruit wings absent. Fruit stipitate; with the stipe 9Ð25 mm long. Fruit Distribution: Vietnam, Hainan, Philippines, and Lesser indehiscent. Replum invisible. Loment indehiscent. Sunda Islands. Loment segments (articles) inconspicuous; 2.5Ð10 mm long; widest across seed area; with all essentially Notes: Merrill (1917) founded the genus without seeing similar in shape; D-shaped, oblong, or quadrangular. fruits; 2 years later he described the fruits (Merrill Epicarp dull; monochrome; brown or tan; pubescent 1919). Ohashi et al. (1996) revised the genus and and indurate; with 1 type of pubescence; puberulent; concluded that it is a distinct, montypic genus in tribe with pubescence gray; with pubescence uniformly Desmodieae. distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; reddish brown; smooth; septate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 3Ð8; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; triangular; straight. Aril dry; rim-aril; reddish brown.

Seed 2Ð4.5 ϫ 1Ð1.4 ϫ 0.7Ð0.8 mm; not overgrown; not Trifidacanthus: T. unifoliolatus E.D. Merrill (AÐE). A, Fruit angular; asymmetrical; reniform; compressed; with (immature and nearly entire) and article (mature) (ϫ 3); surface smooth; without visible radicle and cotyledon B, seed (ϫ 7.6); CÐD, testa (ϫ 50, ϫ 1000); E, lobes; without hilar sinus; without umbo on seed faces. embryos (ϫ 7).

645 Genus: Dendrolobium (R. Wight & G.A.W. Arnott) G. touching; in 1 series. Funiculus measured or less than Bentham 0.5 mm long; 0.5 mm long; of 1 length only; thick; straight. Aril absent. Phylogenetic Number: 11.06. Seed 3Ð3.6 ϫ 2Ð2.8 ϫ 1Ð1.6 mm; not overgrown; not Tribe: Desmodieae. angular; asymmetrical; oblong, ovate, elliptic, or quadrangular; compressed; with surface smooth; Subtribe: Desmodiinae. without visible radicle and cotyledon lobes; without hilar sinus; without or with umbo on seed faces; with Species Studied—Species in Genus: 11 spp.—13 spp. umbo on both faces of seed. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; Fruit a loment (or loment segment) or legume; unilocular; monochrome; reddish brown; glabrous; smooth; 0.9Ð4.5 ϫ 0.5Ð0.85 ϫ 0.12Ð0.3 cm; with deciduous coriaceous. Fracture lines absent. Rim absent. Wings calyx; without orifice formed by curving of fruit or fruit absent. Raphe not visible. Hilum visible or fully segments; slightly curved, straight, or curved; not concealed; concealed by funicular remnant; with faboid plicate; not twisted; asymmetrical; moniliform, linear, split; with the lips of the faboid split the same color as or falcate (sub-); when asymmetrical with both sutures the rest of the hilum; punctiform; marginal according to parallelly curved or 1 straight and 1 curved suture; radicle tip; recessed; within rim. Hilum rim color of narrowing in several places, resembling Desmodium testa. Lens discernible; less than 0.5 mm in length or (11.09) fruit; not inflated; compressed or flattened; with equal to or greater than 0.5 mm in length; 0.5 mm long; or without beak; straight; with solid beak the same with margins straight or curved; triangular or linear; 2 color and texture as fruit; short tapered at apex; apex circular mounds separated by groove; not in groove of aligned with longitudinal axis of fruit; short tapered or raphe; confluent with hilum; mounded; dissimilar color rounded at base; base aligned with longitudinal axis of from testa; darker than testa; dark reddish brown; not fruit; with the apex and base uniform in texture; within corona, halo, or rim. Endosperm thin; covering ligneous; seed chambers externally visible; with the entire embryo; adnate to testa or adnate to embryo. raised seed chambers not torulose. Fruit margin Cotyledons smooth; both outer faces convex; both the constricted along both margins; without sulcus; plain. same thickness; both more or less of equal length; not Fruit substipitate or nonstipitate. Fruit indehiscent. folded; margin entire 180 degrees from base of radicle; Replum invisible. Loment an intact article; indehiscent. similar at apex; not concealing radicle; entire over Loment segments (articles) inconspicuous; 7Ð8 mm radicle; without lobes; with the interface division long; widest across seed area; with all essentially terminating at base of radicle; without margins re- similar in shape; D-shaped or quadrangular. Epicarp cessed; yellow; inner face flat; glabrous around base of dull; monochrome; reddish brown; pubescent and radicle. Embryonic axis deflexed; oblique to length of indurate, glabrous, or glabrate; with hairs appressed or seed; without a joint evident between the radicle and erect; with 1 type of pubescence; tomentose; with the cotyledons. Radicle linear or bulbose; deflexed and pubescence golden; with pubescence uniformly parallel to cotyledon length or width; centered between distributed; with simple hairs; pliable; with hair bases cotyledons; less than 1/2 length of cotyledons. Plumule plain; eglandular; without spines; not smooth; with moderately developed or rudimentary; glabrous. elevated features; not veined or reticulately veined (D. dispermum (B. Hayata) A.K. Schindler and D. Distribution: Tropical Asia, Australia, and Indian Ocean rostratum (A.K. Schindler) A.K. Schindler); not coastal region. tuberculate; wrinkled; not exfoliating; without cracks. Mesocarp thick; surface not veined; 1- or 2-layered; Notes: Ohashi (1973) monographed the genus and provided without balsamic vesicles; without fibers; solid; with illustrations of fruits and seeds. He (Ohashi 1997b,c) honeycomb layer over solid layer; ligneous. Endocarp also reviewed the species of Malesia, Australia, and Fiji glossy; monochrome; tan (to reddish); smooth; septate; and D. cumingianum G. Bentham. with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining Dendrolobium: D. lanceolatum (S.T. Dunn) A.K. Schindler fused to mesocarp and epicarp; entire. Seeds 1Ð8; (CÐE), D. spp. (AÐB). A, Fruit segment and articles length parallel with fruit length; neither overlapping nor (ϫ 2.3); B, seeds (ϫ 2.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

647 Genus: Phyllodium A.N. Desvaux touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened; straight. Aril dry; rim-aril; Phylogenetic Number: 11.07. tan.

Tribe: Desmodieae. Seed 2.2Ð4 ϫ 1.8Ð3 ϫ 1 mm; not overgrown; not angular; symmetrical or asymmetrical; reniform; compressed; Subtribe: Desmodiinae. with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on Species Studied—Species in Genus: 6 spp.—6 spp. seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome; Fruit a loment (or loment segment) or legume; unilocular; reddish brown or tan; glabrous; smooth; chartaceous. 0.4Ð4 ϫ 0.3Ð0.4 ϫ 0.1Ð0.15 cm; with persistent calyx; Fracture lines absent. Rim absent. Wings absent. Raphe with calyx shorter than fruit; without orifice formed by not visible. Hilum visible; with faboid split; with the curving of fruit or fruit segments; straight or curved (or lips of the faboid split the same color as the rest of the slightly curved); not plicate; not twisted; asymmetrical; hilum; larger than punctiform or punctiform; 0.5 mm when asymmetrical with 1 straight and 1 curved suture; long; with curved outline; circular; marginal according widest near middle or D-shaped; not inflated; flattened; to radicle tip; recessed; not within corona, halo, or rim. with beak; straight or coiled; with solid beak the same Lens discernible; less than 0.5 mm in length; with color and texture as fruit; rounded at apex; apex aligned margins straight or curved; linear or wedge-shaped; or oblique with longitudinal axis of fruit; short tapered circular; not in groove of raphe; confluent with hilum; or rounded at base; base aligned with longitudinal axis mounded; dissimilar color from testa; darker than testa; of fruit; with the apex and base uniform in texture; black or tan (reddish); not within corona, halo, or rim. membranous; seed chambers externally visible; with the Endosperm thin; covering entire embryo; adnate to raised seed chambers not torulose. Fruit margin embryo. Cotyledons smooth; both outer faces convex; constricted on 1 margin and slightly constricted on the both the same thickness; both more or less of equal other margin; without sulcus; plain or embellished; with length; not folded; margin entire 180 degrees from base fringe (of hair). Fruit wings absent. Fruit nonstipitate. of radicle; similar at apex; not concealing radicle; entire Fruit indehiscent. Replum invisible. Loment indehis- over radicle; without lobes; with the interface division cent. Loment segments (articles) inconspicuous; 3Ð4 terminating at base of radicle; without margins re- mm long; widest across seed area; with upper and lower cessed; white or tan (to nearly white); inner face flat; 1 different shape than middle ones; D-shaped or glabrous around base of radicle. Embryonic axis quadrangular. Epicarp dull; monochrome; reddish deflexed; oblique to length of seed; without a joint brown; pubescent and indurate; with hairs erect or evident between the radicle and the cotyledons. Radicle appressed; with 1 type of pubescence; pilose; with linear; deflexed and parallel to cotyledon length; pubescence golden or gray; with pubescence uniformly centered between cotyledons; less than 1/2 length of distributed; with simple hairs; pliable; with hair bases cotyledons. Plumule moderately developed; glabrous. plain; eglandular; without spines; not smooth; with elevated features; veined or not veined; reticulately Distribution: Southern Asia and northern Australia. veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without Notes: Ohashi (1973) monographed the genus and provided balsamic vesicles; without fibers; solid; chartaceous. a wider range of fruit and seed characters than our Endocarp dull; monochrome or mottled and streaked plate. (area below seed somewhat streaked and mottled to lighter colored); reddish brown or tan (reddish); with mottling over seed chambers; with brown (reddish) or tan (reddish) overlay; smooth; septate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð9; length Phyllodium: P. pulchellum (C. Linnaeus) A.N. Desvaux transverse to fruit length; neither overlapping nor (CÐE), P. spp. (AÐB). A, Fruits (1- and 2-seeded) (ϫ 5); B, seeds (ϫ 6.8); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

649 Genus: Aphyllodium (A.-P. de Candolle) F. Gagnepain cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; with Phylogenetic Number: 11.08. umbo on both faces of seed. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; Tribe: Desmodieae. monochrome or mottled and streaked; greenish tan, green, or purple; with purple overlay (more frequent Subtribe: Desmodiinae. than background color); glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe Species Studied—Species in Genus: 1 sp.—4 spp. not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the Fruit a loment (or loment segment) or legume; 0.2Ð1.7 ϫ hilum; punctiform (much smaller and much less 0.3Ð0.35 ϫ 0.12Ð0.15 cm; with persistent or deciduous conspicuous than the darker lens on greenish seeds); calyx; with calyx shorter than fruit; without orifice between cotyledon and radicle lobe; recessed; not formed by curving of fruit or fruit segments; straight; within corona, halo, or rim or within halo. Hilum halo not plicate; not twisted; symmetrical; elliptic or color of testa (but not mottled with purple). Lens moniliform; not inflated; flattened; with beak; straight; discernible (greenish seeds) or not discernible (purplish with solid beak the same color and texture as fruit; seeds); less than 0.5 mm in length; with margins tapered, rounded, or truncate (if lower of two articles) straight or curved; square; 2 circular mounds separated at apex; apex aligned or oblique with longitudinal axis by groove; not in groove of raphe; confluent with of fruit; short tapered or truncate (if upper article of hilum; mounded; dissimilar color from testa; darker two) at base; base aligned with longitudinal axis of than testa; reddish brown; not within corona, halo, or fruit; with the apex and base uniform in texture; rim. Endosperm thin; covering entire embryo; adnate to coriaceous; seed chambers externally visible; with the testa. Cotyledons smooth; both outer faces convex; both raised seed chambers not torulose. Fruit margin the same thickness; both more or less of equal length; constricted along both margins; without sulcus; plain. not folded; margin entire 180 degrees from base of Fruit wings absent. Fruit nonstipitate. Fruit indehiscent. radicle; similar at apex; partially concealing radicle; Replum invisible. Loment indehiscent. Loment seg- entire over radicle; without lobes; with the interface ments (articles) inconspicuous; 3.5Ð4 mm long; widest division terminating at base of radicle; without margins across seed area; with all essentially similar in shape; recessed; yellow or green; inner face flat; glabrous elliptic. Epicarp dull; monochrome; reddish brown; around base of radicle. Embryonic axis deflexed; pubescent and indurate; with hairs appressed or erect; oblique to length of seed; without a joint evident with 1 type of pubescence; velutinous; with pubescence between the radicle and the cotyledons. Radicle linear; gray; with pubescence uniformly distributed; with deflexed and parallel to cotyledon length; centered simple hairs; stiff; with hair bases plain; antrorse; between cotyledons; less than 1/2 length of cotyledons. straight at apex; eglandular; without spines; not smooth; Plumule moderately developed; glabrous. with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; Distribution: Tropical Asia to Australia. surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp Notes: Ohashi (1997a) corrected the name of this genus glossy; monochrome; tan; smooth; septate or from Dicerma A.-P. de Candolle to Aphyllodium and nonseptate; with septa thicker than paper, firm; with defined its species more narrowly so that it has four septa eglandular; chartaceous; not exfoliating; remain- species, the count we used. He (Ohashi 1973) also ing fused to mesocarp and epicarp; entire. Seeds 2 or 1; provided excellent drawings of fruits and seeds of the length parallel with fruit length; neither overlapping nor three species when treated as Dicerma. touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened or triangular; straight. Aril present (easily knocked off); dry; rim-aril; greenish tan or green. Aphyllum: A. biarticulatum (C. Linnaeus) F. Gagnepain (AÐE). A, Fruits (1- and 2-seeded: 1-seeded with calyx Seed 1.2Ð1.5 ϫ 1Ð1.2 ϫ 1Ð1.1 mm; not overgrown; not and petal) (ϫ 6.8); B, seeds (ϫ 10); CÐD, testa (ϫ 50, angular; asymmetrical; oblong; compressed; with ϫ 1000); E, embryos (ϫ 10). surface smooth; with or without visible radicle and

651 Genus: Desmodium A.N. Desvaux greenish); with brown (reddish) or purple overlay; glabrate or glabrous; with hairs erect or appressed; with Phylogenetic Number: 11.09. 1 or 2 types of pubescence; pilose, puberulent, or velutinous; with pubescence golden, brown, yellow, or Tribe: Desmodieae. gray-brown; with golden hooked hairs and gray plain hairs; with pubescence uniformly distributed; with Subtribe: Desmodiinae. simple hairs (plain and hooked hairs mixed, plain hairs, or hooked hairs); pliable or stiff; with hair bases plain; Species Studied—Species in Genus: 69 spp.—ca. 300 spp. straight, retrorse, or antrorse; hooked or straight at apex; eglandular; without spines; not smooth; with Fruit a loment (or loment segment) or legume; unilocular; elevated features; veined or not veined; reticulately 0.8Ð5 ϫ 0.08Ð2.5 ϫ 0.02Ð0.09 cm; with persistent veined; not tuberculate; wrinkled; not exfoliating; calyx; with calyx shorter than fruit; without orifice without cracks. Mesocarp present or absent; thin; formed by curving of fruit or fruit segments; straight, surface not veined; 1-layered; without balsamic curved (or slightly curved), or contorted; not plicate or vesicles; without fibers; solid; chartaceous or coria- plicate (D. styracifolium (P. Osbeck) E.D. Merrill ceous. Endocarp glossy or dull; monochrome; tan; plicate when young and becoming straight with age); spongy; with hairs in longitudinal rows (D. auricomum not twisted or twisted; symmetrical or asymmetrical; R. Graham); septate; with septa thicker than paper, linear, circular (with notch at funiculus), dolabriform, firm; with septa eglandular; chartaceous; not exfoliat- irregular, or samaroid; when asymmetrical with 1 ing; remaining fused to mesocarp and epicarp; entire. straight and 1 curved suture, both sutures parallelly Seeds 1Ð12; length parallel with fruit length; neither curved, or both sutures unequally curved; narrowing in overlapping nor touching; in 1 series. Funiculus less several places; not inflated or inflated; flattened or than 0.5 mm long or measured; up to 4 mm long; of 1 compressed; without or with beak; with solid beak the length only; thick, flattened, or filiform; straight, same color and texture as fruit; rounded or short tapered curved, or S-curved. Aril present or absent; dry; rim-aril at apex; apex aligned or oblique with longitudinal axis or cupshaped (only D. microphyllum (C.P. Thunberg) of fruit; short tapered, tapered, or long tapered at base; A.-P. de Candolle); entire; covering less than 1/2 of base aligned with longitudinal axis of fruit; with the seed. apex and base uniform in texture; membranous, coriaceous, or chartaceous; seed chambers externally Seed 0.7Ð12.5 ϫ 0.5Ð5 ϫ 0.5Ð1 mm; not overgrown; not visible; with the raised seed chambers not torulose. angular; asymmetrical; D-shaped, oblong, quadrangu- Fruit margin constricted or not constricted; constricted lar, rectangular, reniform, or linear; flattened or on 1 margin and slightly constricted on the other compressed; with surface grooved (D. oojeinensis (W. margin, slightly constricted only on 1 margin, con- Roxburgh) H. Ohashi) or smooth; with grooves stricted only on 1 margin, or slightly constricted along longitudinal and oblique; without visible radicle and both margins; without sulcus; plain or embellished; cotyledon lobes; without hilar sinus; without umbo on with prickles (at least stiff, usually hooked hairs). Fruit seed faces. Testa not adhering to endocarp; dull or wing 1; up to 10 mm wide; samaroid; on 1 suture. Fruit glossy; not modified by a bloom; colored; monochrome substipitate or stipitate; with the stipe 1Ð20 mm long. or mottled and streaked; with frequent mottles; with Fruit indehiscent or with all layers dehiscing (tardily); frequent streaks; dark to light reddish to brown, tan, splitting along suture. Dehiscence of valves along 1 green, or olive (green); with brown (dark reddish) suture; medial and up and down; passive. Replum overlay; glabrous; smooth or not smooth; with elevated invisible (most spp.) or visible (at least 1 sp. in Panama or recessed features; faintly wrinkled; pitted with small (Schubert 1980)). Loment indehiscent (most spp.) or separate pits; chartaceous or coriaceous. Fracture lines dehiscing along 1 suture (few spp., especially spp. in absent. Rim absent. Wings absent. Raphe not visible. subtribe Sagotia). Loment segments (articles) incon- Hilum visible or fully concealed; concealed by funicu- spicuous; 2Ð30 mm long; widest across seed area; with lar remnant; with or without faboid split; with the lips all essentially similar in shape; D-shaped, triangular, of the faboid split the same color as the rest of the curved, quadrangular, or circular (with notch at hilum; punctiform or larger than punctiform; up to 1 funiculus more or less dolobriform). Epicarp dull; mm long; with curved or straight outline; circular, monochrome or multicolored; mottled; brown (various oblong, or linear; marginal according to radicle tip or shades and combinations, especially reddish) or tan (to between cotyledon and radicle lobe; recessed; within

652 rim or not within corona, halo, or rim. Hilum rim color darker than testa. Lens discernible; less than 0.5 mm in length or equal to or greater than 0.5 mm in length; up to 0.5 mm long; with margins straight or curved; circular, oblong, elliptic, or 2 circular mounds separated by groove; not in groove of raphe; confluent with hilum; flush; dissimilar color from testa; darker than testa; reddish brown, black, or yellow (reddish); not within corona, halo, or rim. Endosperm present or absent; thin; covering entire embryo; adnate to testa or embryo. Cotyledons smooth or not smooth; both outer faces convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing radicle; entire over radicle; without lobes; with the interface division terminating at base of radicle; without margins recessed; yellow, tan, or green; inner face flat; glabrous around base of radicle. Embry- onic axis deflexed; oblique to length of seed; without a joint evident between the radicle and the cotyledons. Radicle linear; deflexed and parallel to cotyledon length; centered between cotyledons; less than 1/2 length of cotyledons. Plumule moderately developed, well developed, or rudimentary; glabrous.

Notes: Several regional or country treatments were reviewed: Schubert (1980), Panama; Oliveira (1983, 1990), Brazil; Verdcourt (1974), Zambia; Schubert (1952, 1954), Congo; Ohashi (1973), Asia; and Ohashi (1988), Hawaii. Verdcourt (1977) synonymized Papilionopsis C.G.G.J. van Steenis with Desmodium, and Ohashi (1982b) transferred Murtonia W.G. Craib to Desmodium as a subgenus. Although most species of Desmodium have indehiscent lomentaceous fruits, a few species have dehiscent loments, especially in the subgenus Sagatia (O.P. Swartz) A.-P. de Candolle, D. heterophyllum (C.L. von Willdenow) A.-P. de Candolle and D. microphyllum (C.P. Thunberg) A.-P. de Candolle. Desmodium auricomum loments have a longitudinal band of dense hairs on the lateral surfaces. All species of Desmodium except one, D. microphyllum, have small, dry rim-arils (Ohashi 1973); D. microphyllum has small, dry, cupshaped arils (Ohashi 1973).

653 Desmodium: D. uncinatum (N. von Jacquin) A.-P. de Candolle (CÐE), D. spp. (AÐB). A, Fruits, partial fruits, and articles (ϫ 1.9); B, seeds (ϫ 3.4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

655 Genus: Codariocalyx J.C. Hasskarl pressed; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; with umbo on Phylogenetic Number: 11.10. seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome or Tribe: Desmodieae. mottled and streaked; with frequent mottles; with frequent streaks; reddish brown; with black overlay; Subtribe: Desmodiinae. glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Species Studied—Species in Genus: 2 spp.—2 spp. visible or partially concealed; concealed by aril; with faboid split; with the lips of the faboid split the same Fruit a legume (resembling a loment); unilocular; 0.7Ð5 ϫ color as the rest of the hilum; larger than punctiform; 1 0.3Ð0.7 ϫ 0.18Ð0.2 cm; with persistent calyx; with mm long; with straight outline; oblong; marginal calyx shorter than fruit; without orifice formed by according to radicle tip; recessed; not within corona, curving of fruit or fruit segments; curved (or slightly halo, or rim. Lens not discernible. Endosperm thin; curved); not plicate; not twisted; asymmetrical; linear covering entire embryo; adnate to testa. Cotyledons (with toothed lower suture); when asymmetrical with 1 smooth; both outer faces convex; both the same straight and 1 curved suture or both sutures nearly thickness; both more or less of equal length; not folded; straight; narrowing in several places, resembling margin entire 180 degrees from base of radicle; similar Desmodium (11.09) fruit (false loment); not inflated; at apex; not concealing radicle; entire over radicle; flattened; without beak; rounded at apex; apex oblique without lobes; with the interface division terminating at or aligned with longitudinal axis of fruit; short tapered base of radicle; without margins recessed; yellow; inner or rounded at base; base oblique with longitudinal axis face flat; glabrous around base of radicle. Embryonic of fruit; with the apex and base uniform in texture; axis deflexed; oblique to length of seed; without a joint coriaceous; seed chambers externally visible; with the evident between the radicle and the cotyledons. Radicle raised seed chambers torulose. Fruit margin constricted bulbose; deflexed and parallel to cotyledon length; only on 1 margin; without sulcus; plain. Fruit wings centered between cotyledons; less than 1/2 length of absent. Fruit nonstipitate. Fruit with all layers dehisc- cotyledons. Plumule moderately developed; glabrous. ing; splitting along suture (lower). Dehiscence of valves along 1 suture (lower margin); passive. Replum Distribution: Southeastern Asia and tropical Australia. invisible. Epicarp dull; monochrome; dark reddish brown; pubescent and indurate; with hairs erect or Notes: Ohashi (1973) monographed the two species and appressed; with 1 type of pubescence; pilose or provided fruit and seed illustrations. The multiseeded tomentose; with pubescence golden (hook tipped) and legumes of Codariocalyx have persistent septa between gray (plain tipped); with pubescence uniformly distrib- the seed compartments, so that the fruit appears to be a uted; with simple hairs (shorter gray plain tipped, loment. It dehisces along the lower suture releasing the longer yellow hooked tipped); with hair bases plain; seeds from the individual seed compartments. The eglandular; without spines; smooth; not veined; not dehisced fruit is a line of open individual seed compart- tuberculate; not exfoliating; without cracks. Mesocarp ments which continue to be joined by the upper suture. thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull or glossy; monochrome; reddish tan or brown (reddish); smooth; septate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð13; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened or triangular; straight. Aril fleshy; cupshaped (with undulate margin); covering less than 1/2 of seed; tan. Codariocalyx: C. gyroides (W. Roxburgh ex J.H.F. Link) J.K. Hasskarl (CÐE), C. spp. (AÐB). A, Fruits (ϫ 2); B, Seed 2.5Ð4.5 ϫ 2.5Ð4 ϫ 1.5Ð1.7 mm; not overgrown; not seeds (ϫ 7.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos angular; asymmetrical; elliptic or reniform; com- (ϫ 8).

657 Genus: Pseudarthria R. Wight & G.A.W. Arnott with visible radicle and cotyledon lobes; without or with external groove (faintly) between radicle and Phylogenetic Number: 11.11. cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar Tribe: Desmodieae. sinus; with umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; Subtribe: Desmodiinae. monochrome; reddish brown, tan (to greenish or reddish), black (P. viscida), or orange (seeds have been Species Studied—Species in Genus: 3 spp.—4–6 spp. described as orange); glabrous; smooth (P. hookeri and P. fagifolia) or not smooth (P. viscida); with elevated Fruit a legume; unilocular; 1.3Ð2 ϫ 0.3Ð0.5 ϫ 0.1Ð0.13 cm; features; shagreen; coriaceous. Fracture lines absent. with persistent calyx; with calyx shorter than fruit; Rim absent. Wings absent. Raphe not visible. Hilum without orifice formed by curving of fruit or fruit visible; with faboid split; with the lips of the faboid segments; straight or curved (slightly); not plicate; not split the same color as the rest of the hilum; punctiform; twisted; asymmetrical; oblong; when asymmetrical with marginal according to radicle tip or between cotyledon both sutures nearly straight; not inflated; flattened; with and radicle lobe; barely recessed; not within corona, or without beak; straight; with solid beak the same halo, or rim or within halo (faint). Hilum halo color of color and texture as fruit; rounded at apex; apex aligned testa. Lens not discernible or discernible (faintly); less with longitudinal axis of fruit; short tapered at base; than 0.5 mm in length; with margins straight or curved; base oblique or aligned with longitudinal axis of fruit; somewhat linear or triangular; not in groove of raphe; with the apex and base uniform in texture; membra- confluent with hilum; flush; similar color as or dissimi- nous; seed chambers externally invisible or visible lar color from testa; darker than testa; reddish brown or (faintly); with the raised seed chambers not torulose. tan (greenish); not within corona, halo, or rim. En- Fruit margin not constricted; without sulcus; plain. Fruit dosperm thin; covering entire embryo; adnate to wings absent. Fruit substipitate. Fruit with all layers embryo. Cotyledons smooth; both outer faces convex; dehiscing; splitting along suture. Dehiscence of valves both the same thickness; both more or less of equal along 1 suture; apical and down (assumed); active; with length; not folded; margin entire 180 degrees from base valves reflexing. Replum invisible. Epicarp dull; of radicle; similar at apex; not concealing radicle; entire monochrome; dark reddish brown; pubescent and over radicle; without lobes; with the interface division indurate; with 1 type of pubescence; puberulent; with terminating at base of radicle; without margins re- pubescence golden (P. viscida (C. Linnaeus) R. Wight cessed; tan; inner face flat; glabrous around base of & G.A.W. Arnott with hooked-tipped hairs) or gray (P. radicle. Embryonic axis deflexed; oblique to length of hookeri without hooked-tipped hairs or P. fagifoli J.G. seed; without a joint evident between the radicle and Baker with straight- and hooked-tipped hairs); with the cotyledons. Radicle linear; deflexed and parallel to pubescence uniformly distributed; with simple hairs; cotyledon length; centered between cotyledons; less pliable; with hair bases plain; eglandular; without than 1/2 length of cotyledons. Plumule moderately spines; not smooth; with elevated features; not veined developed; glabrous. or transversely veined relative to fruit length and reticulately veined; not tuberculate; not exfoliating; Distribution: Old World tropics. without cracks. Mesocarp thin; surface not veined; 1- layered; without balsamic vesicles; without fibers; solid; coriaceous (sub-). Endocarp dull; monochrome; reddish brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 5Ð8; length transverse or oblique to fruit length; touching; in 1 series. Funiculus measured; 10 mm long; of 1 length only; filiform; S-curved or curved. Aril dry; rim-aril; reddish brown. Pseudarthria: P. hookeri R. Wight & G.A.W. Arnott (CÐE), Seed 2.5 ϫ 1.5 ϫ 1 mm; not overgrown; not angular or P. spp. (AÐB). A, Fruits (ϫ 4); B, seeds (ϫ 8.8); CÐD, angular; asymmetrical; reniform, oblong, triangular, or testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10). irregular; compressed; with surface smooth; without or

659 Genus: Pycnospora R. Brown ex R. Wight & G.A.W. without hilar sinus; without umbo on seed faces. Testa Arnott not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; Phylogenetic Number: 11.12. smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with Tribe: Desmodieae. faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; marginal Subtribe: Desmodiinae. according to radicle tip; barely recessed; within halo. Hilum halo color lighter than testa. Lens discernible; Species Studied—Species in Genus: 1 sp.—1 sp. less than 0.5 mm in length; with margins straight or curved; oblong; not in groove of raphe; adjacent to Fruit a legume; unilocular; 1Ð1.3 ϫ 0.5Ð0.6 ϫ 0.5 cm; with hilum; 0.1 mm from hilum; flush; dissimilar color from persistent calyx; with calyx shorter than fruit; without testa; darker than testa; black; not within corona, halo, orifice formed by curving of fruit or fruit segments; or rim. Endosperm thin; covering entire embryo; adnate straight; not plicate; not twisted; asymmetrical; oblong; to testa. Cotyledons smooth; both outer faces convex; when asymmetrical with 1 straight and 1 curved suture both the same thickness; both more or less of equal or both sutures nearly straight; widest near middle or length; not folded; margin entire 180 degrees from base D-shaped; inflated; terete; without or with beak; of radicle; similar at apex; not concealing radicle; entire straight or hooked; with solid beak the same color and over radicle; without lobes; with the interface division texture as fruit; rounded at apex; apex aligned with terminating at base of radicle; without margins re- longitudinal axis of fruit; rounded at base; base aligned cessed; tan; inner face flat; glabrous around base of or oblique with longitudinal axis of fruit; with the apex radicle. Embryonic axis deflexed; oblique to length of and base uniform in texture; membranous; seed seed; without a joint evident between the radicle and chambers externally invisible. Fruit margin not the cotyledons. Radicle linear; deflexed and parallel to constricted; without sulcus; plain. Fruit wings absent. cotyledon length; centered between cotyledons; less Fruit substipitate. Fruit with all layers dehiscing; than 1/2 length of cotyledons. Plumule moderately splitting along suture. Dehiscence of valves along 1 developed; glabrous. suture; apical and down (assumed); passive. Replum invisible. Epicarp dull; monochrome or multicolored; Distribution: Tropical Africa, India to Philippines, New mottled (large areas); blackish brown, tan (blackish), or Guinea, and northern Australia. black; with black overlay; pubescent and indurate; with 2 types of pubescence; puberulent; with pubescence golden or gray (and some hooked); with erect golden hairs and white hooked or not hooked hairs; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; transversely veined relative to fruit length and reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp very thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; chartaceous. Endocarp glossy; monochrome; blackish brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 10; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril dry; rim-aril; dark reddish brown. Pycnospora: P. lutescens (J.L.M. Poiret) A.K. Schindler Seed 1Ð1.3 ϫ 0.8Ð1 ϫ 0.5 mm; not overgrown; not angular; (AÐE). A, Fruits (dehisced and closed) (ϫ 4.6); B, asymmetrical; reniform; compressed; with surface seeds (ϫ 9.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos smooth; without visible radicle and cotyledon lobes; (ϫ 25).

661 Genus: Tadehagi H. Ohashi Seed 2.5Ð3.7 ϫ 2Ð2.5 ϫ 0.7Ð1.5 mm; not overgrown; angular or not angular; asymmetrical; reniform or Phylogenetic Number:11.13. irregular; compressed; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; Tribe: Desmodieae. without or with umbo on seed faces; with umbo on 1 face of seed. Testa not adhering to endocarp; dull; not Subtribe: Desmodiinae. modified by a bloom; colored; monochrome; reddish brown, tan (to reddish or greenish), yellow, green, or Species Studied—Species in Genus: 1 sp.—4 spp. orange; glabrous; smooth; coriaceous. Fracture lines present (easier to see on lighter colored seeds) or Fruit a loment (or loment segment); 2.5Ð4.7 ϫ 0.45Ð0.7 ϫ absent; reticulate. Rim absent. Wings absent. Raphe not 0.07Ð0.09 cm; with persistent calyx; with calyx shorter visible. Hilum visible; with faboid split; with the lips of than fruit; without orifice formed by curving of fruit or the faboid split the same color as the rest of the hilum; fruit segments; straight or curved (slightly); not plicate; larger than punctiform; 0.5 mm long; with curved not twisted; asymmetrical; moniliform; when asym- outline; circular; marginal according to radicle tip or metrical with 1 straight and 1 curved suture or both between cotyledon and radicle lobe; recessed; not sutures parallelly curved; narrowing in several places, within corona, halo, or rim. Lens discernible; less than resembling Desmodium (11.9) fruit; not inflated; 0.5 mm in length; with margins curved; 2 oblong flattened; with beak; straight; with solid beak the same mounds separated by groove; not in groove of raphe; color and texture as fruit; rounded at apex; apex aligned confluent with hilum; mounded; dissimilar color from or oblique with longitudinal axis of fruit; short tapered testa; darker than testa; black; not within corona, halo, at base; base aligned or oblique with longitudinal axis or rim. Endosperm thick; covering entire embryo; of fruit; with the apex and base uniform in texture; adnate to testa. Cotyledons not smooth; both outer faces membranous; seed chambers externally visible; with the convex; both the same thickness; both more or less of raised seed chambers not torulose. Fruit margin equal length; with both folded; sufficiently folded for constricted on 1 margin and slightly constricted on the inner face to touch itself; portions of inner folded face other margin; without sulcus; plain. Fruit wings absent. unequal; margin entire 180 degrees from base of Fruit stipitate, substipitate, or nonstipitate; with the radicle; similar at apex; not concealing radicle; entire stipe up to 14 mm long. Fruit indehiscent. Replum over radicle; without or with lobes; with lobes not invisible. Loment indehiscent. Loment segments touching; without basal groin formed by lobes; with the (articles) inconspicuous; 2.7Ð10 mm long; widest interface division terminating at base of radicle; without across seed area; with upper and lower 1 different shape margins recessed; reddish brown or tan; inner face flat; than middle ones; D-shaped, rectangular, or quadrangu- glabrous around base of radicle. Embryonic axis lar. Epicarp dull; monochrome; brown to reddish or deflexed; oblique to length of seed; with a joint evident dark reddish brown; pubescent and indurate or glabrous between the radicle and the cotyledons. Radicle linear; (except for scattered hairs on suture); with 1 or 2 types deflexed and parallel to cotyledon length; centered of pubescence; pilose; with pubescence yellow; with between cotyledons; less than 1/2 length of cotyledons. long and short yellow hairs intermixed; with pubes- Plumule moderately developed; glabrous. cence uniformly distributed; with simple hairs; longer stiff and pliable (shorter); with hair bases plain; Distribution: India to southern China to New Caledonia and antrorse; straight at apex; eglandular; without spines; northern Australia. not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Notes: Ohashi et al. (1981) noted that Tadehagi integrates Mesocarp absent. Endocarp dull; monochrome; reddish with Droogmansia (11.14) and provided fruit and seed brown; smooth; septate; with septa thicker than paper, drawings of Tadehagi. Ohashi (1982a) and Maiti and firm; with septa eglandular; chartaceous; not exfoliat- Ghosh (1997) reevaluated T. triquetrum subsp. rodgeri ing; remaining fused to epicarp; entire. Seeds 5Ð7; (A.K. Schindler) H. Ohashi, and raised it to the rank of length parallel with fruit length; neither overlapping nor species, T. rogeri (A.K. Schindler) H. Ohashi. There- touching; in 1 series. Funiculus measured; 2 mm long; fore, there are four species in the genus. of 1 length only; filiform; curved or straight. Aril dry; rim-aril; tan.

662 663 Tadehagi: T. triquetrum (C. Linnaeus) H. Ohashi (AÐE). A, Fruits (entire but separated and two united articles) (ϫ 3.3); B, seeds (ϫ 7.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 11).

665 Genus: Droogmansia E.A.J. De Wildeman Seed 2.5Ð4.5 ϫ 2.5Ð3 ϫ 1.3Ð1.5 mm; not overgrown; not angular; asymmetrical; reniform or oblong; flattened; Phylogenetic Number: 11.14. with surface smooth; with or without visible radicle and cotyledon lobes; without external groove between Tribe: Desmodieae. radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to Subtribe: Desmodiinae. endocarp; glossy; not modified by a bloom; colored; monochrome (but note hilar halo); dark reddish brown; Species Studied—Species in Genus: 2 spp.—ca. 10 spp. glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Fruit a loment (or loment segment); 3.3Ð4 ϫ 0.3Ð0.7 ϫ 0.7Ð visible; with faboid split; with the lips of the faboid 0.9 cm; with persistent calyx; with calyx shorter than split the same color as the rest of the hilum; larger than fruit; without orifice formed by curving of fruit or fruit punctiform; 0.5 mm long; with curved outline; elliptic; segments; slightly curved or straight; not plicate; not marginal according to radicle tip or between cotyledon twisted; asymmetrical; moniliform; when asymmetrical and radicle lobe; flush; within halo. Hilum halo color with 1 straight and 1 curved suture or both sutures darker (and irregular in outline) than testa. Lens parallelly curved; narrowing in several places, resem- discernible; less than 0.5 mm in length or equal to or bling Desmodium (11.09) fruit; not inflated; flattened; greater than 0.5 mm in length; 0.3Ð0.8 mm long; with with beak; declined; with solid beak the same color and margins straight or curved; irregular, linear, or wedge- texture as fruit; rounded at apex; apex aligned or shaped; not in groove of raphe; confluent with hilum; oblique with longitudinal axis of fruit; rounded at base; mounded or recessed; dissimilar color from testa; base aligned or oblique with longitudinal axis of fruit; darker than testa; dark reddish brown; not within with the apex and base uniform in texture; coriaceous; corona, halo, or rim. Endosperm thin; covering entire seed chambers externally visible; with the raised seed embryo; adnate to testa. Cotyledons smooth; both outer chambers not torulose. Fruit margin constricted or faces convex; both the same thickness; both more or slightly constricted along both margins; without sulcus; less of equal length; not folded; margin entire 180 plain. Fruit wings absent. Fruit stipitate or substipitate; degrees from base of radicle; similar at apex; not with the stipe up to 70 mm long. Fruit indehiscent. concealing radicle; entire over radicle; without lobes; Replum invisible. Loment indehiscent. Loment seg- with the interface division terminating at base of ments (articles) inconspicuous; 5Ð13 mm long; widest radicle; without margins recessed; tan; inner face flat; across seed area; with upper or lower 1 different shape glabrous around base of radicle. Embryonic axis than middle ones; D-shaped or circular. Epicarp dull; deflexed; oblique to length of seed; without a joint monochrome; dark reddish to light brown or tan; evident between the radicle and the cotyledons. Radicle pubescent and indurate; with 1 type of pubescence; linear; deflexed and parallel to cotyledon length; sericeous; with pubescence gray; with pubescence centered between cotyledons; less than 1/2 length of uniformly distributed; with simple hairs; pliable; with cotyledons. Plumule moderately developed; glabrous. hair bases plain; eglandular; without spines; not smooth; with elevated features; veined or not veined; Distribution: South-central and West Africa and 1 sp. in reticulately veined; not tuberculate; not exfoliating; Indochina. without cracks. Mesocarp thin; surface not veined; 1- layered; without balsamic vesicles; without fibers; Notes: Ohashi et al. (1981) noted that the number of species solid; coriaceous. Endocarp dull; monochrome; reddish is “usually estimated to be about 30 species, but a brown; smooth; septate; with septa thicker than paper, single species complex in south-central Africa with firm; with septa eglandular; chartaceous; not exfoliat- perhaps a few distinct relatives, then four species in ing; remaining fused to mesocarp and epicarp; entire. West Africa intergrading with Tadehagi” (11.13). We Seeds 1Ð5; length parallel with fruit length; neither agree with Ohashi et al. (1981) and Verdcourt (1970b, overlapping nor touching; in 1 series. Funiculus 1974), but not Lock (1989), that there are only a few measured; up to 1 mm long; of 1 length only; flattened; species in Africa, Lock’s recent list of 23 species in straight. Aril absent. Africa notwithstanding. Schubert (1952) recognized nearly 30 species in Africa and noted that most of her new species were collected in flower. Our count of ca.

666 10 species is an estimate that may or may not be accurate. Regardless of the species counts, our fruit and seed material was quite limited.

667 Droogmansia: D. pteropus (J.G. Baker) E.A.J. De Wildemann var. whytei (A.K. Schindler) B. Verdcourt (CÐE), D. spp. (AÐB). A, Fruit and article (ϫ 2.8); B, seeds (ϫ 6.5); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

669 Genus: Mecopus J.J. Bennett punctiform; marginal according to radicle tip; flush; within halo. Hilum halo color of testa. Lens discernible; Phylogenetic Number: 11.15. less than 0.5 mm in length; with margins straight or curved; oblong; not in groove of raphe; confluent with Tribe: Desmodieae. hilum; mounded; dissimilar color from testa; darker than testa; darker reddish brown; not within corona, Subtribe: Desmodiinae. halo, or rim. Endosperm thick; covering entire embryo; adnate to embryo. Cotyledons smooth; both outer faces Species Studied—Species in Genus: 1 sp.—1 sp. convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 degrees Fruit a legume; unilocular; 0.27 ϫ 0.2 ϫ 0.1 cm; with from base of radicle; similar at apex; not concealing deciduous or persistent calyx; with calyx shorter than radicle; entire over radicle; without lobes; with the fruit; without orifice formed by curving of fruit or fruit interface division terminating at base of radicle; without segments; straight; not plicate; not twisted; asymmetri- margins recessed; white; inner face flat; glabrous cal; when asymmetrical with 1 straight and 1 curved around base of radicle. Embryonic axis deflexed; suture; widest near middle or D-shaped; not inflated; oblique to length of seed; without a joint evident compressed; without beak; short tapered at apex; apex between the radicle and the cotyledons. Radicle linear; oblique with longitudinal axis of fruit; short tapered at deflexed and parallel to cotyledon length; centered base; base aligned with longitudinal axis of fruit; with between cotyledons; less than 1/2 length of cotyledons. the apex and base uniform in texture; chartaceous; seed Plumule moderately developed; glabrous. chambers externally visible; with the raised seed chambers not torulose. Fruit margin not constricted; Distribution: India to southern China and Malay Islands. without sulcus; plain. Fruit wings absent. Fruit stipitate; with the stipe 50–60 mm long (and reflexed into “bird Notes: The seeds dehisce from the one-seeded fruits while cage” where fruits are “caught”). Fruit with all layers on the plants. Apparently, the “bird-cage” heads fall dehiscing; splitting along suture. Dehiscence of valves intact, and the seeds are shed as the heads are trans- along 1 suture; apical and down; passive. Replum ported on animal fur. invisible. Epicarp dull; monochrome; reddish brown or tan (reddish); pubescent and indurate; with 1 type of pubescence; puberulent; with pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; reddish brown or tan (reddish); smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching (if 1-seeded); in 1 series. Funiculus less than 0.5 mm long; of 1 length only; straight. Aril dry; rim-aril; reddish brown.

Seed 1.7Ð2 ϫ 1.1Ð1.2 ϫ 0.6Ð0.7 mm; not overgrown; not angular; asymmetrical; reniform (sub-); compressed; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown or tan; glabrous; smooth; coriaceous. Fracture Mecopus: M. nidulans J.J. Bennett (AÐF). A, “bird cage” lines absent. Rim absent. Wings absent. Raphe not (see Notes) (ϫ 2.9); B, fruits (ϫ 10); C, seeds (ϫ 10); visible. Hilum visible; with faboid split; with the lips of DÐE, testa (ϫ 50, ϫ 1000); F, embryos (ϫ 20). the faboid split the same color as the rest of the hilum;

671 Genus: Uraria A.N. Desvaux Seed 1.8Ð2.5 ϫ 1.5Ð2 ϫ 0.7Ð0.8 mm; not overgrown; not angular or angular; asymmetrical; reniform, elliptic, Phylogenetic Number: 11.16. mitaform, or triangular; compressed; with surface smooth; without visible radicle and cotyledon lobes; Tribe: Desmodieae. without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull or glossy; not modified Subtribe: Desmodiinae. by a bloom; colored; monochrome; yellowish to reddish brown or tan; glabrous; smooth; coriaceous. Fracture Species Studied—Species in Genus: 8 spp.—20 spp. lines present or absent; reticulate. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid Fruit a loment (or loment segment); 0.3Ð0.9 ϫ 0.2Ð0.4 ϫ split; with the lips of the faboid split the same color as 0.1Ð0.2 cm; with persistent calyx; with calyx shorter the rest of the hilum; punctiform or larger than puncti- than fruit; without orifice formed by curving of fruit or form; to 0.5 mm long; with curved outline; oval; fruit segments; straight or curved (or slightly curved); marginal according to radicle tip or between cotyledon plicate or not plicate; not twisted; asymmetrical; and radicle lobe; slightly recessed; within halo (faint). oblong; when asymmetrical with both sutures parallelly Hilum halo color darker (slightly) than testa. Lens curved or 1 straight and 1 curved suture; narrowing in discernible; less than 0.5 mm in length; with margins several places, resembling Desmodium (11.9) fruit; not straight or curved; rectangular or circular; not in groove inflated; compressed or terete; without beak; rounded of raphe; confluent with hilum or adjacent to hilum; 0.1 at apex; apex oblique with longitudinal axis of fruit; mm from hilum; mounded; dissimilar color from testa; short tapered or rounded at base; base aligned with darker than testa; reddish brown or black; not within longitudinal axis of fruit; with the apex and base corona, halo, or rim. Endosperm thin; covering entire uniform in texture; membranous; seed chambers embryo; adnate to testa or embryo. Cotyledons smooth; externally visible; with the raised seed chambers not both outer faces convex; both the same thickness; both torulose. Fruit margin constricted along both margins; more or less of equal length; not folded; margin entire without sulcus; plain. Fruit wings absent. Fruit 180 degrees from base of radicle; similar at apex; not substipitate. Fruit indehiscent. Replum invisible. concealing radicle; entire over radicle; without lobes; Loment indehiscent. Loment segments (articles) with the interface division terminating at base of inconspicuous; 2.5Ð3.5 mm long; widest across seed radicle; without margins recessed; tan or yellow; inner area; with all essentially similar in shape; ovate. face flat; glabrous around base of radicle. Embryonic Epicarp dull or glossy; monochrome; brown, tan, black, axis deflexed; oblique to length of seed; without a joint or gray; glabrous (widely scattered hairs may be on evident between the radicle and the cotyledons. Radicle sutures) or pubescent and indurate; with 1 type of linear or bulbose; deflexed and parallel to cotyledon pubescence; puberulent; with pubescence golden or length; centered between cotyledons; less than 1/2 gray (with straight-tipped or straight- and hooked- length of cotyledons. Plumule moderately developed; tipped hairs); with pubescence uniformly distributed; glabrous. with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated Distribution: Old World tropics. features; reticulately veined; not tuberculate; not exfoliating or exfoliating (U. picta leaving a shiny gray Notes: Urariopsis Schindler (11.16) in Ohashi et al. (1981) mesocarp); without cracks. Mesocarp present or absent; is a synonym of Uraria (Haas et al. 1980). Urariopsis thin to very thin; surface not veined; 1-layered; without cordifolia (C.F.W. Wallroth) A.K. Schindler is based balsamic vesicles; without fibers; solid; chartaceous. on U. cordifolia C.F.W. Wallroth. Jha and Pandey Endocarp dull or glossy; monochrome; brown (to (1988) studied the seeds of Uraria and Alysicarpus grayish); smooth; septate; with septa thicker than (11.16 and 11.18). paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; remaining fused to epicarp; entire. Seeds 2Ð6; length parallel or transverse to fruit length; neither overlap- Uraria: U. picta (N. von Jacquin) A.N. Desvaux ex A.-P. de ping nor touching; in 1 series. Funiculus less than 0.5 Candolle (CÐE), U. spp. (AÐB). A, Fruits with calyxes mm long; of 1 length only; more or less thick; straight. and articles (ϫ 4.7); B, seeds (ϫ 8.9); CÐD, testa Aril dry; rim-aril; reddish brown. (ϫ 50, ϫ 1000); E, embryos (ϫ 12).

673 Genus: Christia C. Moench monochrome; dark to light reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Phylogenetic Number: 11.17. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same Tribe: Desmodieae. color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; flush; within rim. Hilum Subtribe: Desmodiinae. rim color darker than testa. Lens discernible; less than 0.5 mm in length; with margins curved; circular or Species Studied—Species in Genus: 2 spp.—ca. 10 spp. elliptic; not in groove of raphe; confluent with hilum; slightly mounded; dissimilar color from testa; darker Fruit a loment (or loment segment); 0.6 ϫ 0.15 ϫ 0.1 cm; than testa; black; not within corona, halo, or rim. with persistent calyx; with calyx longer or shorter than Endosperm thin; covering entire embryo; adnate to fruit; without orifice formed by curving of fruit or fruit testa. Cotyledons not smooth; both outer faces convex; segments; S-curved; plicate; not twisted; asymmetrical; both the same thickness; both more or less of equal irregular; when asymmetrical with 1 straight and 1 length; not folded; margin entire 180 degrees from base curved suture or both sutures unequally curved; of radicle; similar at apex; not concealing radicle; entire narrowing in several places, resembling Desmodium over radicle; without lobes; with the interface division (11.09) fruit (though folded); not inflated; flattened; terminating at base of radicle; without margins re- without beak; truncate at apex; apex aligned with cessed; tan; inner face flat; glabrous around base of longitudinal axis of fruit; truncate at base; base aligned radicle. Embryonic axis deflexed; oblique to length of with longitudinal axis of fruit; with the apex and base seed; without a joint evident between the radicle and uniform in texture; chartaceous (easily eroding); seed the cotyledons. Radicle linear; lobe tip straight; chambers externally visible; with the raised seed deflexed and parallel to cotyledon length; centered chambers not torulose. Fruit margin constricted only on between cotyledons; less than 1/2 length of cotyledons. 1 margin; without sulcus; plain. Fruit wings absent. Plumule moderately developed; glabrous. Fruit substipitate or nonstipitate. Fruit indehiscent. Replum invisible. Loment indehiscent. Loment seg- Distribution: Tropical and subtropical Asia and Australasia. ments (articles) inconspicuous; 2 mm long; widest across seed area; with all essentially similar in shape; D-shaped. Epicarp dull; monochrome; gray, brown, or black; glabrous or pubescent and indurate; with 1 type of pubescence; puberulent; with pubescence gray (with hooked tips); with pubescence uniformly distributed; with simple hairs (with hooked tips); pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; reddish brown or white; smooth; septate; with septa thin (tissue paper-like), flexible; with septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð 3; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 1.6Ð2.5 ϫ 1.2Ð1.8 ϫ 0.6Ð0.9 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with or without visible radicle and cotyledon lobes; without external groove between Christia: C. obcordata (J.L.M. Poiret) R.C. Bakhhuizen van radicle and cotyledon lobes; without hilar sinus; den Brink (CÐE), C. spp. (AÐB). A, Fruit in calyx and without umbo on seed faces. Testa not adhering to articles (ϫ 5.5); B, seeds (ϫ 10.5); CÐD, testa (ϫ 50, endocarp; glossy; not modified by a bloom; colored; ϫ 1000); E, embryos (ϫ 19).

675 Genus: Alysicarpus A.N. Desvaux Seed 1.5Ð2.5 ϫ 1.2Ð1.8 ϫ 0.8Ð1.2 mm; not overgrown; not angular; symmetrical; elliptic (and oblong), oblong, Phylogenetic Number: 11.18. rhombic (rounded), quadrangular, or circular (to sub-); slightly compressed; with surface smooth; without Tribe: Desmodieae. visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to Subtribe: Desmodiinae. endocarp; glossy or dull; not modified by a bloom; colored; monochrome or mottled (faint to conspicuous); Species Studied—Species in Genus: 11 spp.—25–30 spp. with frequent mottles; reddish brown, tan (and greenish- to reddish-tan), yellow, or black; with brown Fruit a loment (or loment segment) or legume; unilocular; (reddish) or purple overlay; glabrous; smooth; coria- 0.5Ð2.5 ϫ 0.1Ð0.3 ϫ 0.1Ð0.2 cm; with persistent or ceous. Fracture lines absent. Rim absent. Wings absent. deciduous calyx; with calyx longer than fruit; without Raphe not visible. Hilum fully concealed or visible; orifice formed by curving of fruit or fruit segments; concealed by funiculus; with faboid split; with the lips straight or curved (slightly); not plicate; not twisted; of the faboid split the same color as the rest of the asymmetrical or symmetrical; linear, moniliform, or hilum; punctiform; marginal according to radicle tip; oblong; when asymmetrical with 1 straight and 1 flush; within rim or within halo. Hilum halo color of curved suture; widest near middle or D-shaped or testa (essentially). Hilum rim color of testa. Lens narrowing in several places, resembling Desmodium discernible; equal to or greater than 0.5 mm in length; (11.09) fruit; not inflated; terete; without or with beak; 0.3 mm long; with margins straight or curved; linear; straight; with solid beak the same color and texture as more or less circular; not in groove of raphe; confluent fruit; rounded at apex; apex aligned with longitudinal with hilum; flush; dissimilar color from testa; darker axis of fruit; rounded at base; base aligned with than testa; dark reddish brown; not within corona, halo, longitudinal axis of fruit; with the apex and base or rim. Endosperm thin; covering entire embryo; adnate uniform in texture; membranous; seed chambers to testa or embryo. Cotyledons not smooth; both outer externally visible; with the raised seed chambers faces convex; both the same thickness; both more or torulose or not torulose. Fruit margin constricted or not less of equal length; not folded; margin entire 180 constricted; constricted along both margins; without degrees from base of radicle; similar at apex; partially sulcus; plain. Fruit wings absent. Fruit nonstipitate. concealing or not concealing radicle; entire over Fruit indehiscent. Replum invisible. Loment indehis- radicle; without or with lobes; with lobes not touching; cent or dehiscing along 1 suture (somewhat). Loment without basal groin formed by lobes; with the interface segments (articles) conspicuous; 1Ð4 mm long; widest division terminating at base of radicle; without margins across seed area; with all essentially similar in shape; recessed; yellow or white; inner face flat; glabrous rectangular or quadrangular. Epicarp dull; monochrome around base of radicle. Embryonic axis deflexed; or multicolored; bichrome; brown (to dirty-, greenish-, oblique to length of seed; without a joint evident dark-, or reddish-brown), tan, or black; with brown between the radicle and the cotyledons. Radicle linear; (reddish) overlay; pubescent and indurate or glabrous; deflexed and parallel to cotyledon length or width; with hairs erect or appressed; puberulent; with pubes- centered between cotyledons; 1/2 to nearly length of cence gray; with simple hairs (with hooked tips that are cotyledons. Plumule moderately developed; glabrous. easily broken off); pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated Distribution: Old World tropics with A. vaginalis (C. features; reticulately veined or transversely veined Linnaeus) A.-P. de Candolle a worldwide crop. relative to fruit length; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; Notes: Jha and Pandey (1988) studied seeds of Alysicarpus monochrome; brown; smooth; septate; with septa and Uraria (11.16). thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð8; length parallel or transverse to fruit length; neither overlapping nor touching; in 1 Alysicarpus: A. rugosus (C.L. von Willdenow) A.-P. de series. Funiculus less than 0.5 mm long; of 1 length Candolle (CÐE), A. spp. (AÐB). A, Fruits (with and only; filiform; straight. Aril absent. without calyx) and articles (ϫ 3.4); B, seeds (ϫ 8.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

677 Genus: Desmodiastrum (D. Prain) A. Pramanik & K. a bloom; colored; monochrome; dark brown or gray Thothathri (dark); glabrous; smooth; coriaceous (assumed). Fracture lines absent. Rim present. Wings absent. Phylogenetic Number: 11.18A. Distribution: India, Indochina, Indonesia. Tribe: Desmodieae. Notes: Pramanik and Thothathri (1986) raised Alysicarpus Subtribe: Desmodiinae. subgenus Desmodiastrum to the genus level. We were unable to obtain fruits and seeds and instead used Species Studied—Species in Genus: 1 sp.—2 spp. Pramanik and Thothathri’s text and the plate from Wight (1838) for our figure. Fruit a loment (or loment segment); 0.8Ð1.5 ϫ 0.35Ð0.5 cm; with persistent calyx; with calyx shorter or longer than fruit; without orifice formed by curving of fruit or fruit segments; straight; not plicate; not twisted; asymmetrical; moniliform; when asymmetrical with both sutures parallelly curved (nearly so) or 1 straight and 1 curved suture; narrowing in several places, resembling Desmodium (11.09) fruit; not inflated; flattened; with beak; nearly straight; with solid beak the same color and texture as fruit; short tapered or rounded at apex; apex aligned with longitudinal axis of fruit; rounded at base; base aligned with longitudinal axis of fruit; with the apex and base uniform in texture; membranous; seed chambers externally visible; with the raised seed chambers not torulose. Fruit margin constricted only on 1 margin; without sulcus; plain. Fruit nonstipitate. Fruit indehiscent. Replum invisible. Loment indehiscent. Loment segments (articles) inconspicuous; widest across seed area; with all essentially similar in shape; D-shaped. Epicarp dull; monochrome; brown (assumed); pubescent and indurate; with hairs appressed; with 1 type of pubescence; with pubescence uniformly distributed; with simple hairs (mixture of hooked and plain tipped hairs); pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp present or absent (assumed to be like Desmodium (11.09)). Endocarp septate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 3Ð6; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only. Aril absent.

Seed 1.2Ð2 ϫ 1.2Ð2 mm; not overgrown; not angular; asymmetrical; reniform or ovate; compressed; with surface smooth; with visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Desmodiastrum: D. belgaumense (W. Wight) A. Pramanik Testa not adhering to endocarp; glossy; not modified by & K. Thothathri (AÐB). A, Fruit (ϫ 4.5); B, seed (ϫ 10), both from Wight (1838).

679 Genus: Melliniella H.A.T. Harms cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between Phylogenetic Number: 11.19. radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo on seed faces. Testa not Tribe: Desmodieae. adhering to endocarp; glossy; not modified by a bloom; colored; monochrome; reddish brown or tan; glabrous; Subtribe: Desmodiinae. smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible (but Species Studied—Species in Genus: 1 sp.—1 sp. difficult to see because of size and color); with faboid split; with the lips of the faboid split the same color as Fruit a legume; unilocular; 1Ð1.3 ϫ 0.2 ϫ 0.5Ð0.7 cm; with the rest of the hilum; punctiform; between cotyledon persistent or deciduous calyx; with calyx shorter than and radicle lobe; flush; within rim. Hilum rim color of fruit; without orifice formed by curving of fruit or fruit testa. Lens discernible or not discernible; less than 0.5 segments; slightly curved or straight; not plicate; not mm in length; with margins curved; circular; not in twisted; asymmetrical; C-shaped or linear; when groove of raphe; confluent with hilum; mounded; asymmetrical with both sutures parallelly curved or dissimilar color from or similar color as testa; darker nearly straight; not inflated; compressed; with beak; than testa; black or brown (darker reddish); not within with solid beak the same color and texture as fruit; short corona, halo, or rim. Endosperm thin; covering entire tapered or tapered at apex; apex oblique or aligned with embryo; adnate to embryo. Cotyledons smooth; both longitudinal axis of fruit; tapered at base; base oblique outer faces convex; both the same thickness; both more or aligned with longitudinal axis of fruit; with the apex or less of equal length; not folded; margin entire 180 and base uniform in texture; membranous; seed degrees from base of radicle; similar at apex; not chambers externally visible; with the raised seed concealing radicle; entire over radicle; without lobes; chambers not torulose. Fruit margin not constricted; with the interface division terminating at base of without sulcus; plain. Fruit wings absent. Fruit radicle; without margins recessed; white; inner face substipitate or nonstipitate. Fruit with all layers flat; glabrous around base of radicle. Embryonic axis dehiscing; splitting along suture. Dehiscence of valves deflexed; oblique to length of seed; without a joint along 1 suture; apical and down; passive. Replum evident between the radicle and the cotyledons. Radicle invisible. Epicarp dull; monochrome; brown or tan; linear; deflexed and parallel to cotyledon length; pubescent and indurate; with 1 type of pubescence; centered between cotyledons; 1/2 to nearly length of puberulent; with pubescence gray; with pubescence cotyledons. Plumule rudimentary; glabrous. uniformly distributed; with simple hairs (with hooked tips); pliable; with hair bases plain; eglandular; without Distribution: Western tropical Africa. spines; not smooth; with elevated features; transversely veined relative to fruit length and reticulately veined; Notes: Harms (1914) founded the genus and provided fruit not tuberculate; not exfoliating; without cracks. and seed drawings. Ohashi et al. (1981) noted that Mesocarp thin; surface not veined; 1-layered; without Melliniella is “a minor segregate of Alysicarpus” balsamic vesicles; without fibers; solid; chartaceous. (11.18). Endocarp dull or glossy; monochrome; brown; smooth; septate or nonseptate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 5Ð8; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril present (but difficult to see) or absent; dry; rim- aril; reddish brown or tan.

Seed 1.2Ð1.3 ϫ 1.1Ð1.2 ϫ 0.6Ð0.7 mm; not overgrown; not angular; asymmetrical; mitaform or oblong; com- Melliniella: M. micrantha H.A.T. Harms (AÐE). A, Fruits pressed; with surface smooth; with visible radicle and with and without calyx (ϫ 6.4); B, seeds (ϫ 13.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 15).

681 Genus: Leptodesmia G. Bentham with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than puncti- Phylogenetic Number: 11.20. form; 0.5 mm long; with curved outline; elliptic; marginal according to radicle tip; flush; within rim. Tribe: Desmodieae. Hilum rim color of testa. Lens discernible (under aril); less than 0.5 mm in length; with margins curved; Subtribe: Desmodiinae. circular; not in groove of raphe; confluent with hilum; mounded; dissimilar color from testa; darker than testa; Species Studied—Species in Genus: 4 spp.—6 spp. black; not within corona, halo, or rim. Endosperm thin; covering entire embryo; adnate to testa. Cotyledons Fruit a legume; unilocular; 0.4Ð0.5 ϫ 0.25 ϫ 0.05 cm; with smooth; both outer faces convex; both the same persistent calyx; with calyx longer than fruit; without thickness; both more or less of equal length; not folded; orifice formed by curving of fruit or fruit segments; margin entire 180 degrees from base of radicle; similar straight; not plicate; not twisted; asymmetrical; oblong; at apex; not concealing radicle; entire over radicle; when asymmetrical with both sutures parallelly curved; without lobes; with the interface division terminating at not inflated; flattened; without beak; rounded at apex; base of radicle; without margins recessed; tan; inner apex oblique to right-angled with longitudinal axis of face flat; glabrous around base of radicle. Embryonic fruit; short tapered at base; base aligned with longitudi- axis deflexed; oblique to length of seed; without a joint nal axis of fruit; with the apex and base uniform in evident between the radicle and the cotyledons. Radicle texture; coriaceous; seed chambers externally visible. linear; deflexed and parallel to cotyledon length; Fruit margin not constricted; without sulcus; plain. Fruit centered between cotyledons; less than 1/2 length of wings absent. Fruit nonstipitate. Fruit with all layers cotyledons or 1/2 to nearly length of cotyledons. dehiscing; splitting along suture. Dehiscence of valves Plumule moderately developed; glabrous. along 1 suture; medial and up and down; active; with valves reflexing. Replum invisible. Epicarp dull; Distribution: Madagascar (5 spp.) and India (L. congesta). monochrome; reddish brown; pubescent and indurate; with 1 type of pubescence; puberulent; with pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1- layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; golden brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seed 1; length parallel with fruit length. Funiculus less than 0.5 mm long; flattened; straight. Aril fleshy; cupshaped; covering less than 1/2 of seed; reddish brown or cream.

Seed 2Ð2.5 ϫ 1Ð1.5 ϫ 0.5Ð0.8 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; mottled and streaked or monochrome; with frequent mottles; with frequent Leptodesmia: L. congesta G. Bentham ex J.G. Baker (CÐE), streaks; dark to lighter reddish brown, green, or olive; L. spp. (AÐB). A, Fruits with and without calyx (ϫ 8); with black overlay; glabrous; smooth; coriaceous. B, seeds (ϫ 10.9); CÐD, testa (ϫ 50, ϫ 1000); E, Fracture lines absent. Rim absent. Wings absent. Raphe embryos (ϫ 10). not visible. Hilum fully concealed; concealed by aril;

683 Genus: Eleiotis A.-P. de Candolle seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled Phylogenetic Number: 11.21. and streaked; with frequent mottles; very dark reddish brown, black, tan (greenish), or green; with purple Tribe: Desmodieae. overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Subtribe: Desmodiinae. Hilum visible; with faboid split; with the lips of the faboid split lighter colored than the rest of the hilum Species Studied—Species in Genus: 1 sp.—2 spp. and therefore conspicuous; larger than punctiform; 0.5 mm long; with straight outline; oblong; marginal Fruit a legume; unilocular; 0.6Ð0.65 ϫ 0.2Ð2.5 ϫ 0.1Ð0.15 according to radicle tip; flush; within halo. Hilum halo cm; with persistent or deciduous calyx; with calyx color darker than testa. Lens discernible; less than 0.5 shorter than fruit; without orifice formed by curving of mm in length or equal to or greater than 0.5 mm in fruit or fruit segments; straight; not plicate; not twisted; length; 0.5 mm long; with margins curved; circular; not asymmetrical; C-shaped; when asymmetrical with 1 in groove of raphe; confluent with hilum; mounded; straight and 1 curved suture or both sutures parallelly dissimilar color from or same color as testa; darker than curved; widest near middle or D-shaped; not inflated; testa; very dark reddish brown, blackish, or tan compressed; with or without beak; straight; with solid (greenish); not within corona, halo, or rim. Endosperm beak the same color and texture as fruit; short tapered at thin; covering entire embryo; adnate to testa. Cotyle- apex; apex aligned with longitudinal axis of fruit; short dons smooth; both outer faces convex; both the same tapered at base; base aligned with longitudinal axis of thickness; both more or less of equal length; not folded; fruit; with the apex and base uniform in texture; margin entire 180 degrees from base of radicle; similar membranous; seed chambers externally visible; with the at apex; completely concealing radicle; entire over raised seed chambers not torulose. Fruit margin not radicle; without lobes; with the interface division constricted; with sulcus; plain. Fruit wings absent. Fruit terminating at base of radicle; without margins re- nonstipitate. Fruit indehiscent. Replum invisible. cessed; reddish brown; inner face flat; glabrous around Epicarp dull; monochrome or multicolored; mottled and base of radicle. Embryonic axis deflexed; oblique to streaked; reddish brown or purple; with green overlay; length of seed; without a joint evident between the glabrous (at first glance glabrous but with careful radicle and the cotyledons. Radicle linear; deflexed and examination puberulent) or pubescent and indurate; parallel to cotyledon length; centered between cotyle- with 1 type of pubescence; puberulent (tiny hairs); with dons; less than 1/2 length of cotyledons. Plumule pubescence gray; with pubescence uniformly distrib- moderately developed; glabrous. uted; pliable; with hair bases plain; glandular or eglandular; with glandular dots (absent to numerous); Distribution: India. without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; Notes: Based on its fruit characters (presence or absence of without cracks. Mesocarp present or absent; trace or hairs and glandular dots), Eleiotis monophylla may be a thin (very); surface not veined; 1-layered; without mixed species. balsamic vesicles; without fibers; solid; chartaceous. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; remaining fused to epicarp; entire. Seed 1; length parallel with fruit length. Funicu- lus less than 0.5 mm long; flattened; straight. Aril present (inconspicuous) or absent; dry; rim-aril and tongue-aril; reddish brown.

Seed 4 ϫ 2.7 ϫ 1Ð1.1 mm; not overgrown; not angular; asymmetrical; reniform (sub-) or oblong; compressed; with surface smooth; without visible radicle and Eleiotis: E. monophylla (N.L. Burman) A.-P. de Candolle cotyledon lobes; without hilar sinus; without umbo on (AÐE). A, Fruits (ϫ 7.8); B, seeds (ϫ 10.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

685 Genus: Phylacium J.J. Bennett endocarp layer P. majus); black, purple, pale pink, or yellow; glabrous; smooth (shagreen) or not smooth; Phylogenetic Number: 11.22. with elevated features; shagreen and warty (small); coriaceous. Fracture lines absent. Rim absent. Wings Tribe: Desmodieae. present. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split lighter colored Subtribe: Lespedezinae. than the rest of the hilum and therefore conspicuous; larger than punctiform; 1 mm long; with curved outline; Species Studied—Species in Genus: 2 spp.—2 spp. elliptic; marginal according to radicle tip or between cotyledon and radicle lobe; slightly recessed; not within Fruit a legume; unilocular; 0.7Ð1.2 ϫ 0.4Ð0.6 ϫ 0.15Ð0.2 corona, halo, or rim or within halo. Hilum halo color cm; with persistent or deciduous calyx; with calyx lighter than testa. Lens discernible; less than 0.5 mm in shorter than fruit; without orifice formed by curving of length or equal to or greater than 0.5 mm in length; 0.5 fruit or fruit segments; straight; not plicate; not twisted; mm long; with margins straight or curved; more or less asymmetrical; when asymmetrical with 1 straight and 1 triangular; elliptic; not in groove of raphe; confluent curved (more or less) suture; widest near middle or D- with hilum; mounded; same color as testa; not within shaped (more or less); not inflated; flattened; with beak; corona, halo, or rim. Endosperm thick; covering entire declined (especially, tip of P. bracteosum J.J. Bennett) embryo; adnate to embryo. Cotyledons smooth; both or coiled (especially tip of P. majus); with solid beak outer faces convex; both the same thickness; both more the same color and texture as fruit; short tapered at or less of equal length; not folded; margin entire 180 apex; apex aligned or oblique with longitudinal axis of degrees from base of radicle; similar at apex; not fruit (especially tip); short tapered at base; base aligned concealing radicle; entire over radicle; without lobes; with longitudinal axis of fruit; with the apex and base with the interface division terminating at base of uniform in texture; membranous; seed chambers radicle; without margins recessed; reddish brown or externally visible. Fruit margin not constricted; without red; inner face flat; glabrous around base of radicle. sulcus; plain. Fruit wings absent. Fruit nonstipitate. Embryonic axis deflexed; oblique to length of seed; Fruit indehiscent. Replum invisible. Epicarp dull; without a joint evident between the radicle and the monochrome; reddish brown to brown; pubescent and cotyledons. Radicle linear; deflexed and parallel to indurate; with hairs erect or appressed; with 1 type of cotyledon length; centered between cotyledons; less pubescence; puberulent; with pubescence gray; with than 1/2 length of cotyledons. Plumule rudimentary; pubescence uniformly distributed; with simple hairs; glabrous. pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately Distribution: China, Indochina, and Philippines to Australia veined; not tuberculate; not exfoliating; without cracks. (northern Queensland). Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Notes: Bresser (1978) monographed the genus and carefully Endocarp dull; monochrome; reddish brown; smooth; described the “most striking” feature of the inflores- nonseptate; chartaceous; not exfoliating; remaining cences: “Large pale green bracts, which cover the fruits fused to mesocarp and epicarp; entire. Seed 1; length and the old flowers.” Figure A was lighted from the parallel with fruit length. Funiculus less than 0.5 mm back to show the two fruits inside the bracts. There may long; filiform or thick and triangular; straight. Aril dry; be one to three fruits in each set of bracts, and this unit rim-aril; reddish brown to brown. may function as a “winged fruit.” Bailey et al. (1997), using the chloroplast rpl2 intron, suggested that Seed 4Ð5 ϫ 1.5Ð3 ϫ 2Ð2.2 mm; not overgrown; not Phylacium is not a member of Desmodieae, but did not angular; symmetrical; reniform, circular, or ovate; indicate where it should be placed. compressed; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering or partially adhering to endocarp (very thin layer P. Phylacium: P. majus H. Collett & W.B. Hemsley (CÐE), P. majus); dull; not modified by a bloom; colored; spp. (AÐB). A, Fruits within large bracts and a freed monochrome (but with very thin reddish-brown fruit (see Notes) (ϫ 1.7); B, seeds (ϫ 9); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

687 Genus: Campylotropis A.A. von Bunge punctiform; 0.4 mm long; with curved outline; circular; marginal according to radicle tip; flush; within rim, or Phylogenetic Number: 11.23. within halo (halo absent or faintly present). Hilum halo color darker than testa. Hilum rim color of testa. Lens Tribe: Desmodieae. discernible; less than 0.5 mm in length; with margins curved; elliptic; not in groove of raphe; confluent with Subtribe: Lespedezinae. hilum (or nearly so); flush; dissimilar color from testa; darker than testa; black; not within corona, halo, or rim. Species Studied—Species in Genus: 15 spp.—ca. 65 spp. Endosperm absent. Cotyledons smooth; both outer faces convex; both the same thickness; both more or Fruit a legume; unilocular; 0.8Ð1.5 ϫ 0.4Ð0.5 ϫ 0.05Ð0.07 less of equal length; not folded; margin entire 180 cm; with deciduous or persistent calyx; with calyx degrees from base of radicle; similar at apex; entire shorter than fruit; without orifice formed by curving of over radicle; without lobes; with the interface division fruit or fruit segments; straight or curved (slightly); not terminating at base of radicle; without margins re- plicate; not twisted; asymmetrical or symmetrical; cessed; reddish to greenish brown; inner face flat; oblong; when asymmetrical with 1 straight and 1 glabrous around base of radicle. Embryonic axis curved suture or both sutures parallelly curved; widest deflexed; oblique to length of seed; without a joint near middle or D-shaped; not inflated; flattened; evident between the radicle and the cotyledons. Radicle without or with beak; straight; with solid beak the same linear; deflexed and parallel to cotyledon length; color and texture as fruit; rounded or short tapered at centered between cotyledons; less than 1/2 length of apex; apex aligned with longitudinal axis of fruit; cotyledons. Plumule rudimentary; glabrous. tapered at base; base aligned with longitudinal axis of fruit; with the apex and base uniform in texture; Distribution: Asia. membranous; seed chambers externally visible. Fruit margin not constricted; without sulcus; plain. Fruit Notes: Ohashi (1974a,b,c) published a trio of papers on wings present. Fruit substipitate. Fruit indehiscent. Campylotropis, and Fu (1987) recognized 29 species in Replum invisible. Epicarp dull; monochrome; brown; the People’s Republic of China. Akiyama and Ohba pubescent and indurate or glabrous (except for stiff (1985) and Nemoto and Ohashi (1988) maintained hairs along sutures); with hairs erect or appressed; with Campylotropis, Kummerowia (11.25), and Lespedeza 1 type of pubescence; pilose; with pubescence brown (11.24) as distinct genera. (to reddish) or golden; with pubescence uniformly distributed; with simple hairs; pliable or stiff (golden hairs along sutures); with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp glossy; monochrome; reddish brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seed 1; length parallel with fruit length. Funiculus less than 0.5 mm long; thick; straight. Aril absent.

Seed 3.7Ð4.5 ϫ 2Ð2.5 ϫ 1.3Ð1.5 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Campylotropis: C. macrocarpa (A.A. von Bunge) A. Rim absent. Wings absent. Raphe not visible. Hilum Rehder (CÐE), C. spp. (AÐB). A, Fruits (with and visible; with faboid split; with the lips of the faboid without calyx) (ϫ 4.3); B, seeds (ϫ 6.3); CÐD, testa split the same color as the rest of the hilum; larger than (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

689 Genus: Lespedeza A. Michaux Seed 1.5Ð5 ϫ 1.3Ð3 ϫ 0.7Ð1.5 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with Phylogenetic Number: 11.24. surface smooth; without or with visible radicle and cotyledon lobes; without or with external groove Tribe: Desmodieae. between radicle and cotyledon lobes; same color as testa; without hilar sinus; without umbo on seed faces. Subtribe: Lespedezinae. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome or mottled Species Studied—Species in Genus: 30 spp.—ca. 40 spp. and streaked; with frequent mottles; light to dark reddish brown, tan, purple, yellow (greenish), green, or Fruit a legume; unilocular; 0.25 ϫ 0.2Ð0.4 ϫ 0.1Ð0.2 cm; black; with purple overlay; glabrous; smooth; coria- with persistent or deciduous calyx; with calyx shorter ceous. Fracture lines absent. Rim absent. Wings absent. than or equal in length to fruit; without orifice formed Raphe not visible. Hilum visible; with faboid split; with by curving of fruit or fruit segments; straight; not the lips of the faboid split the same color as the rest of plicate; not twisted; asymmetrical or symmetrical; the hilum; punctiform or larger than punctiform; 0.3Ð elliptic, oblong, ovate, circular, linear (somewhat), or 0.6 mm long; with straight outline; oblong; marginal C-shaped (slightly); when asymmetrical with 1 straight according to radicle tip; recessed; within halo. Hilum and 1 curved suture or both sutures parallelly curved; halo color lighter or darker than testa. Lens discernible; widest near middle or D-shaped; not inflated; com- less than 0.5 mm in length; with margins straight or pressed; with or without beak; hooked or straight; with curved; oblong, wedge-shaped, or linear (somewhat and solid beak the same color and texture as fruit; short with or without medial groove); circular or oblong; not tapered at apex; apex aligned with longitudinal axis of in groove of raphe; confluent with or adjacent to hilum; fruit; short tapered at base; base aligned with longitudi- up to 0.2 mm from hilum; flush; dissimilar color from nal axis of fruit; with the apex and base uniform in testa; darker or lighter than testa; black or tan; not texture; membranous; seed chambers externally visible. within corona, halo, or rim. Endosperm thin; covering Fruit margin not constricted; without sulcus; plain or entire embryo; adnate to testa or embryo. Cotyledons embellished; with fringe (of hairs). Fruit wings absent. smooth; both outer faces convex; both the same Fruit substipitate. Fruit indehiscent. Replum invisible. thickness; both more or less of equal length; not folded; Epicarp dull; monochrome; reddish brown, tan (to margin entire 180 degrees from base of radicle; similar greenish), green, or gray (because of hairs); pubescent at apex; not concealing radicle; entire over radicle; and indurate, pubescent but soon deciduous, or without lobes; with the interface division terminating at glabrous; with hairs appressed (often missing from base of radicle; without margins recessed; white, center of each valve) or erect; with 1 type of pubes- yellow, green, or tan; inner face flat; glabrous around cence; puberulent; with pubescence gray or with base of radicle. Embryonic axis deflexed; oblique to pubescence golden; with apical pubescence different length of seed; without a joint evident between the from basal pubescence; with apical 1/3Ð1/2 pubescent radicle and the cotyledons. Radicle linear; deflexed and and basal 1/2Ð2/3 glabrous; with simple hairs (either parallel to cotyledon length; centered between cotyle- straight or somewhat hooked at apex); pliable or stiff; dons; less than 1/2 length of cotyledons. Plumule with hair bases plain; antrorse; straight or curved at moderately developed or rudimentary; glabrous. apex; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; Distribution: Temperate North America, eastern and tropical not exfoliating; without cracks. Mesocarp present or Asia, and Australia. absent; trace or thin (usually very); surface not veined; 1-layered; without balsamic vesicles; without fibers; Notes: Akiyama (1988) monographed section solid; chartaceous. Endocarp dull; monochrome; Macrolespedeza, and Akiyama and Ohba (1988) reddish brown or tan; smooth; nonseptate; chartaceous; revised Lespedeza formosa (J.R.T. Vogel) B.A.E. not exfoliating; remaining fused to mesocarp and Koehne. Clewell (1966) monographed the native epicarp; remaining fused to epicarp; entire. Seed 1; species of Lespedeza in North America. Although length parallel with fruit length. Funiculus less than 0.5 cleistogamous and chasmogamous flowers may mm long or measured; up to 1 mm long; flattened, produce slightly different fruits, these differences do thick, or partially filiform and partially thick; curved or not affect the genus circumscription of the fruits and straight. Aril dry; rim-aril; tan. seeds. Nemoto and Ohashi (1993) studied Lespedeza

690 seedling morphology and presented excellent SEM illustrations of the embryo and plumule. They found that the first pair of plumule leaves are either opposite or alternate and that their relative position is positively correlated with the subgeneric classification. Lespedeza stipulacea C.J. Maximowicz and L. striata C.P. Thunberg are now in the segregate genus Kummerowia (11.25). Akiyama and Ohba (1985) and Nemoto and Ohashi (1988) maintained Campylotropis (11.23), Kummerowia (11.25), and Lespedeza as distinct genera. Nemoto et al. (1995), using morphological and molecular evidence, concluded that “Lespedeza is closer to Kummerowia than to Campylotropis.”

691 Lespedeza: L. virginica (C. Linnaeus) N.L. Britton (CÐE), L. spp. (AÐB). A, Fruits (with and without calyx) (ϫ 2.8); B, seeds (ϫ 4.7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 13).

693 Genus: Kummerowia A.K. Schindler lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of Phylogenetic Number: 11.25. the faboid split lighter colored than the rest of the hilum and therefore conspicuous; punctiform or larger than Tribe: Desmodieae. punctiform; 0.3 mm long; with curved outline; circular; between cotyledon and radicle lobe or subapical to Subtribe: Lespedezinae. radicle tip; flush; within halo. Hilum halo color lighter than testa. Lens discernible; less than 0.5 mm in length; Species Studied—Species in Genus: 2 spp.—2 spp. with margins curved; circular; not in groove of raphe; confluent with hilum; recessed; dissimilar color from Fruit a legume; unilocular; 0.28Ð0.45 ϫ 0.15Ð0.23 ϫ 0.1Ð testa; darker or lighter than testa; reddish brown, tan 0.13 cm; with persistent or deciduous calyx; with calyx (greenish), or green; not within corona, halo, or rim. shorter than or equal (nearly) in length to fruit; without Endosperm thin; covering entire embryo; adnate to orifice formed by curving of fruit or fruit segments; testa. Cotyledons smooth; both outer faces convex; both straight; not plicate; not twisted; symmetrical; oblong; the same thickness; both more or less of equal length; not inflated; compressed; without beak; short tapered at not folded; margin entire 180 degrees from base of apex; apex aligned with longitudinal axis of fruit; short radicle; similar at apex; not concealing radicle; entire tapered at base; base aligned with longitudinal axis of over radicle; without lobes; with the interface division fruit; with the apex and base uniform in texture; terminating at base of radicle; without margins re- chartaceous; seed chambers externally visible. Fruit cessed; tan; inner face flat; glabrous around base of margin not constricted; without sulcus; plain. Fruit radicle. Embryonic axis deflexed; oblique to length of wings absent. Fruit nonstipitate. Fruit indehiscent. seed; without a joint evident between the radicle and Replum invisible. Epicarp dull; monochrome; brown or the cotyledons. Radicle linear; deflexed and parallel to tan; pubescent but soon deciduous, pubescent and cotyledon width; centered between cotyledons; 1/2 to indurate, or glabrate; with hairs appressed or erect; with nearly length of cotyledons. Plumule moderately 1 type of pubescence; velutinous; with pubescence developed; glabrous. gray; with pubescence uniformly distributed or with apical pubescence different from basal pubescence; Distribution: Asia and naturalized in temperate North with apical 1/3Ð1/2 pubescent and basal 1/2Ð2/3 America. glabrous; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with Notes: Isely (1948) discussed the two species of annual elevated features; reticulately veined; tuberculate; not Lespedeza naturalized in the United States, which now exfoliating; without cracks. Mesocarp absent. Endocarp are placed in the genus Kummerowia. Akiyama and dull; monochrome; tan; smooth; nonseptate; Ohba (1985) and Nemoto and Ohashi (1988) main- chartaceous; not exfoliating; remaining fused to tained Campylotropis (11.23), Kummerowia, and epicarp; entire. Seed 1; length parallel with fruit length. Lespedeza (11.24) as distinct genera. Funiculus less than 0.5 mm long; more or less filiform; curved. Aril present (barely visible) or absent; dry; rim- aril; reddish tan or brown (reddish).

Seed 1.5Ð2.3 ϫ 1.5Ð1.8 ϫ 0.8Ð1 mm; not overgrown; not angular; symmetrical (except hilum); oblong, ovate, or elliptic; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy; not Kummerowia: K. stipulacea (C.J. Maximowicz) T. Makino modified by a bloom; colored; monochrome or mottled (CÐE), K. spp. (AÐB). A, Fruits with and without calyx and streaked; with frequent mottles; with frequent (ϫ 8.7); B, seeds (ϫ 9.3); CÐD, testa (ϫ 50, ϫ 1000); streaks; dark reddish brown or tan (reddish); with E, embryos (ϫ 10). purple overlay; glabrous; smooth; coriaceous. Fracture

695 Psoraleeae (12.01Ð12.09) faboid split; punctiform; marginal according to radicle tip; recessed; within corona. Hilum corona color lighter Genus: Cullen F.C. Medikus than testa (reddish to yellow). Lens not discernible. Endosperm thin; covering entire embryo; adnate to Phylogenetic Number: 12.01. testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; Tribe: Psoraleeae. not folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing radicle; entire Species Studied—Species in Genus: 10 spp.—32 spp. over radicle; without lobes; with the interface division terminating in radicle tissue; without margins recessed; Fruit a legume; unilocular; 0.25Ð0.7 ϫ 0.15Ð0.35 ϫ 0.1Ð0.5 yellow; inner face flat; glabrous around base of radicle. cm; with persistent calyx; with calyx longer than fruit; Embryonic axis deflexed; parallel to length of seed. without orifice formed by curving of fruit or fruit Radicle linear; deflexed and parallel to cotyledon segments; straight to curved; not plicate; not twisted; length; centered between cotyledons; 1/2 to nearly asymmetrical or symmetrical; oblong or reniform; when length of cotyledons. Plumule rudimentary; glabrous. asymmetrical with both sutures parallelly or unequally curved; partially inflated or not inflated; compressed; Distribution: Africa through India and Sri Lanka to Burma, without beak; short tapered to rounded at apex; apex Philippines, Papua New Guinea, and Australia. aligned to oblique to right-angled (nearly) with longitudinal axis of fruit; rounded at base; base aligned to Notes: Stirton (1981) illustrated seeds and fruits of oblique with longitudinal axis of fruit; with the apex Amorpheae and Psoraleeae. Since Isely (1962) and base uniform in texture; membranous; seed monographed the tribe for north-central United States chambers externally visible. Fruit margin not con- the spelling of the tribe name has changed as well as the stricted; without sulcus; plain. Fruit wings absent. Fruit genera recognized in the tribe. Amorpha (6.04), Dalea nonstipitate. Fruit indehiscent. Replum invisible. (6.08), and Petalostemon A. Michaux (now part of Epicarp dull; monochrome; black or red (brownish Dalea) are now in the Amphoreae, and species in the often concealed by deciduous white hairs); glabrous, remaining genus, Psoralea (12.09), have been assigned pubescent and indurate, or pubescent but soon decidu- to several genera treated here. Grimes (1990) noted that ous; with 1 type of pubescence; with pubescence gray; this tribe “has been described as having indehiscent with pubescence uniformly distributed; with simple fruits. However, in many North American species the hairs; pliable; with hair bases plain; glandular or fruit is secondarily dehiscent (that is, not along sutures) eglandular; without spines; not smooth; with elevated by transverse rupture of the pod.” This technically is or recessed features; veined or not veined; transversely not dehiscence. Stirton (1981) transferred six African veined relative to fruit length; not tuberculate; scaly or species of Psoralea (12.09) to Cullen. The unit of wrinkled; glandularly punctate; exfoliating in part or dispersal may include the soon-deciduous to apparently not exfoliating; without cracks. Mesocarp absent. permanent-papery-to-leathery 5-lobed calyx. The calyx Endocarp absent. Seed 1; length parallel with fruit bears reddish glands and may be glabrous to pubescent length. Funiculus less than 0.5 mm long; filiform; with silvery to golden hairs. Grimes (1997) revised straight. Aril present or absent; dry; rim-aril; white. Cullen, and his species count is used. He accepted it as monophyletic because all of its species have a small Seed 2.5Ð5 ϫ 1.5Ð3.5 ϫ 1Ð2.5 mm; not overgrown; not invagination of the epicarp just above the fruit stalk on angular; asymmetrical; reniform; compressed; with or the ventral side. The invagination only partially without visible radicle and cotyledon lobes; without penetrates the epicarp. The fruit of C. glandulosa (C. umbo on seed faces. Testa with pieces of adhering Linnaeus) J.W. Grimes absices below the calyx and epicarp; partially adhering to endocarp; somewhat travels with it, while in C. americanum the fruit falls glossy; not modified by a bloom; colored; monochrome free of the calyx. The fruit of C. glandulosa is unusual or mottled; with frequent mottles; reddish brown or among the studied species because the upper half is green (brownish); with brownish black overlay; more or less inflated and the lower half is adnate to the glabrous; smooth (when partially adhering endocarp not testa. considered); coriaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum fully concealed; concealed by funiculus or funicular remnant; without

696 697 Cullen: C. americanum (C. Linnaeus) P.A. Rydberg (CÐE), C. spp. (AÐB). A, Fruits with and without calyx (ϫ 3.4); B, embryos with and without epicarp (ϫ 5); CÐD, exocarp (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

699 Genus: Bituminaria L. Heister ex P.C. Fabricius Seed 3.5Ð4.5 ϫ 2.5Ð3.5 ϫ 1.5Ð2.5 mm; not overgrown; not angular; asymmetrical; D-shaped; quadrangular; Phylogenetic Number: 12.02. without visible radicle and cotyledon lobes; without umbo on seed faces. Testa absent. Endosperm thin; Tribe: Psoraleeae. covering entire embryo; adnate to testa. Cotyledons smooth; both outer faces convex; both the same Species Studied—Species in Genus: 1 sp.—2 spp. thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; similar Fruit a legume; unilocular; 0.4Ð0.6 (exclusive of fragile at apex; not concealing radicle; entire over radicle; beak up to 1 cm long) ϫ 0.3Ð0.4 ϫ 0.2Ð0.3 cm; with without lobes; with the interface division terminating in persistent or deciduous calyx; with calyx shorter than radicle tissue; without margins recessed; dark tan; inner fruit; without orifice formed by curving of fruit or fruit face flat; glabrous around base of radicle. Embryonic segments; straight; not plicate; not twisted; asymmetri- axis deflexed; parallel to length of seed. Radicle linear; cal; when asymmetrical with 1 straight and 1 curved deflexed and parallel to cotyledon length; centered suture; widest near middle or D-shaped; not inflated; between cotyledons; 1/2 to nearly length of cotyledons. compressed; with beak; declined; with papery fragile Plumule moderately developed; glabrous. beak up to 1 cm long; truncate at apex (exclusive of beak); apex aligned with longitudinal axis of fruit; Distribution: Mediterranean Europe, North Africa, and rounded at base; base aligned with longitudinal axis of southwestern Asia. fruit; with the apex and base uniform in texture; coriaceous or fragile, thinner than chartaceous like Notes: Intact seeds of B. bituminosa do not exist free of the Trifolium (23.07); seed chambers externally visible. fruit because the testa is fused to the endocarp. There- Fruit margin not constricted; without sulcus; embel- fore, the testa, raphe, hilum, and lens characters could lished; with prickles. Fruit wings absent. Fruit not be scored. The fruits have an aroma reminiscent of nonstipitate. Fruit indehiscent or with all layers fenugreek (Trigonella foenum-graecum, 21.04). dehiscing (secondarily: see Notes). Replum invisible. Epicarp dull; multicolored; mottled; brown or gray; with red overlay (if spines knocked off); with surface texture uniform or not uniform, with patches of different texture not restricted to the base and apex; pubescent but soon deciduous and glabrous (except apex); with 1 type of pubescence; villous; with pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; glandular; with glandular dots (reddish brown); with spines (black on seed chamber); with spines persistent or broken off and their bases evident; with spines same color as or a different color (or their basal remanent) from the rest of the fruit; not smooth; with elevated or recessed features; not veined; not tuberculate; faintly wrinkled; glandularly punctate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 2-layered; without balsamic vesicles; without fibers; with spongy layer over solid layer; coriaceous. Endocarp dull; monochrome; grayish black; smooth; nonseptate; coriaceous; not exfoliating; remaining fused to mesocarp and epicarp. Seed 1; length parallel with fruit Bituminaria: B. bituminosa (C. Linnaeus) C.H. Stirton length. Funiculus less than 0.5 mm long; straight. Aril (AÐE). A, Fruits (without beak and hairs, within calyx, absent. and with beak and hairs) (ϫ 4.9); B, embryo and two fruits functioning as seeds (ϫ 8); CÐD, exocarp (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

701 Genus: Pediomelum P.A. Rydberg Gray) P.A. Rydberg) or not smooth; with elevated or recessed features; wrinkled (P. castoreum (S. Watson) Phylogenetic Number: 12.03. P.A. Rydberg); pitted with small separate pits (faintly and widely scattered); coriaceous. Fracture lines absent. Tribe: Psoraleeae. Rim absent. Raphe not visible. Hilum fully concealed; concealed by funicular remnant; with faboid split; with Species Studied—Species in Genus: 4 spp.—21 spp. the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; up to 0.6 mm long; Fruit a legume; unilocular; 0.8Ð1.6 (including beak) ϫ 0.4Ð with curved outline; circular or elliptic; marginal 0.5 ϫ 0.2Ð 0.25 cm; with persistent calyx; with calyx according to radicle tip; recessed; within corona or not shorter than fruit (but not beak); without orifice formed within corona, halo, or rim. Hilum corona color lighter by curving of fruit or fruit segments; straight; not than testa. Lens discernible; equal to or greater than 0.5 plicate; not twisted; asymmetrical or symmetrical mm in length; up to 0.5 mm long; with margins straight (except beak); oblong; when asymmetrical with both or curved; linear, circular, or oblong; not in groove of sutures parallelly curved; not inflated; compressed; with raphe; adjacent to hilum; up to 0.2 mm from hilum; beak; straight; with solid beak the same color and mounded; dissimilar color from testa; darker than testa; texture as fruit; long tapered or rounded at apex; apex black; not within corona, halo, or rim. Endosperm thin aligned to oblique with longitudinal axis of fruit; (thinnest of all genera in tribe); covering entire embryo; rounded at base; base aligned with longitudinal axis of adnate to testa. Cotyledons smooth; both outer faces fruit; with the apex and base uniform or differing in convex; both the same thickness; both more or less of texture; upper 3/4 barely inflated, reticulate, and equal length; not folded; margin entire 180 degrees pubescent and lower 1/4 not inflated, reticulate, or from base of radicle; similar at apex; not concealing pubescent; chartaceous or fragile, thinner than radicle; entire over radicle; without lobes; with the chartaceous like Trifolium (21.06); seed chambers interface division terminating in radicle tissue; without externally visible. Fruit margin not constricted; without margins recessed; tan; inner face flat; glabrous around sulcus; plain or embellished; with thickened sutural base of radicle. Embryonic axis deflexed; parallel to areas (thickened margins). Fruit wings absent. Fruit length of seed. Radicle linear; deflexed and parallel to substipitate to nonstipitate. Fruit indehiscent. Replum cotyledon length; centered between cotyledons; 1/2 to invisible. Epicarp dull; monochrome; red or tan; nearly length of cotyledons. Plumule rudimentary; pubescent and indurate; with 1 type of pubescence; glabrous. villous; with pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with Distribution: North America, Mexico, and central and hair bases plain; glandular or eglandular; with glandular southern Africa. dots; limited to a portion of fruit; upper 3/4 glandular and lower 1/4 eglandular; without spines; not smooth; Notes: Grimes (1990) recognized 21 species, and we are with elevated features; reticulately veined; not tubercu- following his count. Pediomelum argophyllum (F.T. late; not exfoliating; without cracks. Mesocarp absent. Pursh) J.W. Grimes and P. castoreum have beaks like Endocarp dull; monochrome; tan; smooth; nonseptate; Bitumaria (12.02), but their fruits are too fragile and chartaceous; not exfoliating; remaining fused to were too few in number in our sample for our analysis epicarp; entire. Seed 1; length parallel with fruit length. to be considered complete. Funiculus measured; less than 1 mm long; straight. Aril absent.

Seed 4.5Ð6 ϫ 2.8Ð4 ϫ 1.5Ð2.5 mm; not overgrown, 1 seed filling entire fruit cavity; not angular; symmetrical or asymmetrical; oblong; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa partially adhering to endocarp (may have); Pediomelum: P. esculentum (F.T. Pursh) P.A. Rydberg glossy; not modified by a bloom; colored; monochrome (CÐE), P. spp. (AÐB). A, Fruits with and without calyx or streaked; with frequent streaks; green (ish), red (ϫ 3.2); B, seeds (ϫ 6); CÐD, testa (ϫ 50, ϫ 1000); E, (brownish), tan, or yellow; with black (faintly) or embryos (ϫ 6). brown overlay; glabrous; smooth (P. cyphocalyx (A.

703 Genus: Psoralidium P.A. Rydberg ible; equal to or greater than 0.5 mm in length; up to 0.6 mm long; with margins straight or curved; linear, Phylogenetic Number: 12.04. elliptic, oblong, or key-hole shaped (P. lanceolatum); not in groove of raphe; confluent with hilum; recessed; Tribe: Psoraleeae. dissimilar color from testa; darker than testa; black; not within corona, halo, or rim. Endosperm thin; covering Species Studied—Species in Genus: 2 spp.—13 spp. entire embryo; adnate to testa. Cotyledons outer face of 1 cotyledon flat and other cotyledon concave; 1 thicker Fruit a legume; unilocular; 0.4Ð0.95 (including beak up to 3 than the other; both more or less of equal length; with mm long in P. tenuifolium) ϫ 0.35Ð0.5 ϫ 0.25Ð0.4 cm; both or only 1 folded; margin entire 180 degrees from with persistent or deciduous calyx; with calyx shorter base of radicle; similar at apex; not concealing radicle; than fruit; without orifice formed by curving of fruit or entire over radicle; without lobes; with the interface fruit segments; straight; not plicate; not twisted; sym- division terminating in radicle tissue; without margins metrical; circular, elliptic, or oblong; not inflated; recessed; yellow or green; inner face flat; glabrous compressed to terete; without or with beak; straight; with around base of radicle. Embryonic axis deflexed; solid beak the same color and texture as fruit (P. parallel to length of seed. Radicle linear; deflexed and tenuifolium (F.T. Pursh) P.A. Rydberg); rounded at parallel to cotyledon length; centered between cotyle- apex; apex aligned to oblique with longitudinal axis of dons; less than 1/2 length of cotyledons. Plumule fruit; short tapered to rounded at base; base aligned to moderately developed; glabrous. oblique with longitudinal axis of fruit; with the apex and base uniform in texture; chartaceous; seed chambers Distribution: Western Canada and western United States. externally invisible. Fruit margin not constricted; without sulcus; plain. Fruit nonstipitate. Fruit indehis- Notes: Our species count is based on Grimes (1990). The cent. Replum invisible. Epicarp dull; monochrome (but micropyle of P. tenuiflorum is bright reddish and with well developed reddish-brown glands); tan; therefore more conspicuous than the black lens. pubescent and indurate; with 1 type of pubescence; tomentose; with pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; glandular; with glandular dots; without spines; not smooth; with elevated or recessed features; not veined; tuberculate; glandularly punctate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp. Seed 1; length parallel with fruit length. Funiculus measured; up to 1 mm long; thick; straight. Aril absent.

Seed 3.4Ð5.5 ϫ 3Ð3.5 ϫ 2Ð3 mm; not overgrown; angular to not angular; symmetrical or asymmetrical; circular, irregular (many seeds have large, irregularly placed dimples), or oblong; terete to compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or streaked; with frequent streaks; brownish green or red (brownish); with purple overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.4 mm long; with curved or straight Psoralidium: P. lanceolatum (F.T. Pursh) P.A. Rydberg outline; elliptic or oblong; marginal according to radicle (CÐE), P. spp. (AÐB). A, Fruits (ϫ 6.7); B, seeds tip; flush; not within corona, halo, or rim. Lens discern- (ϫ 7.8); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

705 Genus: Rupertia J.W. Grimes 0.5 mm long; with margins straight; wedge-shaped; not in groove of raphe; confluent with hilum; flush; Phylogenetic Number: 12.05. dissimilar color from testa; darker than testa; black; within corona. Lens corona color darker than testa. Tribe: Psoraleeae. Endosperm absent. Cotyledons smooth; both outer faces convex; both the same thickness; both more or Species Studied—Species in Genus: 2 spp.—3 spp. less of equal length; not folded; margin entire 180 degrees from base of radicle; similar at apex; not Fruit a legume; unilocular; 0.4Ð0.7 ϫ 0.3Ð0.5 ϫ 0.27 cm; concealing radicle; entire over radicle; without lobes; with persistent calyx; with calyx equal in length to fruit; with the interface division terminating in radicle tissue; without orifice formed by curving of fruit or fruit green; inner face flat; glabrous around base of radicle. segments; straight; not plicate; not twisted; symmetri- Embryonic axis deflexed; parallel to length of seed. cal; elliptic; not inflated; compressed; without (R. Radicle linear; deflexed and parallel to cotyledon physodes) or with beak; with solid beak the same color length; centered between cotyledons; less than 1/2 and texture as fruit (beak 1Ð3 mm long in R. hallii (P.A. length of cotyledons. Plumule rudimentary; glabrous. Rydberg) J.W. Grimes and R. rigida (S.B. Parish) J.W. Grimes); short tapered or rounded at apex; apex aligned Distribution: Western Canada and United States to Mexico with longitudinal axis of fruit; rounded at base; base (Baja California). aligned with longitudinal axis of fruit; with the apex and base uniform; chartaceous; seed chambers exter- Notes: Grimes (1990) founded the genus. nally visible. Fruit margin not constricted; without sulcus; plain. Fruit wings absent. Fruit nonstipitate. Fruit indehiscent. Replum invisible. Epicarp dull; monochrome; tan; pubescent and indurate; with 1 type of pubescence; puberulent; with pubescence gray or brown; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; glandular or eglandular; with glandular dots (golden and fading with age); without spines; not smooth; with elevated features; veined or not obliquely veined relative to fruit length (at least lower one-half); not tuberculate; wrinkled; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; spongy; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp. Seed 1; length parallel with fruit length. Funiculus measured; less than 1 mm long; thick; straight. Aril absent.

Seed 4.5Ð7 ϫ 2.7Ð4 ϫ 1.7 mm; not overgrown; not angular; asymmetrical; reniform; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; brownish red; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.8 mm long; with curved outline; circular; marginal Rupertia: R. physodes (D. Douglas ex W.J. Hooker) J.W. according to radicle tip; recessed; within corona. Hilum Grimes (AÐE). A, Fruits with and without calyx (ϫ 6); corona color darker than testa. Lens discernible; equal B, seed (ϫ 8.5); CÐD, testa (ϫ 50, ϫ 1000); E, to or greater than 0.5 mm or less than 0.5 mm in length; embryos (ϫ 10).

707 Genus: Hoita P.A. Rydberg (with conspicuous light tan rim); with faboid split; with the lips of the faboid split the same color as the rest of Phylogenetic Number: 12.06. the hilum; punctiform; marginal according to radicle tip; recessed; within rim. Hilum rim color lighter than Tribe: Psoraleeae. testa. Lens not discernible. Endosperm thin; covering entire embryo; adnate to testa. Cotyledons smooth; both Species Studied—Species in Genus: 3 spp.—3 spp. outer faces convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 Fruit a legume; unilocular; 0.6Ð1 ϫ 0.3Ð0.5 ϫ 0.2Ð0.25 cm; degrees from base of radicle; similar at apex; not with persistent or deciduous calyx; with calyx longer or concealing radicle; entire over radicle; without lobes; shorter than fruit; without orifice formed by curving of with the interface division terminating in radicle tissue; fruit or fruit segments; straight; not plicate; not twisted; without margins recessed; yellow; inner face flat; asymmetrical; when asymmetrical with 1 straight and 1 glabrous around base of radicle. Embryonic axis curved suture; widest near middle or D-shaped; barely deflexed; parallel to length of seed. Radicle linear; inflated or not inflated; compressed; without beak; short deflexed and parallel to cotyledon length; centered tapered at apex; apex aligned with longitudinal axis of between cotyledons; 1/2 to nearly length of cotyledons. fruit; rounded at base; base oblique with longitudinal Plumule moderately developed; glabrous. axis of fruit; with the apex and base differing in texture; upper 3/4 barely inflated, reticulate, and pubescent and Distribution: Canada, western United States, and Mexico lower 1/4 not inflated, reticulate, or pubescent; (Baja California). chartaceous; seed chambers externally invisible. Fruit margin not constricted; without sulcus; plain. Fruit Notes: Our species count is based on Grimes (1990). wings absent. Fruit nonstipitate. Fruit indehiscent. Replum invisible. Epicarp dull; monochrome; brown; pubescent and indurate; with 1 type of pubescence; pilose; with pubescence golden; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; glandular; with glandular dots; limited to a portion of fruit; upper 3/4 glandular and lower 1/4 eglandular; without spines; not smooth; with elevated features; veined or not veined; transversely veined relative to fruit length; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seed 1; length parallel with fruit length. Funiculus measured; less than 1 mm long; thick; straight. Aril absent.

Seed 5.5Ð6.7 ϫ 3.5Ð3.7 ϫ 2Ð2.5 mm; not overgrown; not angular; symmetrical (except for hilum); elliptic; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; monochrome or streaked; with frequent streaks; dark reddish brown; with brown (in dark, broad bands) overlay; glabrous; smooth or not smooth; with recessed features; pitted with small separate pits (some seeds with numerous pits and other seeds with few to no pits Hoita: H. macrostachys (A.-P. de Candolle) P.A. Rydberg H. strobilina (J.W. Hooker & G.A. Arnott) G.A.W. (CÐE), H. spp. (AÐB). A, Fruits and fruit in calyx Rydberg); coriaceous. Fracture lines absent. Rim (ϫ 4.5); B, seeds (ϫ 8); CÐD, testa (ϫ 50, ϫ 1000); E, absent. Wings absent. Raphe not visible. Hilum visible embryos (ϫ 6).

709 Genus: Orbexilum C.S. Rafinesque-Schmaltz the hilum; larger than punctiform; 0.6 mm long; with curved outline; circular; marginal according to radicle Phylogenetic Number: 12.07. tip; flush; not within corona, halo, or rim. Lens not discernible. Endosperm thin; covering entire embryo; Tribe: Psoraleeae. adnate to testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of Species Studied—Species in Genus: 6 spp.—8 spp. equal length; not folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing Fruit a legume; unilocular; 0.3Ð1.2 ϫ 0.3Ð0.65 ϫ 0.3 cm; radicle; entire or split over radicle; without or with with persistent or deciduous calyx; with calyx shorter lobes; with the interface division terminating in radicle than fruit; without orifice formed by curving of fruit or tissue; without margins recessed; tan; inner face flat; fruit segments; straight; not plicate; not twisted; glabrous around base of radicle. Embryonic axis asymmetrical or symmetrical; circular or obovate; when deflexed; parallel to length of seed. Radicle linear; asymmetrical with 1 straight and 1 curved suture; deflexed and parallel to cotyledon length; centered widest near middle or D-shaped; not inflated; com- between cotyledons; less than 1/2 length of cotyledons. pressed; without or with beak; straight; with solid beak Plumule rudimentary; glabrous. the same color and texture as fruit; short tapered to rounded at apex; apex aligned (with 3Ð4 mm long beak) Distribution: North America and Mexico. or right-angled with longitudinal axis of fruit; short tapered to rounded at base; base aligned to oblique with Notes: Stirton (1981) noted that “Orbexilum is all that longitudinal axis of fruit; with the apex and base remains of Psoralea s.l.” when other segregate genera uniform in texture; coriaceous; seed chambers exter- including Psoralea s.s. (12.09) are removed. “Further- nally invisible. Fruit margin not constricted; without more, Orbexilum is clearly an unsatisfactory assem- sulcus; plain. Fruit wings absent. Fruit nonstipitate. blage of plants.” The species count and distribution are Fruit indehiscent. Replum invisible. Epicarp dull; based on Grimes (1990), who noted “Orbexilum is monochrome; brownish black, brown, or green; easily distinguished from all other Psoraleeae by its glabrous to glabrate to pubescent and indurate; with 1 rugose fruits.” Also, he classified fruits as rugose- type of pubescence; puberulent; with pubescence gray; ribbed, but in O. onobrychis (T. Nuttall) P.A. Rydberg with pubescence uniformly distributed; with simple fruits are papillate. hairs; pliable; with hair bases plain; glandular or eglandular; with glandular dots; without spines; not smooth; with elevated features; obliquely veined relative to fruit length or reticulately veined; not tuberculate; papillose or rugose; not exfoliating; without cracks. Mesocarp thin; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seed 1; length parallel with fruit length. Funiculus measured; less than 1 mm long; thick; straight. Aril absent.

Seed 2Ð7 ϫ 2Ð4 ϫ 2.5 mm; not overgrown; not angular; asymmetrical; ovate or reniform; compressed to terete (nearly); without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; black or brown (reddish); glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Orbexilum: O. pedunculatum (P. Miller) A.M. Vail (CÐE), Raphe not visible. Hilum visible; with faboid split; with O. spp. (AÐB). A, Fruits with and without calyx (ϫ 3.8); the lips of the faboid split the same color as the rest of B, seeds (ϫ 7.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

711 Genus: Otholobium C.H. Stirton smooth; with elevated features; heavily wrinkled; coriaceous. Fracture lines absent. Rim absent. Wings Phylogenetic Number: 12.08. absent. Raphe not visible. Hilum fully concealed; concealed by funicular remnant; with faboid split; with Tribe: Psoraleeae. the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.5 mm long; with Species Studied—Species in Genus: 15 spp.—ca. 45 spp. curved outline; circular; marginal according to radicle tip; flush; not within corona, halo, or rim. Lens not Fruit a legume; unilocular; 0.45Ð1.6 (including beak) ϫ discernible. Endosperm thin; covering entire embryo; 0.25Ð0.5 ϫ 0.25Ð0.3 cm; with persistent or deciduous adnate to embryo. Cotyledons smooth; both outer faces calyx; with calyx longer than to equal in length to convex; both the same thickness; both more or less of shorter than fruit; without orifice formed by curving of equal length; not folded; margin entire 180 degrees fruit or fruit segments; straight; not plicate; not twisted; from base of radicle; similar at apex; not concealing asymmetrical or symmetrical; circular, oblong, or radicle; split over radicle; with lobes; with the interface obovate; when asymmetrical with both sutures division terminating in radicle tissue; without margins parallelly curved; not inflated; compressed; with beak; recessed; yellow; inner face flat; glabrous around base straight; with solid beak (flat, 2 mm long) the same of radicle. Embryonic axis deflexed; parallel to length color and texture as fruit; short tapered to rounded at of seed. Radicle linear; deflexed and parallel to cotyle- apex; apex right-angled with longitudinal axis of fruit; don length; centered between cotyledons; 1/2 to nearly rounded at base; base aligned with longitudinal axis of length of cotyledons. Plumule rudimentary; glabrous. fruit; with the apex and base uniform or differing in texture; upper 1/2 inflated and reticulate over seed Distribution: Southeastern and eastern Africa into South cavity and lower 1/2 adnate and wrinkled to scurfy over Africa (zone of Mediterranean vegetation) and South seed cavity (O. glandulosa (C. Linnaeus) J.W. Grimes); America (8 spp.). chartaceous; seed chambers externally invisible. Fruit margin not constricted; without sulcus; plain. Fruit Notes: The species count and distribution are based on wings absent. Fruit nonstipitate. Fruit indehiscent. Grimes (1990), who noted 8 species in South America Replum invisible. Epicarp dull; monochrome; tan; and about 35 species in the Old World. Stirton (1986a) pubescent and indurate; with 1 type (gray) or 2 types of reviewed Otholobium and named two new species from pubescence; pilose or puberulent; with pubescence gray southeastern Africa (Stirton 1990). The Old World and brown or brown (dark); with appressed dark brown species need to be revised. Stirton noted that the hairs and scattered erect gray hairs intermixed; with “morphology of the fruiting calyx and fruit of O. pubescence uniformly distributed; with simple hairs; sericeum (J.L.M. Poiret) C.H. Stirton are quite different pliable; with hair bases plain; eglandular; without from that of the type species” (O. caffrum (C.F. Ecklon spines; not smooth; with elevated features; transversely & C.L.P. Zeyher) C.H. Stirton). veined relative to fruit length; not tuberculate; not exfoliating; with cracks; cracking transverse to fruit length. Mesocarp absent. Endocarp dull; monochrome; tan; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seed 1; length parallel with fruit length. Funiculus measured; less than 1 mm long; straight. Aril absent.

Seed 3Ð6 ϫ 2Ð3 ϫ 2 mm; not overgrown; not angular; asymmetrical; oblong; terete (to subterete); without visible radicle and cotyledon lobes; without umbo on seed faces. Testa partially adhering to endocarp (O. pubescens (J.L.M. Poiret) J.W. Grimes); nearly glossy; Otholobium: O. hirtum (C. Linnaeus) C.H. Stirton (CÐE), O. not modified by a bloom; colored; monochrome or spp. (AÐB). A, Fruits with and without calyx (ϫ 3.3); B, mottled; with frequent mottles; tannish olive or brown seeds some with adhering epicarp (ϫ 6); CÐD, testa (reddish); with black overlay; glabrous; smooth or not (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

713 Genus: Psoralea C. Linnaeus testa; darker than testa; black; within corona. Lens corona color lighter than testa. Endosperm thin; Phylogenetic Number: 12.09. covering entire embryo; adnate to testa. Cotyledons smooth; both outer faces convex; both the same Tribe: Psoraleeae. thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; similar Species Studied—Species in Genus: 7 spp.—ca. 20 spp. at apex; not concealing radicle; entire over radicle; without lobes; with the interface division terminating in Fruit a legume; unilocular; 0.3Ð0.6 (including beak) ϫ 0.2Ð radicle tissue; without margins recessed; yellow; inner 0.4 ϫ 0.15Ð 0.2 cm; with persistent calyx; with calyx face flat; glabrous around base of radicle. Embryonic longer than fruit; without orifice formed by curving of axis straight; parallel to length of seed. Radicle linear; fruit or fruit segments; straight; not plicate; not twisted; deflexed and parallel to cotyledon length; centered symmetrical; elliptic; not inflated; terete; with beak between cotyledons; less than 1/2 length of cotyledons. (0.7-0.8 mm long); straight, or declined; with solid beak Plumule rudimentary; glabrous. the same color and texture as fruit; rounded at apex; apex aligned with longitudinal axis of fruit; rounded at Distribution: South Africa. base; base aligned with longitudinal axis of fruit; with the apex and base uniform in texture; chartaceous or Notes: Stirton (1981) considered Psoralea to be an artificial fragile, thinner than chartaceous like Trifolium (21.06); assemblage and redefined it to include approximately seed chambers externally visible; with the raised seed 20 species, all endemic to South Africa. He accepted chambers not torulose. Fruit margin not constricted; the genus Hallia (12.09) with nine species but later without sulcus; plain. Fruit wings absent. Fruit (Tucker and Stirton 1991, Polhill 1994b, Crow et al. nonstipitate. Fruit indehiscent. Replum invisible. 1997) expressed the opinion that it is a subgenus of Epicarp dull; monochrome; gray or brown; with surface Psoralea, P. subgenus Hallia (C.P. Thunberg) T.M. texture uniform; glabrous or pubescent but soon Salter. Grimes (1990) concurred with Stirton and deciduous; with 1 type of pubescence; eglandular; transferred all New World Psoralea species to various without spines; not smooth; with elevated features; genera. irregularly veined (not aligned); not tuberculate; wrinkled; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; gray; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seed 1; length parallel with fruit length. Funiculus less than 0.5 mm long; straight. Aril dry; rim-aril; tan.

Seed 3Ð3.5 ϫ 1.7Ð2 ϫ 1.5 mm; not overgrown; not angular; symmetrical (except hilum); oblong; terete; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled; with frequent mottles; brown; with black overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum fully concealed; concealed by aril; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.6 mm long; with curved outline; elliptic; marginal according to radicle tip; flush; within corona or not within corona, halo, or rim. Hilum Psoralea: P. aphylla C. Linnaeus (DÐE), P. pinnata C. corona color lighter than testa. Lens discernible; equal Linnaeus (AÐC, F). A, Calyx with a fruit inside and a to or greater than 0.5 mm in length; 0.5 mm long; with cupulum (ϫ 4.6); B, fruit without calyx (ϫ 8.8); C, margins straight; linear; not in groove of raphe; seeds (ϫ 8); DÐE, testa (ϫ 50, ϫ 1000); F, embryos confluent with hilum; recessed; dissimilar color from (ϫ 10).

715 Loteae (13.01Ð13.17) brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Genus: Cytisopsis H.F. Jaubert & É. Spach Hilum fully concealed; concealed by funicular remnant; with faboid split; the same color as the rest of the Phylogenetic Number: 13.01. hilum; punctiform; apical at apex of radicle tip; flush; not within corona, halo, or rim. Lens discernible or not Tribe: Loteae. discernible; equal to or greater than or less than 0.5 mm in length; 0.5 mm long; with margins curved; elliptic; Species Studied—Species in Genus: 1 sp.—ca. 2 spp. not in groove of raphe; adjacent to hilum; 0.3 mm from hilum; mounded; dissimilar color from testa; darker Fruit a legume; unilocular; 1Ð2.2 ϫ 0.4Ð0.5 ϫ 0.4Ð0.5 cm; than testa; black; not within corona, halo, or rim. with persistent calyx; with calyx shorter than fruit; Endosperm thin; not pluglike and resembling tip of without orifice formed by curving of fruit or fruit radicle; covering entire embryo; adnate to embryo and segments; straight; not plicate; not twisted; symmetri- testa. Cotyledons smooth; both outer faces convex; both cal; oblong; not inflated; terete; without beak; short the same thickness; both more or less of equal length; tapered at apex; apex aligned with longitudinal axis of not folded; margin entire 180 degrees from base of fruit; short tapered at base; base aligned with longitudi- radicle; similar at apex; not concealing radicle; entire nal axis of fruit; with the apex and base uniform in over radicle; without lobes; with the interface division texture; ligneous; seed chambers externally visible or terminating at base of radicle; without margins re- invisible; with the raised seed chambers not torulose. cessed; tan or green; inner face flat; glabrous around Fruit margin not constricted or constricted; constricted base of radicle. Embryonic axis deflexed; oblique to along both margins; without sulcus; plain. Fruit wings length of seed; with a joint evident between the radicle absent. Fruit substipitate. Fruit with all layers dehiscing and the cotyledons. Radicle linear; lobe tip straight; (to tardily so); splitting along sutures. Dehiscence of deflexed and parallel to cotyledon length; centered valves along both sutures; apical and down; passive or between cotyledons; 1/2 to nearly length of cotyledons. active; with valves twisting. Replum invisible. Epicarp Plumule rudimentary; glabrous. dull; monochrome; reddish brown; pubescent and indurate; with hairs appressed; with 1 type of pubes- Distribution: Eastern Mediterranean (1 sp.) and perhaps in cence; with pubescence gray; with pubescence uni- north Africa (1 sp.). formly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; Notes: In 1981, Polhill (1981k) accepted much broader with elevated features; somewhat reticulately veined; generic circumscriptions in tribe Loteae and only not tuberculate; wrinkled; not exfoliating; without accepted four genera in the tribe: Cytisopsis, Anthyllis cracks. Mesocarp thick; 1-layered; without balsamic (13.02), Hymenocarpus (13.04), and Lotus (13.07). In vesicles; without fibers; solid; ligneous. Endocarp dull; his most recent classification of Fabaceae (Polhill monochrome or mottled; tan; with mottling above and 1994a,b), he combined the tribes Loteae and below seed chambers; with brown (reddish) overlay; Coronilleae and accepted six segregate genera in smooth; subseptate; with septa thin (tissue paper-like), Loteae, s.s.: Tripodion (13.03), Dorycnopsis (13.05), flexible; with septa eglandular; chartaceous; not Dorycnium (13.06), Podolotus J.F. Royle (13.08), exfoliating; remaining fused to mesocarp and epicarp; Pseudolotus K.H. Rechinger (13.09), and Vermifrux entire. Seeds 2Ð6; length transverse to fruit length; (13.10). Polhill (1981k) noted that the status of the neither overlapping nor touching; in 1 series. Funiculus North African species of Cytisopsis, which also has less than 0.5 mm long; of 1 length only; thick; straight. been placed in the segregate genus Lyauteya R.C.J.E. Aril absent. Maire, is not certain and that perhaps both species of Cytisopsis are better placed in Anthyllis (13.02). We Seed 2.2Ð2.8 ϫ 2 ϫ 2 mm; not overgrown; not angular or studied only the eastern Mediterranean species C. angular; asymmetrical; circular, oblong, or irregular; pseudocytisus. terete; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on Cytisopsis: C. pseudocytisus (P.E. Boissier) Fertig (AÐE). seed faces. Testa not adhering to endocarp; dull; not A, Fruit and valve (ϫ 4); B, seeds (ϫ 7); CÐD, testa modified by a bloom; clear; monochrome; dark reddish (ϫ 50, ϫ1000); E, embryos (ϫ 10).

717 Genus: Anthyllis C. Linnaeus hilum); oblong, mitaform, or triangular; compressed or terete; with surface smooth; without visible radicle and Phylogenetic Number: 13.02. cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not Tribe: Loteae. modified by a bloom; colored; monochrome or mottled and streaked; with frequent mottles; with frequent Species Studied—Species in Genus: 19 spp.—20–25 spp. streaks; brown, tan, or green; with purple overlay; glabrous; smooth or not smooth; with elevated features; Fruit a legume; unilocular; 0.4Ð1.3 ϫ 0.2Ð0.5 ϫ 0.1Ð0.2 shagreen, wrinkled, or tuberculate; coriaceous. Fracture cm; with deciduous or persistent calyx; with calyx lines absent. Rim absent. Wings absent. Raphe not longer than fruit; without or with orifice formed by visible. Hilum visible; with faboid split; with the lips of curving of fruit or fruit segments; straight or 1-coiled; the faboid split the same color as the rest of the hilum; not plicate; not twisted; symmetrical or asymmetrical; punctiform; between cotyledon and radicle lobe; flush; linear or coiled; when asymmetrical with both sutures within rim. Hilum rim color slightly darker than testa. parallelly curved or 1 straight and 1 curved suture; Lens discernible; less than 0.5 mm in length; with widest near middle or D-shaped; inflated; compressed margins straight or curved; wedge-shaped; elliptic, or or terete; without or with beak; with solid beak the hourglass or dumbbell-shaped; not in groove of raphe; same color and texture as fruit; short tapered or confluent with hilum; mounded; dissimilar color from rounded at apex; apex oblique with longitudinal axis of testa; darker than testa; reddish brown; not within fruit; tapered or rounded at base; base right angled with corona, halo, or rim. Endosperm thin; covering entire longitudinal axis of fruit; with the apex and base embryo; adnate to testa. Cotyledons smooth; both outer uniform in texture; coriaceous or fragile, thinner than faces convex; both the same thickness; both more or chartaceous like Trifolium (21.06) (area of “dehis- less of equal length; not folded; margin entire 180 cences”); seed chambers externally visible; with the degrees from base of radicle; similar at apex; not raised seed chambers not torulose. Fruit margin not concealing radicle; entire over radicle; without lobes; constricted or constricted; slightly constricted along with the interface division terminating at base of both margins; without sulcus; plain or embellished. radicle; without margins recessed; tan; inner face flat; Fruit wing absent or present; 1; 0.5 mm wide; sutural; glabrous around base of radicle. Embryonic axis on 1 suture. Fruit nonstipitate or stipitate. Fruit indehis- deflexed; oblique to length of seed; without a joint cent or with all layers dehiscing (tardily); opening by evident between the radicle and the cotyledons. Radicle deterioration of delicate strip of tissue along inner linear; lobe tip straight; deflexed and parallel to suture or both sutures. Dehiscence of valves along 1 cotyledon length; centered between cotyledons; less suture; medial and up and down; passive. Replum than 1/2 length or 1/2 to nearly length of cotyledons. invisible. Epicarp dull or semiglossy; monochrome; Plumule rudimentary; glabrous. brown or tan; with surface texture uniform or not uniform, with patches of different texture not restricted Distribution: Mediterranean region, extending into Europe, to the base and apex; glabrous; eglandular; without Atlantic islands, and North Africa. spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Notes: Polhill (1981k) noted that Anthyllis and Lotus (13.07) Mesocarp thin; surface not veined; 1-layered; without seem to be closely related to Hammatolobium (13.17). balsamic vesicles; without fibers; solid; coriaceous. Polhill also noted that Cytisopsis (13.01) may be Endocarp dull; monochrome; tan; smooth; nonseptate combined with Anthyllis. Tripodion (13.03), previously or septate; with septa thin (tissue paper-like), flexible; A. tetraphylla C. Linnaeus, was not discussed by chartaceous; not exfoliating; remaining fused to Polhill. Anthyllis vulneraria C. Linnaeus is a variable mesocarp and epicarp; entire. Seeds 1Ð3; length parallel species with many subspecies and some economic with fruit length; neither overlapping nor touching; in 1 value. Some species, but not A. vulneraria, possess series. Funiculus less than 0.5 mm long; of 1 length fruits with unusual dehiscences. Either the lower or only; filiform; straight. Aril absent. both sutural areas have much thinner fruit tissue, and this tissue easily ruptures gradually exposing the seeds Seed 1.7Ð3.5 ϫ 1Ð3 ϫ 1Ð2 mm; not overgrown; not angular in situ. Eventually the seeds fall free from one or both or angular; asymmetrical or symmetrical (except valves.

718 719 Anthyllis: A. hermanniae C. Linnaeus (CÐE), A. spp. (AÐB). A, Fruits (ϫ 3); B, seeds (ϫ 7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

721 Genus: Tripodion F.C. Medikus bloom; colored; monochrome or streaked and mottled; with frequent mottles; with frequent streaks; reddish Phylogenetic Number: 13.03. brown or tan; with black overlay; glabrous; not smooth (and distinctly grooved between cotyledons especially Tribe: Loteae. opposite from hilum); with elevated features; tuberculate; coriaceous. Fracture lines absent. Rim absent. Species Studied—Species in Genus: 1 sp.—1 sp. Wings absent. Raphe not visible. Hilum partially concealed; concealed by funicular remnant; with faboid Fruit a legume; unilocular; 0.9Ð1 ϫ 0.3Ð0.4 ϫ 0.2 cm; with split; with the lips of the faboid split the same color as persistent calyx; with calyx longer than fruit (and the rest of the hilum; punctiform; marginal according to inflated); without orifice formed by curving of fruit or radicle tip or between cotyledon and radicle lobe; fruit segments; straight; not plicate; not twisted; recessed; not within corona, halo, or rim. Lens obvi- asymmetrical; oblong; when asymmetrical with both ously or faintly discernible or not discernible; less than sutures unequally or parallelly curved; not inflated (but 0.5 mm in length; with margins curved; circular or calyx inflated); terete; with beak (1 mm long); with elliptic; not in groove of raphe; adjacent to hilum; 0.5 solid beak the same color and texture as fruit; short mm from hilum; mounded; same color as, similar color tapered at apex; apex aligned with longitudinal axis of as, or dissimilar color from testa; lighter than or darker fruit; rounded at base; base aligned with longitudinal than testa; reddish brown, tan (reddish), or black; not axis of fruit; with the apex and base uniform in texture; within corona, halo, or rim. Endosperm thick; covering chartaceous or fragile, thinner than chartaceous like entire embryo; adnate to embryo. Cotyledons smooth; Trifolium (21.06); seed chambers externally visible; both outer faces convex; both the same thickness; both with the raised seed chambers torulose. Fruit margin more or less of equal length; not folded; margin entire constricted along both margins; without sulcus; plain. 180 degrees from base of radicle; similar at apex; not Fruit wings absent. Fruit substipitate. Fruit indehiscent concealing radicle; entire over radicle; without lobes; (but fragile enough to open easily). Replum invisible. with the interface division terminating at base of Epicarp dull; multicolored; streaked (because streaked radicle; without margins recessed; tan; inner face flat; endocarp is seen through epicarp); tan; with brown glabrous around base of radicle. Embryonic axis (reddish) overlay; with mottling over seed chambers; deflexed; oblique to length of seed; without a joint pubescent and indurate; with 1 type of pubescence; evident between the radicle and the cotyledons. Radicle puberulent; with pubescence golden or gray; with linear; deflexed and parallel to cotyledon length; pubescence uniformly distributed; with simple hairs; centered between cotyledons; less than 1/2 length of pliable; with hair bases plain; straight; eglandular; cotyledons. Plumule rudimentary; glabrous. without spines; smooth or not smooth; veined or not veined; reticulately veined (some best seen from Distribution: Mediterranean region, southern Portugal. endocarp view); not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; Notes: This genus was part of Anthyllis (13.02) in the report streaked; tan; with streaking above and below seed of Polhill (1981k), but Tripodion is now recognized as a chambers; with brown (reddish) overlay; smooth; separate genus (Polhill 1994a,b). Previously, Tripodion septate; with septa thin (tissue paper-like), flexible; was also known as Physanthyllis P.E. Boissier. The with septa eglandular; chartaceous; not exfoliating; testa fragments during imbibition as does that of remaining fused to epicarp; entire. Seeds (1Ð)2; length Cyamopsis (9.06). parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 3Ð3.5 ϫ 2.8Ð3.2 ϫ 1.5Ð2.5 mm; not overgrown; not angular or angular (to somewhat); asymmetrical; oblong or reniform; compressed; with surface smooth; without hilar sinus; without umbo on seed faces. Testa without or with pieces of adhering epicarp (occasion- Tripodion: T. teraphyllum (C. Linnaeus) J.P. Fourreau ally); not adhering to endocarp; dull; not modified by a (AÐE). A, Fruits and fruiting calyx (ϫ 1.25); B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 15).

723 Genus: Hymenocarpos C.G. Savi and radicle lobe; somewhat recessed; not within corona, halo, or rim (if testa light colored) or within halo (if Phylogenetic Number: 13.04. testa dark colored). Hilum halo color darker than testa. Lens discernible; less than 0.5 mm in length; with Tribe: Loteae. margins curved; elliptic; not in groove of raphe; confluent with hilum; mounded; dissimilar color from Species Studied—Species in Genus: 1 sp.—1 sp. testa; darker than testa; black; not within corona, halo, or rim. Endosperm absent. Cotyledons smooth; both Fruit a legume; unilocular; 2Ð2.5 ϫ 0.6Ð0.8 ϫ 0.25 cm; outer faces convex; both the same thickness; both more with persistent calyx; with calyx shorter than fruit; with or less of equal length; not folded; margin entire 180 orifice formed by curving of fruit or fruit segments; 1- degrees from base of radicle; similar at apex; not coiled; not plicate; not twisted; asymmetrical; coiled; concealing radicle; entire over radicle; without lobes; when asymmetrical with both sutures parallelly curved; with the interface division terminating at base of not inflated; flattened; without beak; truncate at apex; radicle; without margins recessed; light tan; inner face apex right-angled with longitudinal axis of fruit; flat; glabrous around base of radicle. Embryonic axis truncate at base; base right angled with longitudinal deflexed; oblique to length of seed; with a joint evident axis of fruit; with the apex and base uniform in texture; between the radicle and the cotyledons. Radicle linear; membranous; seed chambers externally invisible. Fruit deflexed and parallel to cotyledon length; centered margin not constricted; without sulcus; embellished; between cotyledons; less than 1/2 length of cotyledons. with wing (margins either entire, dentate, or aculeate). Plumule rudimentary; glabrous. Fruit wing 1; 3 mm wide; sutural; on 1 suture. Fruit nonstipitate. Fruit indehiscent. Replum invisible. Distribution: Mediterranean region and western Asia. Epicarp dull; monochrome (to irregularly brown tinged with purple); brown, tan, or gray and purple; glabrous, pubescent and indurate, or pubescent but soon deciduous; with hairs appressed; with 1 type of pubescence; with pubescence golden; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp present (restricted only to seed chamber); thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; tan; smooth; septate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 1Ð3 ϫ 2Ð2.5 ϫ 1 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown, tan, cream, or yellow; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Hymenocarpos: H. circinnatus (C. Linnaeus) C.G. Savi fully concealed; concealed by funicular remnant; (AÐE). A, Fruits (ϫ 2); B, seeds (ϫ 6); CÐD, testa without faboid split; punctiform; between cotyledon (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

725 Genus: Dorycnopsis P.E. Boissier reddish brown; not within corona, halo, or rim. En- dosperm thin; covering entire embryo; adnate to testa. Phylogenetic Number: 13.05. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; not Tribe: Loteae. folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing radicle; entire over Species Studied—Species in Genus: 1 sp.—1 sp. radicle; without lobes; with the interface division terminating at base of radicle; without margins re- Fruit a legume; unilocular; 0.2Ð0.23 ϫ 0.15 ϫ 0.1 cm; with cessed; yellow; inner face flat; glabrous around base of deciduous calyx (if fruits in a head then longer than radicle. Embryonic axis deflexed; oblique to length of calyx); without orifice formed by curving of fruit or seed; without a joint evident between the radicle and fruit segments; straight; not plicate; not twisted; the cotyledons. Radicle linear; deflexed and parallel to asymmetrical; elliptic; when asymmetrical with both cotyledon length; centered between cotyledons; less sutures parallelly curved; not inflated; terete; without than 1/2 length of cotyledons. Plumule rudimentary; beak; rounded at apex; apex aligned with longitudinal glabrous. axis of fruit; rounded at base; base aligned with longitudinal axis of fruit; with the apex and base Distribution: Spain and southwestern Europe. uniform in texture; coriaceous; seed chambers externally invisible. Fruit margin not constricted; without Notes: This genus was part of Anthyllis (13.02) in the report sulcus; embellished. Fruit wing 1; 0.3 mm wide; of Polhill (1981k), but Dorycnopsis is now recognized sutural; on 1 suture. Fruit nonstipitate. Fruit indehiscent as a separate genus (Polhill 1994a,b). Tikhomirov and (difficult to remove seed from fruit even though fruit Sokoloff (1997) examined the taxonomic status of coat thin). Replum invisible. Epicarp dull; mono- Vermifrux (13.10) abyssinica (A. Richard) J.B. Gillett chrome; brown; glabrous; eglandular; without spines; and concluded that Vermifrux and Dorycnopsis are not smooth; with elevated features; reticulately veined; synonymous. Therefore they made the new combina- not tuberculate; not exfoliating; without cracks. tion D. abyssinica (A. Richard) V.N. Tikhomirov & Mesocarp thin; surface not veined; 1-layered; without D.D. Sokoloff for the only species of Vermifrux. balsamic vesicles; without fibers; solid; coriaceous. Pending further evaluation, we are accepting Vermifrux. Endocarp glossy; monochrome; reddish brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1(Ð2) (though 2-seeded fruits not seen); length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 1.5 ϫ 1 ϫ 0.8 mm; not overgrown; not angular; symmetrical (except hilum); elliptic; terete; with surface smooth; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum fully concealed; concealed by funicular remnant; with faboid split; the same color as the rest of the hilum; punctiform; marginal according to radicle tip; flush; within rim. Hilum rim color darker than testa. Lens discernible; less than 0.5 mm in length; with margins straight or curved; oblong; not in groove of raphe; adjacent to hilum; 0.1 mm from hilum; Dorycnopsis: D. gerardii (C. Linnaeus) P.E. Boissier (AÐE). mounded; dissimilar color from testa; darker than testa; A, Fruits (ϫ 1.25); B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 15).

727 Genus: Dorycnium P. Miller glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum visible or fully Phylogenetic Number: 13.06. concealed; concealed by funicular remnant; with faboid split; with the lips of the faboid split the same color as Tribe: Loteae. the rest of the hilum; punctiform; marginal according to radicle tip or between cotyledon and radicle lobe; flush; Species Studied—Species in Genus: 7 spp.—ca. 12 spp. not within corona, halo, or rim or within halo. Hilum halo color darker than or lighter than testa. Lens Fruit a legume; unilocular; 0.3Ð2 ϫ 0.2Ð1 ϫ 0.2Ð0.32 cm; discernible or not discernible; equal to or greater than with deciduous calyx; without orifice formed by or less than 0.5 mm in length; 0.5 mm long; with curving of fruit or fruit segments; straight; not plicate; margins curved; key-hole shaped; not in groove of not twisted; asymmetrical or symmetrical; oblong, raphe; adjacent to hilum; 0.1 mm from hilum; obovate, ovate, or circular; when asymmetrical with mounded; dissimilar color from testa; darker than testa; both sutures nearly straight or parallelly curved; not black; not within corona, halo, or rim. Endosperm inflated; terete; without or with beak; straight; with thick; covering entire embryo; adnate to embryo. solid beak the same color and texture as fruit; rounded Cotyledons smooth; both outer faces convex; both the or short tapered at apex; apex aligned or oblique with same thickness; both more or less of equal length; not longitudinal axis of fruit; rounded at base; base aligned folded; margin entire 180 degrees from base of radicle; or oblique with longitudinal axis of fruit; with the apex similar at apex; not concealing radicle; entire over and base uniform in texture; coriaceous or ligneous; radicle; without lobes; with the interface division seed chambers externally invisible. Fruit margin not terminating at base of radicle; without margins re- constricted; without sulcus; plain. Fruit wings absent. cessed; tan or white; inner face flat; glabrous around Fruit substipitate. Fruit with all layers dehiscing; base of radicle. Embryonic axis deflexed; oblique to splitting along sutures. Dehiscence of valves along both length of seed; with a joint evident between the radicle sutures; apical and down; active or passive; with valves and the cotyledons. Radicle linear or bulbose (some- twisting or enrolling. Replum invisible. Epicarp dull; what); deflexed and parallel to cotyledon length; monochrome; light to dark reddish brown; glabrous or centered between cotyledons; less than 1/2 length of pubescent and indurate; pilose; with simple hairs; cotyledons. Plumule rudimentary; glabrous. eglandular; without spines; not smooth; with elevated features; not veined; not tuberculate; wrinkled; not Distribution: Central and southern Europe and Mediterra- exfoliating; without cracks. Mesocarp thin; surface not nean region. veined; 1-layered; without balsamic vesicles; without fibers; solid; subligneous. Endocarp glossy; mono- Notes: This genus was part of Lotus (13.07) in the report of chrome or mottled; reddish brown or tan (greenish); Polhill (1981k), but Dorycnium has often been recog- with mottling (dark); with brown (reddish) overlay; nized as a separate genus with two sections: Bonjeania smooth; septate or nonseptate; with septa thin (tissue (H.G.L.R. Reichenbach) P.E. Boissier with 2Ð8-seeded paper-like), flexible; with septa eglandular; legumes and Dorycnium with 1-seeded legumes. chartaceous; exfoliating in part to exfoliating; separating from mesocarp; entire. Seeds 1Ð8; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 1.3Ð2 ϫ 1Ð1.6 ϫ 1Ð1.4 mm; not overgrown; not angular; symmetrical or asymmetrical; elliptic, mitaform, or circular; terete; with surface smooth; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or Dorycnium: D. hirsutum (C. Linnaeus) N.C. Seringe (CÐE), mottled; with frequent mottles; brown to yellowish or D. spp. (AÐB). A, Fruits (dehisced and closed) (ϫ 1.25); reddish brown or green (to tannish); with black overlay; B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 15).

729 Genus: Lotus C. Linnaeus Seed 0.8Ð5 ϫ 0.7Ð5 ϫ 0.3Ð5 mm; not overgrown; not angular or angular; symmetrical or asymmetrical; Phylogenetic Number: 13.07. circular, mitaform, irregular, quadrangular, or triangular; terete, compressed, or quadrangular; with surface Tribe: Loteae. smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa Species Studied—Species in Genus: 62 spp.—ca. 100 spp. not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome or mottled and Fruit a legume; unilocular; 2.5Ð9 ϫ 0.2Ð0.8 ϫ 0.2Ð0.3 cm; streaked; with frequent mottles; with frequent streaks; with deciduous or persistent calyx; with calyx shorter dark brown, tan (to reddish), green, yellow, or purple; than fruit; without orifice formed by curving of fruit or with black or purple overlay; glabrous; smooth or not fruit segments; straight or curved (or slightly curved); smooth; with elevated features; tuberculate; coriaceous. not plicate; not twisted; symmetrical or asymmetrical; Fracture lines absent. Rim absent. Wings absent. Raphe linear, oblong, or falcate; when asymmetrical with both not visible. Hilum partially or fully concealed; con- sutures parallelly curved or nearly straight; not inflated; cealed by funicular remnant; with faboid split; with the terete or compressed; without beak; short tapered or lips of the faboid split the same color as the rest of the rounded at apex; apex aligned with longitudinal axis of hilum; punctiform; marginal according to radicle tip or fruit; short tapered or rounded at base; base aligned between cotyledon and radicle lobe; flush; within rim or with longitudinal axis of fruit; with the apex and base halo. Hilum halo color darker than testa. Hilum rim uniform in texture; coriaceous; seed chambers exter- color darker than testa. Lens discernible; less than 0.5 nally visible or invisible; with the raised seed chambers mm in length; with margins straight or curved; linear; not torulose or torulose. Fruit margin not constricted or circular or elliptic; not in groove of raphe; adjacent to constricted; slightly constricted along both margins; or confluent with hilum; 0.2Ð0.4 mm from hilum; without sulcus; plain or embellished. Fruit wings absent mounded; dissimilar color from or similar color as or present (section Tetragonolobus); 2 or 4 (with 2 testa; darker than testa; black; not within corona, halo, wings on long upper suture and none below or 2 wings or rim. Endosperm thin or thick; covering entire along both sutures); 1Ð2 mm wide; sutural; on both embryo; adnate to embryo or testa. Cotyledons smooth; sutures or 1 suture. Fruit nonstipitate. Fruit with all both outer faces convex; both the same thickness; both layers dehiscing or indehiscent (rarely); splitting along more or less of equal length; not folded; margin entire sutures. Dehiscence of valves along both sutures; apical 180 degrees from base of radicle; similar at apex; not and down; passive or active; with valves twisting or concealing radicle; entire over radicle; without lobes; enrolling. Replum invisible. Epicarp dull; monochrome; with the interface division terminating at base of reddish brown; glabrous or pubescent and indurate; radicle; without margins recessed; green, tan, or yellow; with hairs appressed; with 1 type of pubescence; with inner face flat; glabrous around base of radicle. Embry- pubescence golden; with pubescence uniformly onic axis deflexed; oblique to length of seed; without a distributed; with simple hairs; pliable; with hair bases joint evident between the radicle and the cotyledons. plain; eglandular; without spines; not smooth or Radicle linear; deflexed and parallel to cotyledon smooth; with elevated features; veined or not veined; length; centered between cotyledons; 1/2 to nearly reticulately veined; not tuberculate; not exfoliating or length of cotyledons. Plumule rudimentary; glabrous. checking (of cuticle); without or with cracks (of cuticle); cracking oblique to fruit length. Mesocarp thin; Distribution: North America and extratropical South surface not veined; 1-layered; without balsamic America, Europe, Russia, extending into Africa and vesicles; without fibers; solid; coriaceous. Endocarp Australia. dull; monochrome; tan; smooth; septate; with septa thin (tissue paper-like), flexible; with septa eglandular; Notes: Polhill (1981k) noted that Lotus and Anthyllis (13.02) chartaceous; not exfoliating; remaining fused to seem to be closely related to Hammatolobium (13.17). mesocarp and epicarp; entire. Seeds 1Ð18; length In our treatment, Tetragonolobus G.A. Scopoli is parallel with fruit length; neither overlapping nor included in Lotus even though the species of section touching; in 1 or 2 or more series (section Tetrago- Tetragonolobus have winged fruits and generally larger nolobus). Funiculus less than 0.5 mm long; of 1 length seeds than species in other sections of Lotus. Lassen only; triangular or thick; straight. Aril absent. (1986) transferred L. roudairei E. Bonnet of North

730 Africa to Acmispon (13.07A) because of its glandular stipules and 5Ð6 leaflets per leaf. Kramina and Sokoloff (1997) reexamined L. roudairei and concluded that it is not related to New World taxa and should remain in Lotus. They recognized its unique features among Old World Lotus by placing it in the new section Lotus sect. Pseudosimpeteria T.E. Kramina & D.D. Sokoloff. We left it in Lotus pending further study.

731 Lotus: L. corniculatus C. Linnaeus (DÐF), L. maritimus C. Linnaeus (B), L. spp. (A, C). A, Fruits (ϫ 1.25); B, fruit cross section with four wings (ϫ 10); C, seeds (ϫ 4); DÐE, testa (ϫ 50, ϫ 1000); F, embryos (ϫ 15).

733 Genus: Acmispon C.S. Rafinesque-Schmaltz not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled and streaked Phylogenetic Number: 13.07A. (both faintly); with frequent mottles; with frequent streaks; reddish brown or tan (greenish); with brown Tribe: Loteae. (darker reddish) overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe Species Studied—Species in Genus: 5 spp.—ca. 10 spp. not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the Fruit a legume; unilocular; 0.8Ð3 ϫ 0.1Ð0.3 ϫ 0.1 cm; with hilum; punctiform; between cotyledon and radicle lobe; persistent calyx; with calyx shorter than fruit; without barely recessed; within halo. Hilum halo color darker orifice formed by curving of fruit or fruit segments; than testa. Lens discernible; equal to or greater than or straight or curved (or slightly curved); not plicate; not less than 0.5 mm in length; 0.5 mm long; with margins twisted; symmetrical; linear; not inflated; compressed curved; elliptic or 2 circular mounds separated by or terete; without or with beak; hooked; tapered at apex; groove; not in groove of raphe; adjacent to hilum; 0.2 apex oblique with longitudinal axis of fruit; short mm from hilum; mounded; dissimilar color from testa; tapered at base; base aligned with longitudinal axis of darker than testa; dark reddish brown; not within fruit; with the apex and base uniform in texture; corona, halo, or rim. Endosperm thin or thick; covering coriaceous. Fruit margin not constricted; without entire embryo; adnate to embryo or testa. Cotyledons sulcus; plain. Fruit wings absent. Fruit nonstipitate. smooth; both outer faces convex; both the same Fruit with all layers dehiscing; splitting along sutures. thickness; both more or less of equal length; not folded; Dehiscence of valves along both sutures; apical and margin entire 180 degrees from base of radicle; similar down; active; with valves twisting. Replum invisible. at apex; not concealing radicle; entire over radicle; Epicarp dull or semiglossy; monochrome; reddish without lobes; with the interface division terminating at brown; glabrous or pubescent and indurate; with 1 type base of radicle; without margins recessed; tan, yellow, of pubescence; pilose or villous; with pubescence gray or green; inner face flat; glabrous around base of or golden; with pubescence uniformly distributed; radicle. Embryonic axis deflexed; oblique to length of pliable; with hair bases plain; eglandular or glandular; seed; without a joint evident between the radicle and with glandular dots (in herringbone pattern on valve); the cotyledons. Radicle linear; lobe tip straight; without spines; not smooth; with elevated features; deflexed and parallel to cotyledon length; centered or reticulately veined; not tuberculate; dotted or wrinkled; not centered between cotyledons (radicle outside 1 not exfoliating; without cracks. Mesocarp thin; surface cotyledon and inside other, therefore junctions for each not veined; 1-layered; without balsamic vesicles; cotyledon different); less than 1/2 length or 1/2 to without fibers; solid; coriaceous. Endocarp dull; nearly length of cotyledon. Plumule rudimentary; monochrome; tan; smooth (below and around seed and glabrous. cobweb between “seed chambers” or smooth throughout); septate; with septa thin (tissue paper-like), flexible Distribution: North America, Mexico, South America. (each seed in individual “seed chamber” with 2 septa between seeds or with 1 thin septum between each Notes: Lassen (1986) transferred Lotus roudairei E. Bonnet seed); with septa eglandular; chartaceous; not exfoliat- of North Africa to Acmispon because of its glandular ing; remaining fused to mesocarp and epicarp; entire. stipules and 5Ð6 leaflets per leaf. Kramina and Sokoloff Seeds 5Ð10; length parallel with fruit length; neither (1997) reexamined L. roudairei and concluded that it is overlapping nor touching; in 1 series. Funiculus less not related to New World taxa and should remain in than 0.5 mm long; of 1 length only; filiform; curved. Lotus. Aril absent.

Seed 1Ð2.7 ϫ 1Ð1.5 ϫ 0.6Ð0.8 mm; not overgrown; not angular or angular; asymmetrical; circular, oblong, or mitaform; compressed; with surface smooth; with or Acmispon: A. americanus (T. Nuttall) P.A. Rydberg (CÐE); without visible radicle and cotyledon lobes; without A. spp. (AÐB). A, Fruits (dehisced and closed) (ϫ 1.5); external groove between radicle and cotyledon lobes; B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos without hilar sinus; without umbo on seed faces. Testa (ϫ 15).

735 Genus: Vermifrux J.B. Gillett within corona, halo, or rim. Endosperm thin; covering entire embryo; adnate to testa. Cotyledons smooth; both Phylogenetic Number: 13.10. outer faces convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 Tribe: Loteae. degrees from base of radicle; similar at apex; not concealing radicle; entire over radicle; without lobes; Species Studied—Species in Genus: 1 sp.—1 sp. with the interface division terminating at base of radicle; brown; inner face flat; glabrous around base of Fruit a legume; unilocular; 0.4Ð0.6 ϫ 0.15Ð0.17 ϫ 0.15Ð0.2 radicle. Embryonic axis deflexed; oblique to length of cm; with deciduous or persistent calyx (rarely); with seed; with a joint evident between the radicle and the calyx shorter than fruit; with or without orifice formed cotyledons. Radicle somewhat bulbose or linear; by curving of fruit or fruit segments; 1-coiled, 0.5- deflexed and parallel to cotyledon length; centered coiled, or curved; not plicate; not twisted; asymmetri- between cotyledons; less than 1/2 length of cotyledons. cal; coiled or C-shaped; when asymmetrical with both Plumule rudimentary; glabrous. sutures parallelly curved; not inflated; terete; without beak; short tapered at apex; apex oblique with longitu- Distribution: Africa (Yemen, Sudan, Eritrea, Ethiopia). dinal axis of fruit; short tapered at base; base oblique with longitudinal axis of fruit; with the apex and base Notes: This genus was part of Lotus (13.07) in the report of uniform in texture; coriaceous; seed chambers exter- Polhill (1981k), but Vermifrux often has been recog- nally invisible or visible (faintly). Fruit margin not nized as a separate genus. Tikhomirov and Sokoloff constricted; without sulcus; embellished. Fruit wing (1997) examined the taxonomic status of Vermifrux (very narrow) 1; 0.3 mm wide; sutural; on 1 suture. abyssinica and concluded that Vermifrux and Fruit nonstipitate. Fruit indehiscent. Replum invisible. Dorycnopsis (13.05) are synonymous. Therefore they Epicarp glossy; monochrome; reddish brown or tan; made the new combination D. abyssinica (A. Richard) glabrous; eglandular; without spines; not smooth; with V.N. Tikhomirov & D.D. Sokoloff for the only species elevated features; reticulately veined; not tuberculate; of Vermifrux. Pending further evaluation, we are not exfoliating; without cracks. Mesocarp thin; surface accepting Vermifrux. not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; reddish brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 1.8Ð2 ϫ 0.9Ð1 ϫ 0.7Ð0.8 mm; not overgrown; not angular; asymmetrical; C-shaped; terete; with surface smooth; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; dark reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum fully concealed; concealed by funicular remnant; without faboid split; punctiform; marginal according to radicle tip; flush; within rim. Hilum rim color slightly darker than testa. Lens discernible; less than 0.5 mm in length; with margins curved; elliptic; not in groove of raphe; adjacent to hilum; 0.1 mm from hilum; mounded; Vermifrux: V. abyssinicum (A. Richard) J.B. Gillett (AÐE). dissimilar color from testa; darker than testa; black; not A, Fruits (ϫ 1.25); B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 15).

737 Genus: Coronilla C. Linnaeus or without (Schmidt 1979a) elevated features; with 1 longitudinal ridge on each face; coriaceous. Fracture Phylogenetic Number: 13.11. lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible or fully concealed; concealed by Tribe: Loteae. funicular remnant; with or without faboid split; with the lips of the faboid split the same color as the rest of the Species Studied—Species in Genus: 7 spp.—9 spp. hilum; punctiform; marginal according to radicle tip; flush or recessed (within depression or not); not within Fruit a loment (or a loment segment); 1Ð5 ϫ 0.1Ð0.3 ϫ 0.1Ð corona, halo, or rim. Lens discernible (or barely 0.25 cm; with deciduous corolla; with persistent calyx; discolored or darker than testa); less than 0.5 mm in with calyx shorter than fruit; without orifice formed by length; with margins straight; wedge-shaped; not in curving of fruit or fruit segments; straight to curved; not groove of raphe; adjacent to hilum; mounded; dissimi- plicate; not twisted; symmetrical; linear or moniliform; lar color from testa; darker than testa; black or brown; not inflated; compressed or terete; with beak; straight; not within corona, halo, or rim. Endosperm thick; with solid beak the same color and texture as fruit; short covering entire embryo; adnate to testa or embryo. tapered at apex; apex aligned with longitudinal axis of Cotyledons smooth; both outer faces convex; both the fruit; short tapered at base; base aligned with longitudi- same thickness; both more or less of equal length; not nal axis of fruit; with the apex and base uniform or folded; margin entire 180 degrees from base of radicle; differing in texture; upper 3/4 barely inflated, reticulate, similar at apex; not concealing radicle; entire over and pubescent and lower 1/4 not inflated, reticulate, or radicle; without lobes; with the interface division pubescent; coriaceous; seed chambers externally visible terminating at base of radicle; without margins re- or invisible; with the raised seed chambers not torulose. cessed; yellow; inner face flat; glabrous around base of Fruit margin constricted or not constricted; constricted radicle. Embryonic axis deflexed; parallel to length of along both margins; without sulcus; plain. Fruit wings seed. Radicle linear; deflexed and parallel to cotyledon absent. Fruit nonstipitate. Fruit with all layers dehisc- length; centered between cotyledons; less than 1/2 or ing. Replum invisible. Loment indehiscent. Loment 1/2 to nearly length of cotyledons. Plumule rudimen- segments (articles) inconspicuous; 4.5Ð5.5 mm long; tary; glabrous. widest across each end or widest across seed area; with upper or lower 1 different shape than middle ones; Distribution: Europe, Mediterranean region, Atlantic oblong or quadrangular. Epicarp dull or glaucous; islands, West Asia, northeastern Africa. monochrome; brown or tan; glabrous; glandular or eglandular; with glandular dots; limited to a portion of Notes: Schmidt (1979b) monographed Coronilla and fruit; upper 3/4 glandular and lower 1/4 eglandular; documented (Schmidt 1979a) the reduction of without spines; not smooth; with elevated features; Securigera. We accept Securigera (13.12) as a separate longitudinally veined relative to fruit length; not genus based on Lassen (1989) and use it as the basis of tuberculate; not exfoliating; without cracks. Mesocarp our species count. Schmidt (1979a) illustrated the seeds absent. Endocarp glossy; monochrome or bichrome of 31 species, showing which ones bear a longitudinal (with central reddish-brown band); tan; smooth; line on each face. nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð7; length parallel with fruit length; neither overlapping nor touching; in 2 or more series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 3Ð5 ϫ 1Ð2 ϫ 1Ð1.5 mm; not overgrown; not angular; symmetrical or asymmetrical; rhombic; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; Coronilla: C. scorpioides (C. Linnaeus) W.D.J. Koch dark reddish brown or tan; glabrous; smooth (except for (CÐE), C. spp. (AÐB). A, Fruits and articles (ϫ 2.3); B, 1 longitudinal ridge on each face) or not smooth; with seeds (ϫ 6.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

739 Genus: Securigera A.P. de Candolle cal; linear, rectangular, or quadrangular; compressed or flattened; without visible radicle and cotyledon lobes; Phylogenetic Number: 13.12. without umbo on seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; Tribe: Loteae. colored; monochrome; light to dark reddish brown or tan (reddish near hilum); glabrous; smooth; coriaceous. Species Studied—Species in Genus: 5 spp.—12 spp. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum fully concealed; concealed by Fruit a legume or loment (or a loment segment); unilocular; funicular remnant; without faboid split; punctiform; 8.5Ð9.5 ϫ 0.15Ð0.6 ϫ 0.1Ð1.6 cm; with deciduous marginal according to radicle tip; flush; not within corolla; with deciduous calyx; without orifice formed corona, halo, or rim or within halo. Hilum halo color by curving of fruit or fruit segments; straight to curved darker than testa. Lens discernible or not discernible; (slightly); not plicate; not twisted; symmetrical; linear; less than 0.5 mm in length; with margins curved; not inflated; flattened or terete; with beak; declined or circular; not in groove of raphe; adjacent to hilum; 0.3 hooked; with solid beak the same color and texture as mm from hilum; mounded; same color as testa; within fruit; long tapered at apex; apex aligned with longitudi- halo. Lens halo color darker than testa. Endosperm nal axis of fruit; short tapered at base; base aligned with thick; covering entire embryo; adnate to embryo. longitudinal axis of fruit; with the apex and base Cotyledons smooth; both outer faces convex; both the uniform in texture; coriaceous; seed chambers exter- same thickness; both more or less of equal length; not nally visible; with the raised seed chambers not folded; margin entire 180 degrees from base of radicle; torulose. Fruit margin not constricted; with sulcus; plain similar at apex; not concealing radicle; entire over or embellished; with thickened sutural areas. Fruit radicle; without lobes; with the interface division wings absent. Fruit substipitate or nonstipitate. Fruit terminating at base of radicle; without margins re- indehiscent (the chartaceous valve surface between cessed; yellow or tan; inner face flat; glabrous around thickened sutures open transversely and irregularly in S. base of radicle. Embryonic axis deflexed; oblique to securidaca). Replum invisible. Loment indehiscent. length of seed. Radicle bulbose or linear; deflexed and Loment segments (articles) inconspicuous; 5Ð10 mm parallel to cotyledon length; centered between cotyle- long; widest across seed area; with upper 1 different dons; less than 1/2 or 1/2 to nearly length of cotyledons. shape than middle ones; linear. Epicarp dull; mono- Plumule rudimentary; glabrous. chrome or multicolored; bichrome (sutures thickened and a lighter or darker shade); brown (including dirty- Distribution: Southern Europe. brown), tan (and greenish-tan), or green; glabrous or pubescent and indurate; with 1 type of pubescence; Notes: Securigera has been maintained separate from pilose (but not on thickened sutures); with pubescence Coronilla (13.11) because of its fruits being heavy, golden; with pubescence uniformly distributed; with flattened, and apparently not divided and thus not simple hairs; pliable; with hair bases plain; eglandular; breaking into articles at maturity. It was united with without spines; not smooth; with elevated or recessed Coronilla (13.11) for diverse reasons by Schmidt features; veined or not veined; longitudinally veined (1979a). Lassen (1989) realigned Coronilla (13.11), relative to fruit length (1 medial on each valve) or Hippocrepis (13.13), and Securigera and in so doing reticulately veined (faint); not tuberculate; wrinkled recognized 12 species of Securigera, the species count (faintly); grooved (faintly); not exfoliating; without that we are using. When doing this, Lassen devaluated cracks. Mesocarp absent. Endocarp dull; monochrome; the loment versus legume character. tan; smooth; septate or subseptate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 7Ð9; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent. Securigera: S. securidaca (C. Linnaeus) A. von Degen & I. Doerfler (CÐE), S. spp. (AÐB). A, Fruits and articles Seed 3.5Ð4 ϫ 1.5Ð4 ϫ 1Ð1.5 mm; not overgrown; angular (ϫ 1.2); B, seeds (ϫ 6.5); CÐD, testa (ϫ 50, ϫ 1000); or not angular (some are square in outline); symmetri- E, embryos (ϫ 11.4).

741 Genus: Hippocrepis C. Linnaeus without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; Phylogenetic Number: 13.13. glossy; not modified by a bloom; colored; monochrome; brown (brown to reddish), green (yellowish), Tribe: Loteae. tan, or blue (reddish); glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe Species Studied—Species in Genus: 10 spp.—30 spp. not visible. Hilum visible; without faboid split; punctiform; between cotyledon and radicle lobe; slightly Fruit a loment (or a loment segment); 1.2Ð4 ϫ 0.1Ð0.6 ϫ recessed; within corona (H. ciliata C.L. von 0.07Ð0.16 cm; with deciduous corolla; with persistent Willdenow) or not within corona, halo, or rim. Hilum calyx; with calyx shorter than fruit; with orifice formed corona color darker than testa (black). Lens discernible; by curving of fruit or fruit segments (more than 1 per less than 0.5 mm in length; with margins curved; fruit) or without orifice formed by curving of fruit or circular; not in groove of raphe; adjacent to hilum; fruit segments (H. emerus); straight to curved (to mounded or flush; same color as testa or dissimilar slightly curved) or 0.5-coiled to 1-coiled to 1.5-coiled; color from testa; darker than testa; red; not within not plicate; not twisted; asymmetrical or symmetrical; corona, halo, or rim. Endosperm thin; covering entire linear or quadrangular; when asymmetrical with both embryo; adnate to embryo. Cotyledons smooth; both sutures parallelly or unequally curved; not inflated; outer faces convex; both the same thickness; both more flattened or compressed; without beak; long tapered to or less of equal length; not folded; margin entire 180 tapered at apex; apex aligned to oblique with longitudi- degrees from base of radicle; similar at apex; not nal axis of fruit; long tapered at base; base aligned to concealing radicle; entire over radicle; without lobes; oblique with longitudinal axis of fruit; with the apex with the interface division terminating at base of and base uniform in texture; coriaceous; seed chambers radicle; without margins recessed; yellow; inner face externally visible; with the raised seed chambers not flat; glabrous around base of radicle. Embryonic axis torulose. Fruit margin constricted or not constricted; deflexed; parallel to length of seed. Radicle linear; with constricted along both margins; without sulcus; plain or 180-degree turn; not centered between cotyledons embellished; with wing. Fruit wing present or absent; 1; (radicle outside 1 cotyledon and inside other, therefore 2 mm wide; sutural; on both sutures. Fruit substipitate. junctions for each cotyledon different); less than 1/2 Fruit indehiscent. Replum invisible. Loment indehis- length of cotyledons. Plumule moderately developed; cent. Loment segments (articles) inconspicuous; 5Ð25 glabrous. mm long; widest across seed area; with all essentially similar in shape; hippocrepiform. Epicarp semiglossy; Distribution: Mediterranean region into Europe and western monochrome; reddish brown; pubescent and indurate; Asia. with 1 type of pubescence; puberulent; with pubescence gray; with pubescence uniformly distributed; with Notes: Domínguez (1976) monographed the annual species glandular hairs (restricted to top of arch between of Hippocrepis and illustrated their fruits. Our species winged margins); stiff; with hair bases plain; glandular; count is based on Lassen (1989), who gave the genus a with glandular hairs; without spines; smooth or not traditional treatment except for the transfer of Coronilla smooth; with elevated features; longitudinally veined emerus C. Linnaeus into Hippocrepis. This transfer is relative to fruit length; not tuberculate; not exfoliating; particularly disturbing to us because the fruit characters without cracks. Mesocarp thin; surface not veined; 1- and to some degree the seed characters are unlike those layered; without balsamic vesicles; without fibers; of the other species in Hippocrepis (compare figs. A solid; coriaceous. Endocarp glossy; monochrome; tan; and B). smooth; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 3Ð12; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent. Hippocrepis: H. emerus (C. Linnaeus) P. Lassen (B), H. unisiliquosa C. Linnaeus (DÐF), H. spp. (A, C). AÐB, Seed 2Ð4.5 ϫ 0.7Ð0.8 ϫ 0.7Ð0.8 mm; not overgrown; not Fruits and fruit segments (ϫ 2, ϫ 2.3); C, seeds (ϫ 6); angular; asymmetrical; hippocrepiform or linear; terete; DÐE, testa (ϫ 50, ϫ 1000); F, embryos (ϫ 10).

743 Genus: Scorpiurus C. Linnaeus brous; smooth or not smooth; with elevated and recessed (present or absent) features; shagreen; striate Phylogenetic Number: 13.14. (faintly and when present caused by ribs on fruit); coriaceous. Fracture lines absent. Rim absent. Raphe Tribe: Loteae. not visible. Hilum fully concealed; concealed by funicular remnant; without faboid split; punctiform; Species Studied—Species in Genus: 2 spp.—2 spp. marginal according to radicle tip (but on side opposite lobes); flush; not within corona, halo, or rim or within Fruit a loment (or a loment segment); 8 ϫ 0.3 ϫ 0.3 cm; halo. Hilum halo color darker than testa (blackish- with deciduous corolla; with deciduous calyx; with or brown). Lens discernible; less than 0.5 mm in length; without orifice formed by curving of fruit or fruit with margins curved; elliptic; not in groove of raphe; segments; 4-coiled; not plicate to plicate (at most adjacent to hilum; 0.4 mm from hilum; mounded; loosely plicate); loosely to tightly twisted or not dissimilar color from testa; darker than testa; dark twisted; asymmetrical or symmetrical; linear; when brown; not within corona, halo, or rim. Endosperm asymmetrical with both sutures nearly straight; not thick; covering entire embryo; adnate to embryo. inflated; terete; without beak; long tapered or short Cotyledons smooth or not smooth; both outer faces tapered at apex; apex exceeding (crossing) longitudinal convex; both the same thickness; both more or less of axis of fruit; long to short tapered at base; base aligned equal length; with both folded; not sufficiently folded with longitudinal axis of fruit; with the apex and base for inner face to touch itself; portions of inner folded uniform in texture; coriaceous; seed chambers exter- face unequal; margin entire 180 degrees from base of nally invisible. Fruit margin constricted or not con- radicle; similar at apex; not concealing radicle; entire stricted; slightly constricted along both margins; over radicle; without lobes; with the interface division without sulcus; plain. Fruit wings absent. Fruit terminating at base of radicle; without margins re- nonstipitate. Fruit indehiscent. Replum invisible. cessed; white; inner face flat; glabrous around base of Loment indehiscent. Loment segments (articles) radicle. Embryonic axis deflexed; oblique to length of inconspicuous; 5Ð10 mm long; with all essentially seed. Radicle bulbose; deflexed and parallel to cotyle- similar in shape; curved. Epicarp dull; monochrome or don length; centered between cotyledons; less than 1/2 multicolored; bichrome (spines may be darker colored length of cotyledons. Plumule rudimentary; glabrous. than surface and ribs below spines may be darker colored); brown or tan; glabrous; eglandular; with Distribution: Europe, Mediterranean region, western Asia, spines (aligned in longitudinal ribs) or without spines northeastern Africa. (longitudinal ribs present); with spines persistent; with spines same color or spines (or their basal remanent) a Notes: Domínguez and Galiano (1974) monographed different color from the rest of the fruit; not smooth; the genus and recognized four species; however, Heyn with elevated features; not veined; tuberculate or not and Raviv (1966) recognized only two of the four tuberculate; tuberculate (blunt spines in longitudinal species: S. vermiculatus C. Linnaeus and S. muricatus. rows); not exfoliating; without cracks. Mesocarp Heyn and Raviv reorganized the S. muricatus complex absent. Endocarp dull; monochrome; tan; smooth; with two varieties: var. muricatus (S. muricatus, s.s., subseptate; with septa thicker than paper, firm; with the and S. sulcatus, p.p.) and var. subvillosus (C. Linnaeus) septa eglandular; chartaceous; not exfoliating; remain- J.B.A.P.M. de Lamarck (S. subvillosus and S. sulcatus ing fused to epicarp; entire. Seed length parallel with C. Linnaeus, p.p.). Kaniewski and Miszkiel (1980) fruit length; neither overlapping nor touching; in 1 studied the histogenesis of the pericarp of Scorpiurus series. Funiculus less than 0.5 mm long; of 1 length and noted that S. muricatus var. muricatus had “pods only; filiform; straight. Aril absent. with smooth or tubercled ribs, usually coiled in one plane,” while S. muricatus var. subvillosus had “pods Seed 2.3Ð5 ϫ 1.5Ð3 ϫ 1.5Ð3 mm; not overgrown; not spiny, usually twisted.” Heyn and Raviv noted that the angular; asymmetrical; C-shaped, circular (subcircular), two seed shapes (straight with truncate margin and or reniform; terete; without visible radicle and cotyle- reniform with apiculate margin) faithfully reflect the don lobes; without umbo on seed faces. Testa not degree of coiling of the loment segment, or article in adhering to endocarp; dull; not modified by a bloom; which it developed. Upon soaking the seeds, the colored; monochrome; orangish red to orange; gla- endosperm greatly expands (up to twice the diameter of the seed) and ruptures the testa, as it does in Antopetita (13.16). 744 745 Scorpiurus: S. muricatus C. Linnaeus (CÐE), S. spp. (AÐB). A, Fruits and one article (ϫ 2.2); B, seeds (ϫ 4.8); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

747 Genus: Ornithopus C. Linnaeus bloom; colored; monochrome; orange (reddish), red (dish), or yellow (pale); glabrous; smooth; coriaceous. Phylogenetic Number: 13.15. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the Tribe: Loteae. lips of the faboid split the same color as the rest of the hilum; punctiform; marginal according to radicle tip; Species Studied—Species in Genus: 6 spp.—6 spp. flush; within halo. Hilum halo color darker than testa. Lens discernible or not discernible; less than 0.5 mm in Fruit a loment (or a loment segment); 1.5Ð5 ϫ 0.1Ð0.23 ϫ length; with margins curved; circular (sub); not in 0.08Ð0.13 cm; with deciduous corolla; with persistent groove of raphe; adjacent to hilum; mounded; dissimi- or deciduous calyx; with calyx shorter than fruit; lar color from testa; darker than testa; black; not within without orifice formed by curving of fruit or fruit corona, halo, or rim. Endosperm thick or thin; covering segments; curved to 0.5-coiled; not plicate; not twisted; entire embryo; adnate to testa. Cotyledons smooth; both asymmetrical or symmetrical; linear or falcate; when outer faces convex; both the same thickness; both more asymmetrical with both sutures parallelly curved; not or less of equal length; not folded; margin entire 180 inflated; compressed; with beak; hooked; with solid degrees from base of radicle; similar at apex; not beak the same color and texture as fruit; long tapered at concealing radicle; entire over radicle; without lobes; apex; apex oblique with longitudinal axis of fruit; with the interface division terminating at base of tapered at base; base aligned with longitudinal axis of radicle; without margins recessed; yellow; inner face fruit; with the apex and base uniform in texture; flat; glabrous around base of radicle. Embryonic axis coriaceous; seed chambers externally visible; with the deflexed; parallel to length of seed. Radicle bulbose; raised seed chambers not torulose. Fruit margin deflexed and parallel to cotyledon length; centered constricted or not constricted; slightly constricted along between cotyledons; less than 1/2 or 1/2 to nearly both margins; without sulcus; plain. Fruit wings absent. length of cotyledons. Plumule rudimentary or moder- Fruit nonstipitate. Replum invisible. Loment indehis- ately developed; glabrous. cent. Loment segments (articles) inconspicuous or conspicuous; 4Ð11 mm long; widest across seed area; Distribution: Five species in Europe, Mediterranean region, with upper or lower 1 different shape than middle ones; western Asia, and Atlantic islands; O. micranthus (G. circular, oblong, or rectangular. Epicarp dull; mono- Bentham) J. Arechaveleta y Bolpardo in southern South chrome; brown or tan; glabrous or pubescent and America. indurate; with 1 type of pubescence; puberulent; with pubescence brown; with pubescence uniformly distrib- Notes: Alsina (1988) described the morphology and uted; with simple hairs; pliable; with hair bases plain; anatomy of seeds of four species and illustrated their eglandular; without spines; smooth or not smooth; with testa with micrographs at ϫ320. The monotypic genus elevated features; reticulately veined; not tuberculate; Antopetitia (13.16) differs by having dehiscent fruit not exfoliating; without cracks. Mesocarp thin; surface segments and a persistent dorsal suture (Verdcourt not veined; 1-layered; without balsamic vesicles; 1974). without fibers; solid; coriaceous. Endocarp glossy; monochrome; tan; smooth; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 4Ð12; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 1.5Ð3 ϫ 1Ð2 ϫ 0.7Ð1.1 mm; not overgrown; not angular (except hilum); symmetrical (except hilum); circular, oblong, ovate, or reniform; compressed; without visible radicle and cotyledon lobes; without Ornithopus: O. sativus F.d’A. Brotero (CÐE), O. spp. (AÐB). umbo on seed faces. Testa dull; not modified by a A, Fruits and articles (ϫ 2.2); B, seeds (ϫ 5.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

749 Genus: Antopetitia A. Richard visible; without faboid split (hilum too small); punctiform; between cotyledon and radicle lobe; flush; not Phylogenetic Number: 13.16. within corona, halo, or rim. Lens not discernible. Endosperm thick; covering entire embryo; adnate to Tribe: Loteae. testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; Species Studied—Species in Genus: 1 sp.—1 sp. not folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing radicle; entire Fruit a legume or a loment (not a true loment); unilocular; over radicle; without lobes; with the interface division 0.7Ð1.2 ϫ 0.2Ð0.25 ϫ 0.23 cm; with deciduous corolla; terminating at base of radicle; without margins re- with persistent calyx; with calyx shorter than fruit; cessed; yellow; inner face flat; glabrous around base of without orifice formed by curving of fruit or fruit radicle. Embryonic axis deflexed; oblique to length of segments; slightly curved to curved; not plicate; not seed. Radicle bulbose; oblique to cotyledons; centered twisted; asymmetrical; falcate or moniliform; when between cotyledons; 1/2 to nearly length of cotyledons. asymmetrical with both sutures parallelly curved; not Plumule rudimentary; glabrous. inflated; terete; with beak (small); straight; with solid beak the same color and texture as fruit; rounded at Distribution: Tropical Africa. apex; apex aligned with longitudinal axis of fruit; short tapered at base; base oblique with longitudinal axis of Notes: Polhill (1981l) noted that this genus is closely related fruit; with the apex and base uniform in texture; to Ornithopus (13.15) and that the genera of the coriaceous; seed chambers externally visible; with the Coronilleae only differ from those of the Loteae (13) by raised seed chambers not torulose. Fruit margin the presence of jointed fruits in the former. Verdcourt constricted along both margins; without sulcus; plain. (1974) noted that Antopetitia is “a monotypic genus Fruit wings absent. Fruit substipitate (may be longer closely allied to Ornithopus L., but differing in the pod than 2Ð3 mm if lowest locule aborted). Fruit with all segments being dehiscent and in the persistent dorsal layers dehiscing; fenestrating, opening by a coiling strip suture.” Both authors alluded to an anomaly in fruit of of tissue. Dehiscence of valves along 1 suture; passive. Antopetita. These fruits are unique in the Fabaceae. At Replum invisible. Loment dehiscing along 1 suture. first glance they appear to be loments, but they are not. Loment segments (articles) inconspicuous; 2Ð2.5 mm At most they are loment-like or lomentaceous because long; widest across seed area; with upper or lower 1 the fruits dehisce, albeit by a unique mechanism. In different shape than middle ones; circular. Epicarp addition, Léonard (1954, fig. 10) clearly illustrated that glossy; monochrome; light brown to tan; glabrous; the fruit dehisces on the plant, the seeds falling free of eglandular; without spines; not smooth; with elevated the fruiting cluster. By definition a loment separates at features; reticulately veined; not tuberculate; not maturity into 1-seeded indehiscent articles. The unique exfoliating; without cracks. Mesocarp thin; surface not dehiscence mechanism is the opening of the valves by veined; 1-layered; without fibers; solid; coriaceous. the rolling up of the vertical suture. During the process, Endocarp glossy; monochrome; tan or white; smooth; the individual fruit segments become fenestrated and nonseptate; chartaceous; not exfoliating; remaining then the seed may fall out or one valve may fall off fused to epicarp; entire. Seeds 2Ð4; length parallel with (Léonard 1954, fig. 10). Upon soaking the testa fruit length; neither overlapping nor touching; in 2 or fractures as the endosperm rapidly expands (as in more series. Funiculus less than 0.5 mm long; of 1 Scorpiurus (13.14)). Both the seed and fruit dehis- length only; thick; straight. Aril absent. cences are related to edaphic factors in the habitat of the species. Seed 1.5Ð2 ϫ 1.2Ð1.7 ϫ 0.5Ð1.3 mm; not overgrown; angular to not angular; asymmetrical; circular to irregular; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; dark to reddish brown to tan (reddish); glabrous; not smooth; with elevated features; Antopetitia: A. abyssinica A. Richard (AÐE). A, Fruits and rugose or tuberculate; coriaceous. Fracture lines absent. fruit segments (ϫ 4.5); B, seeds (ϫ 6.3); CÐD, testa Rim absent. Wings absent. Raphe not visible. Hilum (ϫ 50, ϫ 1000); E, embryos (ϫ 20).

751 Genus: Hammatolobium E. Fenzl and radicle lobe; recessed; not within corona, halo, or rim. Lens discernible; less than 0.5 mm in length; with Phylogenetic Number: 13.17. margins curved; elliptic; not in groove of raphe; adjacent to hilum; mounded; dissimilar color from Tribe: Loteae. testa; darker than testa; black; not within corona, halo, or rim. Endosperm thin; covering entire embryo; adnate Species Studied—Species in Genus: 2 spp.—2 spp. to testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal Fruit a loment (or a loment segment); 2Ð6 ϫ 0.15Ð0.4 ϫ 0.2 length; not folded; margin entire 180 degrees from base cm; with deciduous corolla; with persistent calyx; with of radicle; similar at apex; not concealing radicle; entire calyx shorter than fruit; without orifice formed by over radicle; without lobes; with the interface division curving of fruit or fruit segments; straight; not plicate; terminating at base of radicle; without margins re- not twisted; symmetrical; linear and moniliform; not cessed; yellow; inner face flat; glabrous around base of inflated; compressed; with beak; straight; with solid radicle. Embryonic axis deflexed; oblique to length of beak the same color and texture as fruit; short tapered at seed. Radicle linear; oblique to cotyledons; centered apex; apex oblique with longitudinal axis of fruit; short between cotyledons; 1/2 to nearly length of cotyledons. tapered at base; base aligned with longitudinal axis of Plumule rudimentary; glabrous. fruit; with the apex and base uniform in texture; coriaceous; seed chambers externally visible; with the Distribution: One species at either end of the Mediterranean raised seed chambers not torulose. Fruit margin region. constricted along both margins; without sulcus; plain. Fruit wings absent. Fruit substipitate. Fruit indehiscent. Notes: Polhill (1981l) noted that this genus seems to be Replum invisible. Loment indehiscent. Loment seg- more closely related to Anthyllis (13.02) and Lotus ments (articles) inconspicuous; 3Ð6 mm long; widest (13.07) in the Loteae than to Coronilla (13.11). across seed area; with upper or lower 1 different shape than middle ones; oblong. Epicarp dull; monochrome; dark brown; pubescent and indurate; with 1 type of pubescence; pilose or villous; with pubescence golden; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; smooth or not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1- layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; white; smooth; septate; with septa thicker than paper, firm; with the septa eglandular; coriaceous; not exfoliating; remaining fused to epicarp; entire. Seeds up to 11; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 2.5 ϫ 2.3 ϫ 1 mm; not overgrown; not angular; asymmetrical; nearly circular; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hammatolobium: H. kremerianum (E.St.-C. Cosson) C.H. Hilum fully concealed; concealed by funicular remnant; Muller (CÐE), H. spp. (AÐB). A, Fruits with calyx and 1 without faboid split; punctiform; between cotyledon article (ϫ 1.8); B, seeds (ϫ 7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

753 Aeschynomeneae (14.01Ð14.26) chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð6; length parallel with fruit Genus: Ormocarpum A.M.F.J. Palisot de Beauvois length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; Phylogenetic Number: 14.01. thick; straight or curved. Aril absent.

Tribe: Aeschynomeneae. Seed 2Ð9 ϫ 1.5Ð4 ϫ 1.2Ð2 mm; not overgrown; not angular; asymmetrical; circular to elliptic to oblong; Subtribe: Ormocarpinae. compressed to flattened; with visible radicle and cotyledon lobes; with external groove between radicle Species Studied—Species in Genus: 12 spp.—ca. 20 spp. and cotyledon lobes; same color as testa; without umbo on seed faces. Testa not adhering to endocarp; dull; Fruit a legume or loment; unilocular; 3.5Ð6 ϫ 0.5Ð0.8 ϫ colored; monochrome; pinkish to reddish brown or tan; 0.2Ð0.3 cm; with persistent (O. kirkii S. le M. Moore) or glabrous; smooth; chartaceous. Fracture lines absent. deciduous corolla; with persistent (O. kirkii) or decidu- Rim absent. Wings absent. Raphe from hilum to near ous calyx; with calyx shorter than fruit; with or without base of seed and terminating (from hilum to nearly base orifice formed by curving of fruit or fruit segments (O. of seed); not bifurcating; darker than testa; brown; kirkii); straight to curved to 1-coiled (O. kirkii); not flush. Hilum fully concealed; concealed by funicular plicate; not twisted; asymmetrical or symmetrical; remnant; without faboid split; larger than punctiform; moniliform, C-shaped, or coiled; when asymmetrical 1Ð1.3 mm long; with curved outline; elliptic; between with 1 straight and 1 curved suture or both sutures cotyledon and radicle lobe; flush; within halo. Hilum parallelly curved; widest near middle or D-shaped; not halo color of or darker than testa. Lens not discernible. inflated; flattened; without or with beak; straight; with Endosperm trace;1/2 covering entire embryo; adnate to solid beak the same color and texture as fruit; long testa. Cotyledons smooth; both outer faces convex; both tapered, short tapered, or rounded at apex; apex aligned the same thickness; both more or less of equal length; to oblique with longitudinal axis of fruit; long or short not folded; margin entire 180 degrees from base of tapered at base; base aligned to oblique with longitudi- radicle; similar at apex; partially concealing radicle; nal axis of fruit; with the apex and base uniform in split over radicle; with lobes; with lobes not touching; texture; coriaceous; seed chambers externally visible; with the interface division terminating at base of with the raised seed chambers not torulose. Fruit margin radicle; without margins recessed; reddish tan; inner constricted along both margins; without sulcus; plain. face flat; glabrous around base of radicle. Embryonic Fruit wings absent. Fruit stipitate to substipitate; with axis deflexed; oblique to length of seed. Radicle linear; the stipe 3Ð8 mm long. Fruit indehiscent. Replum oblique to cotyledons; centered between cotyledons; invisible. Loment indehiscent. Loment segments less than 1/2 length of cotyledons. Plumule well (articles) inconspicuous; 6Ð20 mm long; widest across developed; glabrous. seed area; with upper 1 or lower 1 different shape than middle ones; oblong or D-shaped. Epicarp dull; Distribution: Tropical and South Africa, Madagascar, monochrome; dark green; glabrous or pubescent and southern Asia to Philippines, and Fiji. indurate; with 1 type of pubescence; puberulent; with pubescence yellow; with pubescence uniformly distrib- Notes: Rudd (1981a) recognized four subtribes of uted; with simple hairs (often with swollen bases); stiff; Aeschynomeneae: Ormocarpinae Rudd (genera 14.01Ð with hair bases swollen; retrorse; straight at apex; 14.08), Aeschynomeninae (genera 14.09Ð14.16), glandular; with glandular hairs (minutely tubercule Discolobiinae (Burkart) Rudd (genus 14.17: bases); with spines; not smooth; with elevated features; Discolobium), Poiretiinae (Burkart) Rudd (genera longitudinally veined relative to fruit length or reticu- 14.18Ð14.21), and Stylosanthinae (Bentham) Rudd lately veined; tuberculate or not tuberculate; with solid (genera 14.22Ð14.26). Tribal and subtribal placement of tubercles on each valve; tuberculate; not exfoliating; Diphysa is based on Lavin (1987) and Polhill (1994a, without cracks. Mesocarp thick; surface not veined; 1- 1994b) and not on Polhill and Sousa (1981), who layered; without balsamic vesicles; without fibers; placed Diphysa in Robinieae. Because Lavin did not solid; coriaceous. Endocarp nearly glossy; mono- relate Diphysa to a genus in subtribe Ormocarpinae, we chrome; brown; smooth; septate or nonseptate; with have assigned the genus the phylogenetic number septa thicker than paper, firm; with the septa eglandular; 14.03. Bailey et al. (1997), using the chloroplast rpl2

754 intron and ORF184, suggested that Brya (11.01), Cranocarpus (11.02), Phylacium (11.22), and Neocollettia (11.26) are not members of Desmodieae (11) and that they probably belong in Aeschynomeneae. Gillett (1966b) described the 17 species of Ormocarpum from southwestern Asia and Africa (excluding Madagascar) and presented some fruit and seed data and some fruit illustrations.

755 Ormocarpum: O. orientale (C.P.J. Sprengel) E.D. Merrill (CÐE), O. spp. (AÐB). A, Article and partial fruit, without or with calyx (ϫ 1.8); B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

757 Genus: Ormocarpopsis R. Viguier wrinkled and 1Ð3 grooves on each face; both outer faces convex; both the same thickness; both more or Phylogenetic Number: 14.02. less of equal length; not folded; margin not entire 180 degrees from base of radicle; notched or notched and Tribe: Aeschynomeneae. bearing flaps; similar at apex; completely concealing radicle; split over radicle; with lobes; with lobes Subtribe: Ormocarpinae. touching (auriculate) or not touching; with basal groin formed by lobes; with the interface division terminating Species Studied—Species in Genus: 1 sp.—5 spp. at base of radicle; without margins recessed; brown; inner face concave (in center with varying width rim Fruit a legume; unilocular; 2.3Ð3 ϫ 1.3Ð2 ϫ 1.3Ð2 cm; with around the depression); glabrous around base of radicle. deciduous corolla; with deciduous calyx; without Embryonic axis straight; parallel to oblique to length of orifice formed by curving of fruit or fruit segments; seed; with a joint evident between the radicle and the straight; not plicate; not twisted; asymmetrical; oblong; cotyledons. Radicle differentiated from cotyledon; when asymmetrical with both sutures parallelly curved; triangular; lobe tip straight; straight with embryonic not inflated; terete; with beak; straight; with the apex axis; centered between cotyledons; less than 1/2 length and base uniform in texture; seed chambers externally of cotyledons. Plumule well developed; pubescent at invisible. Fruit margin not constricted; without sulcus; the base and glabrous at the apex. embellished. Fruit wings absent. Fruit stipitate or substipitate; with the stipe up to 8 mm long. Fruit Distribution: Madagascar. indehiscent. Replum invisible. Fruit a nutlet; entire. Epicarp dull; monochrome; greenish brown; glabrous; glandular; with spines (glandular) or without spines; not smooth; with elevated features; not veined; tuberculate or not tuberculate; with solid tubercles on each valve (minute); warty (bases of spines); not exfoliating; without cracks. Mesocarp present (and streaked with purple); thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; exfoliating; separating from mesocarp. Seed 1; length parallel with fruit length. Funiculus less than 0.5 mm long; thick; straight. Aril absent.

Seed 18 ϫ 13 ϫ 10 mm; overgrown, 1 seed filling entire fruit cavity; not angular; asymmetrical; ovate; terete; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; not smooth; with elevated features; shagreen; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; without faboid split; larger than punctiform; 1.5 mm long; with straight outline; linear; marginal according to radicle tip; flush; not within corona, halo, or rim. Lens discernible; equal to or greater than 0.5 mm in length; 2.5 mm long; with margins straight or curved; oblong; not in groove of raphe; confluent with hilum; mounded; similar color as testa; darker than testa; reddish brown; Ormocarpopsis: O. aspera R. Viguier (AÐE). A, Fruit not within corona, halo, or rim. Endosperm absent. (ϫ 2.1); B, seed (ϫ 2.3); CÐD, testa (ϫ 50, ϫ 1000); Cotyledons not smooth or smooth; wrinkled or E, embryos (ϫ 2.5).

759 Genus: Diphysa N. von Jacquin hilum; punctiform to larger than punctiform; up to 0.5 mm long; with curved outline; circular; subapical to Phylogenetic Number: 14.03. radicle tip or between cotyledon and radicle lobe; recessed; within rim or not within corona, halo, or rim. Tribe: Aeschynomeneae. Hilum rim color of testa. Lens not discernible. En- dosperm thick to thin; covering entire embryo; adnate Subtribe: Ormocarpinae. to testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal Species Studied—Species in Genus: 8 spp.—15 spp. or 20 length; not folded; margin entire 180 degrees from base spp. of radicle; similar at apex; partially concealing radicle; split over radicle; with lobes; with lobes not touching; Fruit a legume; unilocular; 3Ð6.5 ϫ 0.7Ð3 ϫ 0.7Ð3 (esti- with the interface division terminating at base of mated) cm; with deciduous corolla; with persistent radicle; without margins recessed; yellow or green calyx; with calyx shorter than fruit; without orifice (ish); inner face flat; glabrous around base of radicle. formed by curving of fruit or fruit segments; straight; Embryonic axis right angled; perpendicular to length of not plicate; not twisted; symmetrical; linear or oblong; seed. Radicle linear; lobe tip straight; deflexed and inflated; terete; without beak; short tapered to rounded parallel to cotyledon width; centered between cotyle- at apex; apex aligned with longitudinal axis of fruit; dons; less than 1/2 length of cotyledons. Plumule long tapered or rounded at base; base aligned with moderately developed to well developed; glabrous. longitudinal axis of fruit; with the apex and base uniform in texture; membranous; seed chambers Distribution: Mexico, Central America, northern South externally invisible. Fruit margin not constricted; America. without sulcus; plain or embellished; with wing. Fruit wing present or absent; 1; sutural (subwinged or Notes: Tribal and subtribal placement of Diphysa is based keeled); on 1 suture. Fruit stipitate to substipitate; with on Lavin (1987, Polhill 1994a,b), and not on Polhill the stipe up to 15 mm long. Fruit indehiscent. Replum and Sousa (1981), who placed Diphysa in Robinieae. invisible. Epicarp dull; monochrome; brown or tan; glabrous; without spines; not smooth; with elevated features; longitudinally veined relative to fruit length (with faint cross veining) or reticulately veined; not tuberculate; not exfoliating; with to without cracks; cracking irregular. Mesocarp absent. Endocarp dull; monochrome; tan; scurfy; nonseptate; coriaceous; not exfoliating; remaining fused to epicarp; entire. Seeds few, probably less than 5; length oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 4Ð7 ϫ 2.7Ð4.5 ϫ 1.5Ð3.3 mm; not overgrown; not angular; asymmetrical; oblong; terete to compressed; with or without visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; same color as testa; without umbo on seed faces. Testa not adhering to endocarp; dull; modified by a bloom; colored; monochrome; dark reddish brown to tan; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe from hilum to near base of seed and terminating; not bifurcating; color of testa; raised to Diphysa: D. minutifolia J.N. Rose (E), D. robinioides G. recessed. Hilum visible to partially concealed; con- Bentham (CÐD), D. spp. (AÐB). A, Fruits (ϫ 0.9); B, cealed by funicular remnant; with faboid split; with the seeds (ϫ 2.4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos lips of the faboid split the same color as the rest of the (ϫ 6).

761 Genus: Belairia A. Richard ing at base of radicle; without margins recessed; tan; inner face flat; glabrous around base of radicle. Embry- Phylogenetic Number: 14.04. onic axis deflexed; oblique to length of seed. Radicle bulbose; lobe tip straight; deflexed and parallel to Tribe: Aeschynomeneae. cotyledon length or width; centered between cotyledons; less than 1/2 length of cotyledons. Plumule well Subtribe: Ormocarpinae. developed; glabrous.

Fruit a legume; unilocular; 1.5Ð2 ϫ 0.4Ð0.5 ϫ 0.03 cm; Notes: Data from the meager (at the U.S. National Seed with deciduous corolla; with deciduous calyx; without Herbarium [BARC]) samples were supplemented by orifice formed by curving of fruit or fruit segments; fruit notes and illustrations in Lewis (1988). The straight; not plicate; not twisted; symmetrical; linear or number of species in the genus is based on Lewis, not oblong; not inflated; flattened; without or with beak; on Rudd (1981a). with solid beak the same color and texture as fruit; short tapered to rounded at apex; apex aligned with longitudinal axis of fruit; short tapered at base; base aligned with longitudinal axis of fruit; with the apex and base uniform in texture; coriaceous; seed chambers externally invisible. Fruit margin not constricted; without sulcus; plain. Fruit wings absent. Fruit substipitate. Fruit indehiscent. Replum invisible. Epicarp dull; monochrome; brown; glabrous; eglandular; without spines; not smooth; with elevated features; longitudinally veined relative to fruit length or reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; spongy; coriaceous. Endocarp dull; monochrome; brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 3.5Ð3.7 ϫ 2.5Ð2.7 ϫ 0.6 mm; not overgrown; not angular; asymmetrical; circular (with radicle lobe); flattened; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; tan; smooth; chartaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum fully concealed; concealed by funicular remnant; without faboid split; punctiform; marginal according to radicle tip; flush (and dark brown). Lens not discernible. Endosperm absent. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; similar at apex; not Belairia: B. mucronata A.H.R. Grisebach (AÐE). A, Fruits concealing radicle; split over radicle; with lobes; with (ϫ 3.2); B, seed (ϫ 4.5); CÐD, testa (ϫ 50, ϫ 1000); E, lobes not touching; with the interface division terminat- embryos (ϫ 10).

763 Genus: Pictetia A.-P. de Candolle glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum fully concealed; Phylogenetic Number: 14.05. concealed by aril; without faboid split (visible); punctiform; between cotyledon and radicle lobe; flush; Tribe: Aeschynomeneae. not within corona, halo, or rim. Lens discernible; less than 0.5 mm in length; with margins straight or curved; Subtribe: Ormocarpinae. oblong; not in groove of raphe; 180 degrees from hilum; flush; dissimilar color from testa; darker than Species Studied—Species in Genus: 4 spp.—ca. 6 spp. testa; black (ish); not within corona, halo, or rim. Endosperm thin; 1/2 covering entire embryo; adnate to Fruit a legume or loment; unilocular; 2.5Ð4 ϫ 0.5Ð1 ϫ testa. Cotyledons smooth; both outer faces convex; both 0.09Ð0.2 cm; with deciduous corolla; with persistent or the same thickness; both more or less of equal length; deciduous calyx; with calyx shorter than fruit; without not folded; margin entire 180 degrees from base of orifice formed by curving of fruit or fruit segments; radicle; similar at apex; partially concealing radicle; straight to curved (to slightly curved); not plicate; not split over radicle; with lobes; with lobes not touching; twisted; asymmetrical or symmetrical; broadly linear to with the interface division terminating at base of oblong; when asymmetrical with both sutures parallelly radicle; without margins recessed; white; inner face curved; not inflated; flattened to compressed; without flat; glabrous around base of radicle. Embryonic axis beak; short tapered to rounded at apex; apex aligned deflexed; oblique to length of seed. Radicle linear; lobe with longitudinal axis of fruit; short tapered to rounded tip straight; deflexed and parallel to cotyledon width; at base; base aligned with longitudinal axis of fruit; centered between cotyledons; less than 1/2 length of with the apex and base uniform in texture; coriaceous; cotyledons. Plumule moderately developed; glabrous. seed chambers externally visible or invisible; with the raised seed chambers not torulose. Fruit margin Distribution: West Indies. constricted or not constricted; constricted along both margins; without sulcus; plain. Fruit wings absent. Fruit substipitate to nonstipitate. Fruit indehiscent. Replum invisible. Loment indehiscent. Loment segments (articles) inconspicuous; 7Ð10 mm long; widest across seed area; with all essentially similar in shape; oblong. Epicarp dull to glossy; monochrome; brown; pubescent and indurate; with 1 type of pubescence; puberulent; with pubescence gray; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; tuberculate or not tuberculate; with solid tubercles on each valve; tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; mealy; chartaceous. Endocarp dull; monochrome; tan; smooth; septate or nonseptate; with septa thicker than paper, firm; with the septa eglandular; coriaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð5; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; contorted. Aril dry; rim-aril; tan.

Seed 5Ð5.3 ϫ 3Ð3.2 ϫ 0.3Ð0.4 mm; not overgrown; not angular; asymmetrical; C-shaped; flattened; with visible radicle and cotyledon lobes (barely); without umbo on Pictetia: P. spinifolia (A.N. Desvaux) I. Urban (BÐE), P. seed faces. Testa not adhering to endocarp; glossy; not spp. (A). A, Fruit and partial fruit (ϫ 2.6); B, seeds modified by a bloom; colored; monochrome; tan; (ϫ 5.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

765 Genus: Fiebrigiella H.A.T. Harms brown; glabrous; smooth or not smooth; with elevated features; wrinkled (1 longitudinal wrinkle); Phylogenetic Number: 14.06. chartaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; Tribe: Aeschynomeneae. concealed by aril (rim); without faboid split; punctiform; marginal according to radicle tip; flush; not Subtribe: Ormocarpinae. within corona, halo, or rim. Lens discernible; less than 0.5 mm in length; with margins curved; circular; not in Species Studied—Species in Genus: 1 sp.—1 sp. groove of raphe; adjacent to hilum; 1 mm from hilum; mounded; dissimilar color from testa; darker than testa; Fruit a loment; 3Ð3.1 ϫ 0.3Ð0.4 ϫ 0.08Ð0.1 cm; with black (ish); not within corona, halo, or rim. Endosperm deciduous corolla; with persistent or deciduous calyx; absent. Cotyledons smooth; both outer faces convex; with calyx shorter than fruit; without orifice formed by both the same thickness; both more or less of equal curving of fruit or fruit segments; straight to curved length; not folded; margin entire 180 degrees from base (slightly); not plicate; not twisted; asymmetrical; when of radicle; similar at apex; not concealing radicle; split asymmetrical with 1 straight and 1 curved suture; over radicle; with lobes; with lobes not touching; narrowing in several places, resembling Desmodium without basal groin formed by lobes; with the interface fruit; not inflated; flattened; without beak; short tapered division terminating at base of radicle; without margins at apex; apex aligned to oblique with longitudinal axis recessed; brown; inner face flat; glabrous around base of fruit; tapered to short tapered at base; base aligned of radicle. Embryonic axis straight; oblique to length of with longitudinal axis of fruit; with the apex and base seed; without a joint evident between the radicle and uniform in texture; coriaceous; seed chambers exter- the cotyledons. Radicle differentiated from cotyledon; nally visible; with the raised seed chambers not bulbose; lobe tip straight; oblique to cotyledons; torulose. Fruit margin constricted only on 1 margin; centered between cotyledons; less than 1/2 length of without sulcus; plain. Fruit wings absent. Fruit cotyledons. Plumule rudimentary; glabrous. substipitate. Fruit indehiscent. Replum invisible. Loment indehiscent. Loment segments (articles) Distribution: Bolivia. inconspicuous; 5Ð6 mm long; widest across seed area; with all essentially similar in shape; D-shaped. Epicarp Notes: Burkart and Vilchez (1971) monographed the genus, dull; monochrome; dirty brown; pubescent but soon and their figure 2 was consulted in preparing our plate. deciduous; with 1 type of pubescence; puberulent; with Scant seeds and fruits were available for our study. pubescence golden; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined (with 1 major vein and 1 minor longitudinal vein); not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; tan; spongy; septate; with septa thicker than paper, firm; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 2Ð4; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril dry; rim-aril; reddish brown.

Seed 4.2Ð5 ϫ 2.5Ð3 ϫ 0.7Ð1 mm; not overgrown; not angular; asymmetrical; D-shaped; flattened; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not Fiebrigiella: F. gracilis H.A.T. Harms (AÐF). A, Fruit modified by a bloom; colored; monochrome; reddish (ϫ 5.9); B, fruit and article (ϫ 3); C, seed (ϫ 8); DÐE, testa (ϫ 50, ϫ 1000); F, embryos (ϫ 8).

767 Genus: Chaetocalyx A.-P. de Candolle visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to partially adhering to Phylogenetic Number: 14.07. endocarp; glossy; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; Tribe: Aeschynomeneae. coriaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum visible; with faboid split; with the Subtribe: Ormocarpinae. lips of the faboid split the same color as the rest of the hilum; punctiform or larger than punctiform; 0.3 mm Species Studied—Species in Genus: 10 spp.—12 spp. long; with curved outline; circular; between cotyledon and radicle lobe; flush; within rim. Hilum rim color Fruit a loment; 7Ð16 ϫ 0.2Ð2.2 ϫ 0.07Ð0.2 cm; with darker than testa (slightly). Lens discernible; less than deciduous corolla; with persistent or deciduous calyx; 0.5 mm in length (ca. 0.3 mm); with margins curved; with calyx shorter than fruit; without orifice formed by circular; not in groove of raphe; adjacent to hilum; 0.1 curving of fruit or fruit segments; straight, curved (or mm from hilum; mounded; similar color as testa; darker slightly curved), or S-curved; not plicate; not twisted; than testa; brown; not within corona, halo, or rim. asymmetrical or symmetrical; linear; when asymmetri- Endosperm thick; 1/2 covering entire embryo; adnate to cal with both sutures parallelly curved; not inflated; testa. Cotyledons smooth; both outer faces convex; both flattened or compressed; without beak; tapered or short the same thickness; both more or less of equal length; tapered at apex; apex aligned or oblique with longitudi- not folded; margin entire 180 degrees from base of nal axis of fruit; short tapered or rounded at base; base radicle; similar at apex; not concealing radicle; entire aligned with longitudinal axis of fruit; with the apex over radicle; without lobes; with the interface division and base uniform in texture; membranous; seed terminating at base of radicle; without margins re- chambers externally invisible. Fruit margin not con- cessed; yellow; inner face flat; glabrous around base of stricted; without sulcus; plain or embellished; with radicle. Embryonic axis deflexed; oblique to length of wing. Fruit wing 1; 8Ð10 mm wide; continuous wing seed. Radicle bulbose; lobe tip curved; deflexed and around fruit. Fruit stipitate, substipitate, or nonstipitate; parallel to cotyledon width; centered between cotyle- with the stipe 5Ð15 mm long. Fruit indehiscent. Replum dons; less than 1/2 length of cotyledons. Plumule invisible. Loment indehiscent. Loment segments rudimentary; glabrous. (articles) inconspicuous; 4Ð17 mm long; widest across seed area; with upper 1 different shape than middle Distribution: Tropical America. ones; oblong or quadrangular. Epicarp dull; monochrome or multicolored; bichrome (flattened fruit have Notes: Rudd (1958) monographed the genus and Diatloff a darker center (over seed chambers) that fades to and Diatloff (1977) noted that Chaetocalyx is a non- lighter margins); brown or tan; glabrous, glabrate, or nodulating faboid legume genus. pubescent and indurate; with 1 type of pubescence; velutinous; with pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; glandular; with glandular setae (few); without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thick; surface not veined; 1- layered; without balsamic vesicles; without fibers; solid; ligneous (sub). Endocarp dull; monochrome; gray; cobwebby; septate; with septa thicker than paper, firm; with the septa eglandular; coriaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 6Ð16; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; Chaetocalyx: C. brasiliensis (J.R.T. Vogel) G. Bentham (E), of 1 length only; straight. Aril absent. C. latisiliqua (J.L.M. Poiret) G. Bentham ex W.B. Hemsley (CÐD), C. spp. (AÐB). A, Fruits (none entire) Seed 2.5Ð7 ϫ 2Ð2.5 ϫ 2Ð2.5 mm; not overgrown; not (ϫ 1.4); B, seeds (ϫ 6); CÐD, testa (ϫ 50, ϫ 1000); E, angular; asymmetrical; linear or reniform; terete; with embryos (ϫ 10).

769 Genus: Nissolia N. von Jacquin Seed 2Ð5 ϫ 1.5Ð3.5 ϫ 1.7 mm; not overgrown; not angular; asymmetrical; oblong, quadrangular (nearly), or Phylogenetic Number: 14.08. reniform; compressed; with visible radicle and cotyledon lobes; without external groove between radicle and Tribe: Aeschynomeneae. cotyledon lobes; without umbo on seed faces. Testa partially adhering to endocarp (hairs); more or less Subtribe: Ormocarpinae. glossy; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; coriaceous. Species Studied—Species in Genus: 11 spp.—13 spp. Fracture lines absent. Rim absent. Wings absent. Raphe from hilum to lens; not bifurcating; darker than testa; Fruit a loment; 1.5Ð4.3 ϫ 0.5Ð1.5 ϫ 0.09Ð0.3 cm; with black; flush. Hilum visible; with faboid split; with the persistent or deciduous androecial sheath; with decidu- lips of the faboid split the same color as the rest of the ous corolla; with persistent or deciduous calyx (rarely); hilum; punctiform to larger than punctiform; up to 0.3 with calyx shorter than fruit; without orifice formed by mm long; with curved outline; circular; between curving of fruit or fruit segments; straight or curved; cotyledon and radicle lobe; flush; not within corona, not plicate; not twisted; asymmetrical or symmetrical; halo, or rim or within halo. Hilum halo color lighter oblong or falcate; when asymmetrical with both sutures than testa. Lens not discernible. Endosperm thick; 1/2 parallelly curved; not inflated; compressed; without covering entire embryo; adnate to testa. Cotyledons beak; tapered to rounded at apex; apex aligned to smooth; both outer faces convex; both the same oblique with longitudinal axis of fruit; short tapered to thickness; both more or less of equal length; not folded; rounded at base; base aligned to oblique with longitudi- margin entire 180 degrees from base of radicle; similar nal axis of fruit; with the apex and base uniform in at apex; partially concealing or not concealing radicle; texture; coriaceous; seed chambers externally visible; split over radicle; with lobes; with lobes not touching; with the raised seed chambers not torulose. Fruit with the interface division terminating at base of margin slightly constricted along both margins; without radicle; without margins recessed; yellow; inner face sulcus; plain or embellished; with wing (restricted to flat; glabrous around base of radicle. Embryonic axis apical article). Fruit wing present or absent; 1; 5Ð15 deflexed; oblique to length of seed. Radicle bulbose mm wide; samaroid; apical. Fruit stipitate to (nearly); lobe tip straight; deflexed and parallel to substipitate to nonstipitate; with the stipe up to 6 mm cotyledon width; centered between cotyledons; less long. Fruit indehiscent (if 1-seeded). Replum invisible. than 1/2 length of cotyledons. Plumule moderately Loment indehiscent. Loment segments (articles) developed; glabrous. conspicuous; 3Ð10 mm long (sterile (winged) 10Ð25 mm); widest across seed area; with upper 1 different Distribution: Mexico, West Indies, Central America, South shape than middle ones; oblong (with apical article with America. terminal wing) or rectangular. Epicarp dull; monochrome; reddish brown or tan; glabrous or pubescent Notes: Rudd (1956) monographed the genus and noted that and indurate; with 1 type of pubescence; crinkled the “fruits are samaralike, 2–5-articulate loments with pilose; with pubescence gray; with pubescence uni- terminal joint sterile, flat, and winglike.” formly distributed; with simple or glandular hairs; pliable; with hair bases plain; glandular or eglandular; with glandular setae; without spines; not smooth; with elevated features; longitudinally veined relative to fruit length (with 3 to many well developed veins) or reticulately veined; not tuberculate; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; cobwebby; septate; with septa thicker than paper, firm; coriaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð Nissolia: N. leiogyne N.Y. Sandwith (E), N. schottii (J. 6; length parallel with fruit length; neither overlapping Torrey) A. Gray (CÐD), N. spp. (AÐB). A, Fruits and 1 nor touching; in 1 series. Funiculus less than 0.5 mm article (ϫ 1.9); B, seeds (ϫ 5.3); CÐD, testa (ϫ 50, long; of 1 length only; straight. Aril dry; rim-aril; tan. ϫ 1000); E, embryos (ϫ 5).

771 Genus: Aeschynomene C. Linnaeus septa thicker than paper, firm; with the septa eglandular; ligneous (sub); not exfoliating; remaining Phylogenetic Number: 14.09. fused to epicarp; entire. Seeds 1Ð18; length parallel with fruit length; neither overlapping nor touching; in 1 Tribe: Aeschynomeneae. series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent. Subtribe: Aeschynomeninae. Seed 1Ð6 ϫ 0.7Ð5 ϫ 2Ð2.5 mm; not overgrown; not Species Studied—Species in Genus: 21 spp.—ca. 150 spp. angular; asymmetrical; reniform; compressed; with visible radicle and cotyledon lobes; without umbo on Fruit a loment; 0.8Ð2 ϫ 0.4Ð0.5 ϫ 0.1Ð0.3 cm; with seed faces. Testa not adhering to endocarp; glossy; deciduous corolla; with persistent or deciduous calyx; colored; monochrome; black (and greenish) or brown with calyx shorter than fruit (in loments that do not (light); smooth; coriaceous. Fracture lines absent. Rim articulate); with or without orifice formed by curving of absent. Raphe not visible. Hilum fully concealed; fruit or fruit segments; straight, curved (or slightly concealed by funicular remnant; with faboid split; with curved), S-curved, or 0.5-, 1-, 1.5-, 4-, or 2-coiled; not the lips of the faboid split the same color or lighter plicate; not twisted; asymmetrical or symmetrical; colored than the rest of the hilum and therefore con- circular, linear, moniliform, or falcate; when asym- spicuous; larger than punctiform; 1.5 mm long; with metrical with 1 straight and 1 curved suture or both curved outline; elliptic; marginal according to radicle sutures parallelly curved; narrowing in several places, tip or between cotyledon and radicle lobe; flush; within resembling Desmodium (11.09) fruit; not inflated; halo. Hilum halo color lighter than testa. Lens not compressed or quadrangular; without beak; tapered at discernible. Endosperm thin; 1/2 covering entire apex; apex aligned with longitudinal axis of fruit; embryo; adnate to testa. Cotyledons smooth; both outer tapered at base; base aligned with longitudinal axis of faces convex; both the same thickness; both more or fruit; with the apex and base uniform in texture; less of equal length; not folded; margin entire 180 coriaceous or fragile, thinner than chartaceous like degrees from base of radicle; similar at apex; not Trifolium (21.06); seed chambers externally visible; concealing radicle; split over radicle; with lobes; with with the raised seed chambers not torulose. Fruit lobes not touching; with the interface division terminat- margin constricted or not constricted; constricted along ing at base of radicle; without margins recessed; inner both margins or only on 1 margin; without sulcus; face flat; glabrous around base of radicle. Embryonic plain. Fruit wings absent. Fruit stipitate to substipitate; axis deflexed; perpendicular to length of seed. Radicle with the stipe 1Ð25 mm long. Fruit indehiscent. Replum bulbose; lobe tip straight or curved; deflexed and occasionally visible. Fruit a segment; an intact article; parallel to cotyledon width; centered between cotyle- loment segment. Loment indehiscent. Loment segments dons; 1/2 to nearly length of cotyledons. Plumule (articles) inconspicuous; 3Ð7 mm long; widest across rudimentary or well developed; glabrous. seed area; with all essentially similar in shape; D- shaped or rectangular. Epicarp dull; monochrome; Distribution: Tropics to warm temperate America, Africa, black, brown, green (ish-black), or tan; with surface and Asia. texture uniform or not uniform, with patches of different texture not restricted to the base and apex; Notes: Rudd (1955) monographed the American species glabrous or pubescent and indurate; with 1 type of and, according to her subgeneric categories, the U.S. pubescence; appressed puberulent or tomentose; with National Seed Herbarium (BARC) has representatives pubescence gray; with pubescence uniformly distrib- of both subgenera and seven of eight series. The uted; with simple or glandular hairs; pliable; with hair missing series included two species: A. fluminensis J. bases plain; glandular (sections Aeschynomene and Velloso de Miranda and A. parviflora M. Micheli. Of Ochopodium); with glandular hairs; without spines; the studied species, only three are restricted to the Old smooth or not smooth; with elevated features; veined or World. Rudd also provided seed and fruit data for each not veined; obliquely veined relative to fruit length; not American taxon. Verdcourt (1974) monographed tuberculate; verrucose-rugose, muricate, or faveolate; Aeschynomene for the Flora of Zambia and included not exfoliating; without cracks. Mesocarp absent. seed and fruit data. Pandey and Jha (1989) reported on Endocarp dull; monochrome; tan; smooth; septate; with the seed structure of A. aspera C. Linnaeus (type for genus) and A. indica C. Linnaeus. Rudd (1955) re-

772 corded the variable epicarp surfaces of articles using these descriptors: (Hairs) appressed-pubescent, ciliate, crisp-pubescent, glabrate, glabrous, glandular- hispidulous, hispid, hispidulous, pubescent, sericeous, subappressed hairs, subglabrous, villous-hispid, white- pubescent; (surface) faveolate, muricate, muricate in center of article, reticulate-veiny, rugose, smooth, tuberculate, tuberculate bases of hairs..., veiny, ventri- cose, verrucose, and verrucose at center of article. All species of Aeschynomene are noxious weeds in rice (Oryza sativa C. Linnaeus) in Arkansas.

773 Aeschynomene: A. virginica (C. Linnaeus) N.L. Britton, E.E. Sterns & J.F. Poggenberg (CÐE), A. spp. (AÐB). A, Articles and broken and entire fruits (ϫ 1.9); B, seeds (ϫ 3.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

775 Genus: Soemmeringia C.F.P. von Martius within corona, halo, or rim. Lens discernible; less than 0.5 mm in length; with margins straight or curved; Phylogenetic Number: 14.10. irregular; irregular; not in groove of raphe; adjacent to hilum; 0.2 mm from hilum; mounded; dissimilar color Tribe: Aeschynomeneae. from testa; darker than testa; black (on dark brown testa) or brown (on green testa); not within corona, Subtribe: Aeschynomeninae. halo, or rim. Endosperm thick; 1/2 covering entire embryo; adnate to testa. Cotyledons smooth; both outer Species Studied—Species in Genus: 1 sp.—1 sp. faces convex; both the same thickness; both more or less of equal length; not folded; margin not entire 180 Fruit a loment; 0.4Ð0.5 ϫ 0.18Ð0.2 ϫ 0.05Ð0.06 cm; with degrees from base of radicle; similar at apex; partially persistent or deciduous androecial sheath; with persis- concealing radicle; split over radicle; with lobes; with tent or deciduous corolla; with various petals; with lobes not touching; with the interface division terminat- persistent calyx; with calyx longer than fruit; without ing at base of radicle; without margins recessed; orifice formed by curving of fruit or fruit segments; yellow; inner face flat; glabrous around base of radicle. somewhat contorted; somewhat plicate to not plicate; Embryonic axis deflexed; oblique to length of seed. not twisted; asymmetrical; moniliform; when asym- Radicle nearly bulbose; lobe tip straight; deflexed and metrical with both sutures parallelly curved; not parallel to cotyledon width; centered between cotyle- inflated; compressed; without beak; rounded at apex; dons; less than 1/2 length of cotyledons. Plumule apex aligned with longitudinal axis of fruit; rounded at moderately developed; glabrous. base; base aligned with longitudinal axis of fruit; with the apex and base uniform in texture; chartaceous; seed Distribution: Venezuela, Brazil, and Bolivia. chambers externally visible; with the raised seed chambers not torulose. Fruit margin constricted along Notes: Apparently the marcescent corolla acts as a wing in both margins; without sulcus; plain. Fruit wings absent. fruit dispersal and therefore aids in seed dissemination. Fruit nonstipitate. Fruit indehiscent. Replum invisible. Loment indehiscent. Loment segments (articles) inconspicuous; 1.5Ð2 mm long; widest across seed area; with all essentially similar in shape; circular. Epicarp dull; monochrome; brown; glabrous; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 2Ð4; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened; straight. Aril dry; rim-aril; reddish brown.

Seed 1 ϫ 1 ϫ 0.3 mm; not overgrown; not angular; asymmetrical; mitaform; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome; brown (dark brown) or green; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible or fully concealed; concealed by funicu- Soemmeringia: S. semperflorens C.F.P. von Martius (AÐF). lar remnant; with faboid split; with the lips of the faboid A, Partial fruit (ϫ 14); B, fruit (left) and fruits in split the same color as the rest of the hilum; punctiform; marcescent corollas (right) (ϫ 3.7); C, seeds (ϫ 9); between cotyledon and radicle lobe; recessed; not DÐE, testa (ϫ 50, ϫ 1000); F, embryos (ϫ 15).

777 Genus: Kotschya S.F.L. Endlicher black, brown (reddish or gray), green, or yellow; glabrous; smooth; osseous. Fracture lines absent. Rim Phylogenetic Number: 14.11. absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the Tribe: Aeschynomeneae. same color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; flush; not within Subtribe: Aeschynomeninae. corona, halo, or rim. Lens discernible; equal to or greater than 0.5 mm in length or less than 0.5 mm in Species Studied—Species in Genus: 7 spp.—ca. 30 spp. length; 0.5 mm long; with margins straight; linear; not in groove of raphe; adjacent to hilum; less than 0.5 mm Fruit a loment; 0.2Ð3.9 (see Notes) 0.2Ð0.4 0.15 cm; from hilum; recessed; dissimilar color from testa; with persistent corolla; with standard; with persistent darker than testa; black; not within corona, halo, or rim. calyx; with calyx longer than fruit; without orifice Endosperm absent. Cotyledons smooth; both outer formed by curving of fruit or fruit segments; straight; faces convex; both the same thickness; both more or plicate (see Notes); not twisted; asymmetrical; when less of equal length; not folded; margin entire 180 asymmetrical with 1 straight and 1 curved suture; degrees from base of radicle; similar at apex; partially narrowing in several places, resembling Desmodium or not concealing radicle; split over radicle; with lobes; fruit; not inflated; compressed; without beak; rounded with lobes not touching; with the interface division at apex; apex aligned with longitudinal axis of fruit; terminating at base of radicle; without margins re- rounded at base; base aligned with longitudinal axis of cessed; red; inner face flat; glabrous around base of fruit; with the apex and base uniform in texture; radicle. Embryonic axis deflexed; oblique to length of chartaceous; seed chambers externally visible; with the seed. Radicle linear; lobe tip straight or curved; oblique raised seed chambers not torulose. Fruit margin to cotyledons; centered between cotyledons; less than constricted only on 1 margin; without sulcus; plain. 1/2 length of cotyledons. Plumule rudimentary. Fruit wings absent. Fruit substipitate. Fruit indehiscent. Replum invisible. Loment indehiscent. Loment Distribution: Tropical Africa and Madagascar. segments (articles) inconspicuous; 2Ð6 mm long; widest across seed area; with all essentially similar in Notes: Verdcourt (1974) described 19 species from Zambia shape; D-shaped. Epicarp dull; monochrome; brown and noted that if the fruit is composed of several (including reddish) or tan; glabrous or pubescent and articles, then the fruit is “folded like a concertina.” indurate; with 2 types of pubescence; pilose; with Upon opening the calyx, the fruit disarticulates, making pubescence gray and yellow; with golden hooked hairs length measurements difficult. Our fruit length range is and gray plain hairs; with pubescence uniformly based on maximum and minimum number of articles distributed; with simple hairs; pliable; with hair bases multiplied by the maximum and minimum article swollen or plain; without spines; not smooth; with lengths. elevated features; reticulately veined; not tuberculate; tuberculate (bases of hairs); not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; septate; with septa thin (tissue paper-like), flexible; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð9; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril dry; rim-aril; yellow.

Seed 2Ð4.7 1.2Ð3 0.5Ð1.7 mm; not overgrown; not Kotschya: K. africana S.F.L. Endlicher var. bequaertii angular; asymmetrical; elliptic, circular (sub), reniform (E.A.J. De Wildeman) B. Verdcourt (DÐF), K. spp. (to ellipsoid-reniform), or triangular; compressed; with (AÐC). A, Articles, fruits in calyx and corolla ( 4.3); visible radicle and cotyledon lobes; without umbo on B, fruit in flower ( 3); C, seeds ( 5.8); DÐE, testa seed faces. Testa not adhering to endocarp; glossy; not ( 50, 1000); F, embryos ( 8). modified by a bloom; colored; monochrome; reddish

779 Genus: Smithia W. Aiton raphe; adjacent to hilum; mounded; dissimilar color from testa; darker than testa; black; not within corona, Phylogenetic Number: 14.12. halo, or rim. Endosperm thin; 1/2 covering entire embryo; adnate to testa. Cotyledons smooth; both outer Tribe: Aeschynomeneae. faces convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 Subtribe: Aeschynomeninae. degrees from base of radicle; differing at apex (1 concealed by overarching radicle and other auriculate Species Studied—Species in Genus: 12 spp.—ca. 30 spp. and concealing radicle); partially concealing radicle; entire over radicle; without lobes; with the interface Fruit a loment; 1Ð1.5 (estimated) 0.15Ð0.3 0.2Ð0.22 division terminating at base of radicle; without margins cm; with deciduous corolla; with persistent calyx; with recessed; yellow; inner face flat; glabrous around base calyx longer than fruit; without orifice formed by of radicle. Embryonic axis deflexed; oblique to length curving of fruit or fruit segments; contorted; plicate; not of seed. Radicle linear; lobe tip straight; deflexed and twisted; asymmetrical; moniliform; when asymmetrical parallel to cotyledon length or width; not centered with both sutures parallelly curved; not inflated; between cotyledons (radicle outside 1 cotyledon and compressed; without beak; rounded at apex; apex inside other, therefore junctions for each cotyledon aligned with longitudinal axis of fruit (probably); different); 1/2 to nearly length of cotyledons. Plumule rounded at base; base aligned with longitudinal axis of moderately developed; glabrous. fruit (probably); with the apex and base uniform in texture; chartaceous; seed chambers externally visible. Distribution: Old World tropics, mainly Asia, Madagascar, Fruit margin constricted along both margins; without and Africa. sulcus; plain. Fruit wings absent. Fruit substipitate. Fruit indehiscent. Replum invisible. Loment indehis- Notes: Dewit and Duvigneaud (1954) evaluated the species cent. Loment segments (articles) inconspicuous; 2Ð3 of Smithia in the Congo. The testa is thin, and the mm long; widest across seed area; with all essentially cotyledons are soft, making it difficult to remove the similar in shape; circular. Epicarp dull; monochrome; embryo. This is unlike most faboid legume seeds, glabrous; without spines; smooth or not smooth; with whose embryos are easy to remove from the testa. elevated features; veined or not reticulately veined; tuberculate or not tuberculate; with solid tubercles on each valve; tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 4Ð7; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only. Aril absent.

Seed 1.5Ð1.8 1.3 0.5 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; Smithia: S. elliotti E.G. Baker var. elliotti (B), S. purpurea between cotyledon and radicle lobe; flush; not within W.J. Hooker (DÐF), S. spp. (A, C). A, Fruits and fruits corona, halo, or rim or within halo. Hilum halo color within calyx and bracts ( 5); B, young fruit ( 10, darker than testa. Lens discernible; less than 0.5 mm in modified from Gillett et al. 1971); C, seeds ( 8); DÐE, length; with margins curved; elliptic; not in groove of testa ( 50, 1000); F, embryos ( 15).

781 Genus: Geissaspis R. Wight & G.A.W. Arnott visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not Phylogenetic Number: 14.13. modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; osseous. Fracture lines Tribe: Aeschynomeneae. absent. Rim absent. Wings absent. Raphe visible or not visible; from hilum to lens; not bifurcating; color of Subtribe: Aeschynomeninae. testa or darker than testa; black (ish); recessed. Hilum visible; with faboid split; with the lips of the faboid Species Studied—Species in Genus: 2 spp.—3 spp. split the same color as the rest of the hilum; punctiform; marginal according to radicle tip or between cotyledon Fruit a legume or loment; unilocular; 0.35Ð0.8 0.25Ð0.35 and radicle lobe; recessed; within rim or not within 0.1Ð0.15 cm; with persistent androecial sheath; with corona, halo, or rim. Hilum rim color of testa. Lens deciduous corolla; with persistent or deciduous calyx; discernible; equal to or greater than 0.5 mm in length; with calyx shorter than fruit; without orifice formed by 1Ð2 mm long; with margins curved; more or less curving of fruit or fruit segments; straight; not plicate; circular; not in groove of raphe; adjacent to hilum; 2 not twisted; asymmetrical; when asymmetrical with 1 mm from hilum; mounded; dissimilar color from testa; straight and 1 curved suture; widest near middle or D- darker than testa; black (ish); not within corona, halo, shaped; not inflated; compressed; without or with beak; or rim. Endosperm thin; 1/2 covering entire embryo; straight or declined; with solid beak the same color and adnate to testa. Cotyledons smooth; both outer faces texture as fruit (if present, quite small); rounded at convex; both the same thickness; both more or less of apex; apex oblique with longitudinal axis of fruit; equal length; not folded; margin entire 180 degrees rounded at base; base aligned with longitudinal axis of from base of radicle; similar at apex; partially conceal- fruit (more or less); with the apex and base uniform in ing to not concealing radicle; split over radicle; with texture; chartaceous; seed chambers externally visible; lobes; with lobes touching (auriculate); with the with the raised seed chambers not torulose. Fruit interface division terminating at base of radicle; without margin constricted along both margins; without sulcus; margins recessed; yellow; inner face flat; glabrous plain or embellished; with wing. Fruit wing 1 (article); around base of radicle. Embryonic axis deflexed; 0.5Ð0.6 mm wide; continuous wing around fruit; on 1 oblique to length of seed. Radicle more or less bulbose; suture. Fruit substipitate. Fruit indehiscent. Replum lobe tip straight; oblique to cotyledons; centered invisible. Loment indehiscent. Loment segments between cotyledons; less than 1/2 length of cotyledons. (articles) inconspicuous; 5Ð6 mm long; widest across Plumule rudimentary; glabrous. seed area; with all essentially similar in shape; D- shaped. Epicarp dull; monochrome or multicolored; Distribution: Tropical and subtropical Asia. streaked; brown or tan; with red overlay; with mottling over seed chambers; pubescent and indurate; with 1 Notes: Duvigneaud (1954b) evaluated the genus. type of pubescence; with pubescence brown; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; with cracks (if cracked because fruit is fragile) or without cracks; cracking irregular. Mesocarp absent. Endocarp dull; monochrome; brown or tan; smooth; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent. Geissaspis: G. cristata R. Wight & G.A.W. Arnott (CÐE), Seed 2.2Ð2.8 1.8Ð2.2 0.9Ð1.1 mm; not overgrown; not G. spp. (AÐB). A, Articles and fruits concealed in bracts angular; asymmetrical; reniform; compressed; with ( 4.5); B, seeds ( 8.6); CÐD, testa ( 50, 1000); E, embryos ( 8).

783 Genus: Bryaspis P. Duvigneaud equal to or greater than 0.5 mm in length; 0.5 mm long; with margins straight; linear (centered within a discol- Phylogenetic Number: 14.14. ored area); not in groove of raphe; confluent with hilum; recessed; dissimilar color from testa; darker than Tribe: Aeschynomeneae. testa; black; not within corona, halo, or rim. Endosperm absent. Cotyledons smooth; both outer faces convex; Subtribe: Aeschynomeninae. both the same thickness; both more or less of equal length; not folded; margin entire 180 degrees from base Species Studied—Species in Genus: 1 sp.—2 spp. of radicle; similar at apex; partially concealing radicle; split over radicle; with lobes; with lobes not touching; Fruit a loment; 0.8Ð1.3 0.2 0.14Ð0.15 cm; with with the interface division terminating at base of deciduous corolla; with deciduous calyx (within large radicle; without margins recessed; yellow; inner face bract); without orifice formed by curving of fruit or flat; glabrous around base of radicle. Embryonic axis fruit segments; straight; not plicate; not twisted; deflexed; oblique to length of seed. Radicle bulbose; asymmetrical; when asymmetrical with 1 straight and 1 lobe tip straight; deflexed and parallel to cotyledon curved suture; narrowing in several places, resembling width; centered between cotyledons; less than 1/2 Desmodium fruit; not inflated; compressed; without length of cotyledons. Plumule rudimentary; glabrous. beak; rounded at apex; apex oblique with longitudinal axis of fruit; rounded at base; base aligned with Distribution: Western tropical Africa. longitudinal axis of fruit; with the apex and base uniform in texture; chartaceous; seed chambers Notes: Duvigneaud (1954b) separated Geissaspis (14.13) externally visible; with the raised seed chambers not from Bryaspis by stipule, bract, standard, keel, and fruit torulose. Fruit margin constricted only on 1 margin; characters. His fruit key characters were: Fruit mem- without sulcus; plain. Fruit wings absent. Fruit branaceous and article densely reticulate for Geissaspis nonstipitate. Fruit indehiscent. Replum invisible. Fruit a and fruit firmer and article faintly reticulate for segment; an intact article; entire. Loment indehiscent. Bryaspis. Loment segments (articles) inconspicuous; 2Ð3 mm long; widest across seed area; with all essentially similar in shape; D-shaped. Epicarp dull; monochrome; reddish brown; glabrous; eglandular; without spines; not smooth; with elevated features; irregularly veined; not tuberculate; subvesicular; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; reddish brown; smooth; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 2Ð2.5 1.6Ð2 1.2Ð1.5 mm; not overgrown; not angular; asymmetrical; D-shaped (with radicle lobe); terete; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; tan; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than Bryaspis: B. lupulina (L.D. Planchon ex G. Bentham) P. punctiform or punctiform; 0.3 mm long; with curved Duvigneaud (AÐE). A, Articles and fruit within bracts outline; circular; between cotyledon and radicle lobe; ( 1.9); B, seeds ( 5); CÐD, testa ( 50, 1000); E, flush; not within corona, halo, or rim. Lens discernible; embryos ( 10).

785 Genus: Humularia P. Duvigneaud visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not Phylogenetic Number: 14.15. modified by a bloom; colored; monochrome; purplish black or brown (reddish); smooth; coriaceous. Fracture Tribe: Aeschynomeneae. lines absent. Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by Subtribe: Aeschynomeninae. funicular remnant; without faboid split; larger than punctiform; 0.3Ð0.4 mm long; with curved outline; Species Studied—Species in Genus: 18 spp.—ca. 40 spp. circular; between cotyledon and radicle lobe; flush; not within corona, halo, or rim. Lens discernible or not Fruit a legume or loment; unilocular; 0.5Ð1.5 0.4Ð0.55 discernible; less than 0.5 mm in length (0.1Ð0.2 mm); 0.3 cm; with persistent or deciduous androecial sheath; with margins straight or curved; irregular, linear, or with persistent or deciduous corolla; with various wedge-shaped; irregular; not in or in groove of raphe; petals; with persistent calyx; with calyx shorter than adjacent to hilum; flush or recessed; dissimilar color fruit; without orifice formed by curving of fruit or fruit from testa; darker than testa; black; not within corona, segments; straight; not plicate; not twisted; asymmetri- halo, or rim. Endosperm thin;1/2 covering entire cal; when asymmetrical with 1 straight and 1 curved embryo; adnate to testa. Cotyledons smooth; both outer suture; narrowing in several places, resembling faces convex; both the same thickness; both more or Desmodium (11.09) fruit; not inflated; flattened; less of equal length; not folded; margin entire 180 without or with beak; with solid beak the same color degrees from base of radicle; similar at apex; partially and texture as fruit; rounded at apex; apex aligned with concealing radicle; split over radicle; with lobes; with longitudinal axis of fruit; short tapered at base; base lobes not touching; with the interface division terminat- aligned with longitudinal axis of fruit; with the apex ing at base of radicle; without margins recessed; red; and base uniform in texture; coriaceous; seed chambers inner face flat; glabrous around base of radicle. Embry- externally visible; with the raised seed chambers not onic axis oblique; oblique to length of seed. Radicle torulose. Fruit margin constricted on 1 margin and bulbose; lobe tip straight; oblique to cotyledons; slightly constricted on the other margin; without sulcus; centered between cotyledons; less than 1/2 or 1/2 to plain. Fruit wings absent. Fruit nonstipitate. Fruit nearly length of cotyledons. Plumule rudimentary; indehiscent. Replum invisible. Loment indehiscent. glabrous. Loment segments (articles) inconspicuous; 6.5 mm long; widest across seed area; with all essentially Distribution: Africa. similar in shape; D-shaped. Epicarp dull; monochrome; brown; glabrous or pubescent but soon deciduous; with Notes: Duvigneaud (1954a) analyzed the tropical African 1 type of pubescence; puberulent; with pubescence species and Verdcourt (1974) described the species for gray; with pubescence uniformly distributed; with Flora of Zambia. simple or glandular hairs; pliable; with hair bases plain; glandular or eglandular; with glandular hairs; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; warty (scattered to numerous); not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; septate, subseptate, or nonseptate; with septa thicker than paper, firm; with the septa eglandular; coriaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril dry; rim-aril; Humularia: H. chevalieri (E.A.J. De Wildeman) P.A. covering less than 1/2 of seed; dark reddish brown. Duvigneaud (DÐF), H. drepanocephalus (E.G. Baker) P.A. Duvigneaud (B), H. spp. (A, C). A, Articles and Seed 2.4Ð4 2.2Ð3.5 1Ð2 mm; not overgrown; not fruits concealed in bracts ( 2.8); B, fruit ( 2.8, angular; asymmetrical; nearly circular, D-shaped, modified from Gillett et al. 1971); C, seeds ( 5.7); quadrangular (nearly), or reniform; compressed; with DÐE, testa ( 50, 1000); F, embryos ( 5).

787 Genus: Cyclocarpa A. Afzelius ex J.G. Baker in groove of raphe; adjacent to hilum; 0.5 mm from hilum; flush; dissimilar color from testa; darker than Phylogenetic Number: 14.16. testa; black (ish); not within corona, halo, or rim. Endosperm thick; 1/2 covering entire embryo; adnate to Tribe: Aeschynomeneae. testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; Subtribe: Aeschynomeninae. not folded; margin entire 180 degrees from base of radicle; similar at apex; completely concealing radicle; Species Studied—Species in Genus: 1 sp.—1 sp. split over radicle; with lobes; with lobes not touching; with the interface division terminating at base of Fruit a loment; 1.5Ð2.2 (diameter of coiled fruit 0.4Ð0.6 cm) radicle; without margins recessed; tan; inner face flat; 0.15Ð0.27 0.03Ð0.04 cm; with deciduous corolla; glabrous around base of radicle. Embryonic axis right with deciduous calyx; with orifice formed by curving of angled; perpendicular to length of seed. Radicle fruit or fruit segments; 1.5-coiled; not plicate; not bulbose; lobe tip straight; deflexed and parallel to twisted; symmetrical; circular; not inflated; flattened; cotyledon width; centered between cotyledons; 1/2 to without beak; rounded at apex; apex oblique with nearly length of cotyledons. Plumule rudimentary; longitudinal axis of fruit (slightly); rounded at base; glabrous. base oblique with longitudinal axis of fruit; with the apex and base uniform in texture; coriaceous; seed Distribution: Tropical Africa, Indochina, Indonesia (Java), chambers externally invisible. Fruit margin not con- and Australia (Queensland). stricted; without sulcus; plain. Fruit wings absent. Fruit substipitate. Fruit with all layers dehiscing; separating Notes: Gillett et al. (1971) have an excellent fruit illustra- from sutures and along loment segment margins. tion. Verdcourt (1974) noted that the species is “very Dehiscence of valves along both sutures (fig. B); local.” Because the fruits are coiled 1.5 times, their passive. Replum visible (when articles separate). lengths, though determined, are of little diagnostic Loment dehiscing along 1 suture. Loment segments value. Their diameters are noted. This unique dehis- (articles) inconspicuous; 1.5Ð2 mm long; widest across cence in the Fabaceae is illustrated in figure B. The seed area; with upper or lower 1 different shape than fruit breaks into articles that in turn dehisce along their middle ones; trapezoid. Epicarp dull; monochrome; margins. dark brown; glabrous; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp glossy; monochrome; dark brown; spongy; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 8Ð11; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 0.7Ð1.2 0.5Ð1 0.25Ð0.5 mm; not overgrown; not angular; asymmetrical; mitaform; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; nearly glossy; colored; monochrome; dark brown, green, or tan; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same Cyclocarpa: C. stellaris A. Afzelius ex J.G. Baker (AÐG). A, color as the rest of the hilum; punctiform; between Articles and fruit ( 7.1); B, article with marginal cotyledon and radicle lobe; flush; not within corona, dehiscence ( 10 and see Notes); C, segment with seed halo, or rim. Lens discernible; less than 0.5 mm in dispersing ( 13); D, seeds ( 11); EÐF, testa ( 50, length; with margins curved; more or less circular; not 1000); G, embryos ( 20).

789 Genus: Discolobium G. Bentham chartaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum fully concealed; concealed by Phylogenetic Number: 14.17. funiculus; without faboid split; larger than punctiform; 1 mm long; with curved outline; elliptic; marginal Tribe: Aeschynomeneae. according to radicle tip or between cotyledon and radicle lobe; flush; not within corona, halo, or rim. Lens Subtribe: Discolobiinae. not discernible. Endosperm absent. Cotyledons smooth; both outer faces convex; both the same thickness; both Species Studied—Species in Genus: 3 spp.—8 spp. more or less of equal length; not folded; margin entire 180 degrees from base of radicle; similar at apex; not Fruit a legume; unilocular; 2.7Ð3.7 0.8Ð1.2 0.07Ð0.09 concealing radicle; entire over radicle or notched at cm; with persistent or deciduous androecial sheath; radicle; without lobes; with the interface division with deciduous corolla; with persistent calyx; with terminating at base of radicle; without margins re- calyx shorter than fruit; without orifice formed by cessed; red; inner face flat; glabrous around base of curving of fruit or fruit segments (because of the upper radicle. Embryonic axis deflexed; oblique to length of and lower coils arising from a twisted suture); 1- to 3- seed. Radicle bulbose; lobe tip curved; oblique to coiled (when single coiled, with one major central coil cotyledons; centered between cotyledons; less than 1/2 with upper and lower much smaller, sterile coils); not length of cotyledons. Plumule rudimentary; glabrous. plicate; twisted (top and bottom coils twisted when compared to center coil); symmetrical; circular; not Distribution: Brazil, Paraguay, and Argentina. inflated; flattened; without beak; rounded at apex (with persistent long, thick style); apex oblique with longitu- Notes: Burkart (1939) monographed the four species in dinal axis of fruit; rounded at base; right angled with Argentina and provided excellent seed and fruit longitudinal axis of fruit; with the apex and base illustrations of Discolobium psoraleifolium. The uniform in texture; coriaceous; seed chambers exter- curiously formed fruit of Discolobium are unusual, nally invisible. Fruit margin not constricted; without though not as unique as Eligmocarpus cynometroides sulcus; embellished; with wing. Fruit wing 1 (per coil); R. Capuron (Caesalpinioideae: Cassieae) (Gunn 1991). 3Ð4 mm wide; continuous wing around fruit; on 1 The Discolobium fruit is composed of three whorls. The suture. Fruit nonstipitate. Fruit indehiscent. Replum upper and lower whorls are smaller in diameter and invisible. Epicarp glossy (nearly); monochrome; brown, sterile, and the middle whorl is larger in diameter and red (dish), or tan; glabrous or pubescent but soon one-seeded (fig. F). This arrangement indicates wind deciduous; with 1 type of pubescence; puberulent; with dispersal of the seeds in the indehiscent fruits. pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; with spines (short and widely spaced on valves of D. pulchellum G. Bentham); not smooth; with elevated features; reticulately veined; tuberculate or not tuberculate; with solid tubercles on each valve; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; spongy; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seed 1; length parallel with fruit length. Funiculus less than 0.5 mm long; thick; straight. Aril absent.

Seed 8 5 0.4 mm; not overgrown; not angular; asymmetrical; reniform (to lunate); flattened; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modi- Discolobium: D. psoraleifolium G. Bentham (BÐF), D. spp. fied by a bloom; colored; monochrome; reddish brown; (A). A, Fruits ( 2.2); B, fruit diagram showing coiling not smooth; with elevated features; wrinkled; ( 2); C, seed ( 7.3); DÐE, testa ( 50, 1000); F, embryos ( 5).

791 Genus: Amicia K.S. Kunth glabrous; smooth; chartaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Phylogenetic Number: 14.19. visible; without faboid split (perhaps too small to see split); punctiform; between cotyledon and radicle lobe; Tribe: Aeschynomeneae. recessed; not within corona, halo, or rim. Lens not discernible. Endosperm absent. Cotyledons smooth; Subtribe: Poiretiinae. both outer faces convex; both the same thickness; both more or less of equal length; not folded; margin entire Species Studied—Species in Genus: 4 spp.—7 spp. 180 degrees from base of radicle; similar at apex; not concealing radicle; split over radicle; with lobes; with Fruit a loment; 1Ð1.5 0.2Ð0.3 0.05Ð0.09 cm; with lobes not touching; with the interface division terminat- deciduous corolla; with persistent or deciduous calyx; ing at base of radicle; without margins recessed; red with calyx shorter than fruit; without orifice formed by (dish); inner face flat; glabrous around base of radicle. curving of fruit or fruit segments; straight; not plicate; Embryonic axis deflexed; parallel to length of seed. not twisted; asymmetrical; when asymmetrical with 1 Radicle linear; deflexed and parallel to cotyledon straight and 1 curved suture; narrowing in several length; centered between cotyledons; 1/2 to nearly places, resembling Desmodium fruit; not inflated; length of cotyledons. Plumule rudimentary; glabrous. compressed; without beak; rounded at apex; apex aligned with longitudinal axis of fruit; short tapered at Distribution: Mexico, Ecuador, Peru, Bolivia, and Argen- base; base aligned with longitudinal axis of fruit; with tina. the apex and base uniform in texture; ligneous; seed chambers externally visible; with the raised seed Notes: Burkart (1939) provided a key to the seven species of chambers not torulose. Fruit margin constricted along Amicia and described the seeds and fruits of the three both margins; with sulcus (margins so thickened to species found in Argentina. Our fruit data were some- create appearance of a sulcus) or without sulcus; plain what limited and readers should not infer from our plate or embellished; with prickles or with wings. Fruit wings that the calyx falls with the mature fruit. The relation of present or absent; 2; 1 mm wide; valvular; on both the calyx to the mature fruit has not been recorded in valves (and compressed to valves). Fruit nonstipitate. the field. The fruits of A. zygomeris A.-P. de Candolle Fruit indehiscent. Replum invisible. Fruit a segment; an are unlike those of other species in the genus. Compare intact article; loment segment. Loment indehiscent. the extreme left article in figure A with the other Loment segments (articles) inconspicuous; 3Ð4 mm articles. long; widest across seed area; with all essentially similar in shape; D-shaped. Epicarp dull; monochrome or multicolored; mottled; brown or gray; with black or brown overlay; with mottling over seed chambers; glabrous; eglandular; without spines; not smooth; with elevated features; veined or not veined; reticulately veined; not tuberculate or tuberculate (faintly); not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð5; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril dry; rim-aril; tan.

Seed 1.8Ð3 1.8Ð3 0.8 mm; not overgrown; not angular; asymmetrical; mitaform; compressed; with visible Amicia: A. medicaginea A.H.R. Grisebach (BÐF), A. spp. radicle and cotyledon lobes; without umbo on seed (A). A, Articles and fruit exserted from calyx ( 3.1); faces. Testa not adhering to endocarp; dull; not modi- B, drawing of two fruits ( 4); C, seed ( 6.8); DÐE, fied by a bloom; colored; monochrome; reddish brown; testa ( 50, 1000); F, embryos ( 15).

793 Genus: Poiretia E.P. Ventenat split the same color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; flush; within rim. Phylogenetic Number: 14.20. Hilum rim color of testa. Lens discernible; less than 0.5 mm in length; with margins curved; circular; not in Tribe: Aeschynomeneae. groove of raphe; adjacent to hilum; 0.4 mm from hilum; mounded; dissimilar color from testa; darker than testa; Subtribe: Poiretiinae. black (ish); not within corona, halo, or rim. Endosperm thin; 1/2 covering entire embryo; adnate to testa. Species Studied—Species in Genus: 4 spp.—6 spp. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; not Fruit a loment; 1.2Ð4 0.3Ð0.6 0.5 cm; with persistent folded; margin entire 180 degrees from base of radicle; or deciduous corolla; with persistent or deciduous similar at apex; partially concealing radicle; split over calyx; with calyx shorter than fruit; without orifice radicle; with lobes; with lobes not touching; with the formed by curving of fruit or fruit segments; straight to interface division terminating at base of radicle; without curved (slightly); not plicate; not twisted; asymmetrical margins recessed; yellow; inner face flat; glabrous or symmetrical; linear to oblong; when asymmetrical around base of radicle. Embryonic axis deflexed; with both sutures parallelly curved; not inflated; oblique to length of seed. Radicle linear; lobe tip flattened; without beak; short tapered or rounded at straight or curved; deflexed and parallel to cotyledon apex; apex aligned with longitudinal axis of fruit; short width; centered between cotyledons; less than 1/2 tapered at base; base aligned with longitudinal axis of length of cotyledons. Plumule rudimentary; glabrous. fruit; with the apex and base uniform in texture; membranous to coriaceous; seed chambers externally Distribution: Mexico, Central America, tropical South visible; with the raised seed chambers not torulose. America. Fruit margin constricted along both margins; without sulcus; plain. Fruit wings absent. Fruit nonstipitate. Notes: Rudd (1972b) summarized the genus and Janke et al. Fruit indehiscent. Replum invisible. Loment indehis- (1988) discussed the pharmacognostic value of P. cent. Loment segments (articles) inconspicuous; 3Ð8 latifolia J.R.T. Vogel and P. tetraphylla. mm long; widest across seed area; with all essentially similar in shape; rectangular. Epicarp semiglossy; monochrome; reddish to greenish brown or tan; glabrous; eglandular; without spines; not smooth; with elevated or recessed features; veined or not veined; reticulately veined; not tuberculate; dotted; pitted; not exfoliating; without cracks. Mesocarp absent. Endocarp glossy; monochrome; tan; smooth; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð8; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened; straight. Aril absent.

Seed 1.5Ð3.5 1Ð2 mm; not overgrown; not angular; asymmetrical; ovate or reniform; flattened; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; not smooth; with recessed features; pitted with small separate pits; coriaceous. Fracture lines Poiretia: P. tetraphylla (J.L.M. Poiret) A.E. Burkart (CÐE), absent. Rim absent. Wings absent. Raphe not visible. P. spp. (AÐB). A, Fruits and article ( 2.2); B, seeds Hilum partially concealed; concealed by funicular ( 6.8); CÐD, testa ( 50, 1000); E, embryos ( 8). remnant; with faboid split; with the lips of the faboid

795 Genus: Zornia J.F. Gmelin Seed 1.5Ð2.3 1Ð1.5 0.5Ð0.8 mm; not overgrown; angular or not angular; asymmetrical; elliptic (reni- Phylogenetic Number: 14.21. form), irregular (reniform), or reniform (with or without beak); compressed; with visible radicle and cotyledon Tribe: Aeschynomeneae. lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; Subtribe: Poiretiinae. monochrome or mottled; with frequent or infrequent mottles; brown (to reddish-brown or purplish-brown) or Species Studied—Species in Genus: 77 spp.—ca. 80 spp. tan; with red overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe Fruit a loment; 1.1Ð1.8 0.1Ð0.35 0.1Ð0.2 cm; with not visible. Hilum visible; with faboid split; with the deciduous corolla; with persistent calyx; with calyx lips of the faboid split the same color as the rest of the longer, equal in length to, or shorter than fruit; without hilum; punctiform; between cotyledon and radicle lobe; orifice formed by curving of fruit or fruit segments; flush; within rim or halo. Hilum halo color lighter than straight to curved (slightly); not plicate; not twisted; testa. Hilum rim color lighter than testa. Lens discern- asymmetrical or symmetrical; moniliform; when ible or not discernible; less than 0.5 mm in length; with asymmetrical with 1 straight and 1 curved suture or margins curved; 2 circular mounds separated by with both sutures parallelly curved; narrowing in groove; not in groove of raphe; confluent with hilum several places, resembling Desmodium fruit; not (or at least to the hilar rim); mounded; dissimilar color inflated; compressed; without beak; rounded at apex; from testa; darker than testa; reddish brown; not within apex aligned with longitudinal axis of fruit; rounded at corona, halo, or rim. Endosperm thick or thin; 1/2 base; base aligned with longitudinal axis of fruit; with covering entire embryo; adnate to testa. Cotyledons the apex and base uniform in texture; coriaceous; seed smooth; both outer faces convex; both the same chambers externally visible; with the raised seed thickness; both more or less of equal length; not folded; chambers not torulose. Fruit margin constricted along margin entire 180 degrees from base of radicle; similar both margins or constricted only on 1 margin; without at apex; partially concealing radicle; split over radicle; sulcus; plain or embellished; with prickles. Fruit wings with lobes; with lobes not touching; with the interface absent. Fruit nonstipitate. Fruit indehiscent. Replum division terminating at base of radicle; without margins invisible. Loment indehiscent. Loment segments recessed; yellow; inner face flat; glabrous around base (articles) inconspicuous; 1.2Ð5 mm long; widest across of radicle. Embryonic axis deflexed; oblique to length seed area; with all essentially similar in shape; circular of seed. Radicle bulbose or linear; lobe tip straight; or D-shaped. Epicarp dull; monochrome (with or deflexed and parallel to cotyledon width or oblique to without different colored reticulum); brown, green, tan, cotyledons; centered between cotyledons; less than 1/2 or yellow; glabrous or pubescent and indurate; with 1 or 1/2 to nearly length of cotyledons. Plumule rudimen- type of pubescence; pilose or villous; with pubescence tary; glabrous. gray or brown; with pubescence uniformly distributed; with simple or complex hairs; with plumose hairs, Distribution: Tropical and warm temperate regions of Old bristle-like hairs, or setae; pliable; with hair bases plain; and New Worlds. antrorse or retrorse; straight, hooked, or coiled at apex; glandular or eglandular; with glandular dots; without Notes: Mohlenbrock (1961) monographed the genus and spines; not smooth; with elevated features; veined or revised his monograph in Mohlenbrock (1962a). not veined; reticulately veined; not tuberculate; Reynolds and Holland (1989) corrected and expanded glandular dotted; not exfoliating; without cracks. the Mohlenbrock monograph for the 17 species that Mesocarp absent. Endocarp dull; monochrome; brown; occur in Australia. spongy; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð15; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; Zornia: Z. glochidiata H.G.L. Reichenbach ex A.-P. de of 1 length only; thick; straight. Aril absent. Candolle (CÐE), Z. spp. (AÐB). A, Fruits and articles ( 2.6); B, seeds ( 5.6); CÐD, testa ( 50, 1000); E, embryos ( 10).

Fruit a loment; 2Ð3 (estimated) 0.5Ð1 (estimated) 0.15 cm; with deciduous corolla; with deciduous calyx; without orifice formed by curving of fruit or fruit segments; straight; not plicate; not twisted; asymmetrical; irregular; when asymmetrical with both sutures unequally curved; not inflated; flattened; with beak; with solid beak the same color and texture as fruit; rounded at apex; apex oblique with longitudinal axis of fruit; rounded at base; base oblique with longitudinal axis of fruit; with the apex and base uniform in texture; coriaceous; seed chambers externally invisible. Fruit margin constricted along both margins; with sulcus (broad shallow depression on 1 face); plain. Fruit wings absent. Fruit substipitate. Fruit indehiscent. Replum invisible. Loment indehiscent. Loment segments (articles) inconspicuous; 1.8Ð2 mm long; widest across seed area; with all essentially similar in shape; oblong. Epicarp dull; monochrome; brown; pubescent and indurate; with 1 type of pubescence; puberulent; with pubescence golden; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; longitudinally veined relative to fruit length; not tuberculate; not exfoliating; without cracks (2Ð3 prominent veins on each valve). Mesocarp present (spongy according to Balfour (1888) who labelled it endocarp). Seeds 1Ð5.

Notes: Gillett (1966b) recognized two species that are rarely collected, A. gracile and A. somalense J.O.D. Hillcoat & J.B. Gillett. Fruits of the latter species are unknown. Unfortunately, Gillett did not discuss the fruit or seed characters of A. gracile of Socotra. Our data presented above are based on two fruit samples and data from the text and illustrations of Balfour (1888). Balfour also noted that A. gracile is a “very beautiful tree.” Our fruit data are limited to external characters as well as the Balfour plate. We have no seed data. Arthrocarpum: A. gracile I.M. Balfour (AÐB). A, Fruits ( 4.4); B, fruit drawing ( 3.5, modified from Polhill and Raven 1981).

799 Genus: Pachecoa P.C. Standley & J.A. Steyermark Rim absent. Raphe from hilum through lens to base of seed and terminating; not bifurcating; darker than testa; Phylogenetic Number: 14.23. brown; flush. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the Tribe: Aeschynomeneae. hilum; punctiform; subapical to radicle tip; flush; within halo. Hilum halo color of or darker than testa. Subtribe: Stylosanthinae. Lens discernible; less than 0.5 mm in length; with margins straight; wedge-shaped; not in groove of raphe; Species Studied—Species in Genus: 1 sp.—1 sp. adjacent to hilum; mounded; dissimilar color from testa; darker than testa; dark brown; not within corona, Fruit a loment; 1.5Ð3 0.4Ð0.5 0.3Ð0.37 cm; with halo, or rim. Endosperm absent. Cotyledons smooth; deciduous corolla; with deciduous calyx; without both outer faces convex; both the same thickness; both orifice formed by curving of fruit or fruit segments; more or less of equal length; not folded; margin entire straight to curved (slightly); not plicate; not twisted; 180 degrees from base of radicle; similar at apex; barely asymmetrical to symmetrical; linear or oblong; partially concealing radicle; split over radicle; with when asymmetrical with both sutures parallelly curved; lobes; with lobes overlapping; with the interface not inflated; terete; without or with beak; straight; with division terminating at base of radicle; without margins solid beak the same color and texture as fruit; long recessed; inner face flat; glabrous around base of tapered at apex; apex aligned with longitudinal axis of radicle. Embryonic axis oblique; oblique to length of fruit; short tapered at base; base aligned with longitudi- seed. Radicle linear; lobe tip straight; oblique to nal axis of fruit; with the apex and base uniform in cotyledons; centered between cotyledons; less than 1/2 texture; ligneous; seed chambers externally invisible. length of cotyledons. Plumule moderately developed; Fruit margin not constricted; without sulcus; plain. Fruit glabrous. wings absent. Fruit nonstipitate. Fruit indehiscent. Replum invisible. Loment indehiscent. Loment seg- Distribution: Mexico and Guatemala; introduced in Venezu- ments (articles) conspicuous; 2.5Ð10 mm long; widest ela. across seed area; with upper or lower 1 different shape than middle ones; oblong or triangular. Epicarp dull; Notes: Norman and Gunn (1985) monographed Pachecoa. monochrome; dirty brown; glabrate or pubescent but soon deciduous; with 1 type of pubescence; villous; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; without spines; not smooth; with elevated features; longitudinally veined relative to fruit length (8Ð9 veins) or reticulately veined (interstices); not tuberculate; not exfoliating; without cracks. Mesocarp thick; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; ligneous (sub). Endocarp dull; monochrome; tan; smooth; septate; with septa thicker than paper, firm; with the septa eglandular; ligneous (sub); not exfoliating; remaining fused to epicarp; entire. Seeds 2Ð4; length oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril dry; rim-aril; tan.

Seed 3Ð3.5 2.3Ð2.5 0.9Ð1 mm; not overgrown; not angular; asymmetrical; rhombic; compressed; without Pachecoa: P. prismatica (M. Sessé y Lacastra & J.M. visible radicle and cotyledon lobes; without umbo on Mociño) P.C. Standley & B.G. Schubert (AÐB); P. seed faces. Testa not adhering to endocarp; glossy; not venezuelensis A. Burkart (CÐE). A, Articles and fruits modified by a bloom; colored; monochrome; tan; ( 2); B, seeds ( 7.3); CÐD, testa ( 50, 1000); E, glabrous; smooth; coriaceous. Fracture lines absent. embryos ( 6).

801 Genus: Chapmannia J. Torrey & A. Gray on seed faces. Testa not adhering to endocarp; nearly glossy; not modified by a bloom; colored; mono- Phylogenetic Number: 14.24. chrome; yellow; glabrous; not smooth; with elevated features; wrinkled; chartaceous. Fracture lines absent. Tribe: Aeschynomeneae. Rim absent. Wings absent. Raphe from hilum to near base of seed and terminating; not bifurcating; color of Subtribe: Stylosanthinae. testa; raised. Hilum fully concealed; concealed by funicular remnant; without faboid split; punctiform; Species Studied—Species in Genus: 1 sp.—1 sp. between cotyledon and radicle lobe; flush; not within corona, halo, or rim. Lens not discernible. Endosperm Fruit a loment; 1Ð3 0.25Ð0.3 0.25Ð0.3 cm; with absent. Cotyledons smooth; both outer faces convex; deciduous corolla; with deciduous calyx; without both the same thickness; both more or less of equal orifice formed by curving of fruit or fruit segments; length; not folded; margin entire 180 degrees from base straight; not plicate; not twisted; asymmetrical; more or of radicle; similar at apex; completely concealing less linear; when asymmetrical with 1 straight and 1 radicle (nearly); split over radicle; with lobes; with curved suture or both sutures parallelly curved; widest lobes touching (auriculate); with the interface division near middle or D-shaped (more or less); not inflated; terminating at base of radicle; tan; inner face flat; terete; without or with beak; straight; with solid beak glabrous around base of radicle. Embryonic axis the same color and texture as fruit; long tapered at apex; oblique; oblique to length of seed. Radicle nearly apex aligned with longitudinal axis of fruit; rounded at bulbose; lobe tip curved; oblique to cotyledons; base; base aligned with longitudinal axis of fruit; with centered between cotyledons; less than 1/2 length of the apex and base uniform in texture; coriaceous; seed cotyledons. Plumule moderately developed; glabrous. chambers externally visible; with the raised seed chambers not torulose. Fruit margin constricted along Distribution: United States (Florida). both margins; without sulcus; plain. Fruit wings absent. Fruit nonstipitate. Fruit indehiscent. Replum invisible. Notes: Gunn et al. (1980) monographed the genus. Loment indehiscent. Loment segments (articles) conspicuous; 5Ð8 mm long; widest across seed area; with upper or lower 1 different shape than middle ones; more or less oblong. Epicarp dull; monochrome; pubescent but soon deciduous; with 1 type of pubescence; villous; with pubescence red (dish-brown and turning brown at maturity) or brown; with pubescence uniformly distributed; with simple hairs and with glandular hairs; pliable; with hair bases plain; glandular; with glandular hairs (reddish, turning brown on maturation); not smooth; with elevated features; longitudinally veined relative to fruit length; not tuberculate; not exfoliating; without cracks. Mesocarp thick; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; white (ish); smooth; septate; with septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð4; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 3.5Ð4 2Ð2.5 2 mm; not overgrown; not angular; Chapmannia: C. floridana J. Torrey & A. Gray (AÐE). A, asymmetrical; oblong (with radicle end tapered); terete; Articles and incomplete fruits ( 2.1); B, seeds ( 6.5); with visible radicle and cotyledon lobes; without umbo CÐD, testa ( 50, 1000); E, embryos ( 10).

803 Genus: Stylosanthes O.P. Swartz exfoliating; remaining fused to epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlap- Phylogenetic Number: 14.25. ping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent. Tribe: Aeschynomeneae. Seed 1.9Ð3 1.4Ð2.5 0.6Ð1.2 mm; not overgrown; not Subtribe: Stylosanthinae. angular; asymmetrical; ovate or reniform (with prominent radicle lobe (beaklike)); compressed; with visible Species Studied—Species in Genus: 25 spp.—ca. 25 spp. radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; glossy; not Fruit a loment; 0.4Ð0.9 0.08Ð0.25 0.01Ð0.02 cm; with modified by a bloom; colored; monochrome; black, deciduous corolla; with persistent or deciduous calyx; brown (to reddish-brown), or tan; smooth; chartaceous. with calyx shorter than fruit; without orifice formed by Fracture lines absent. Rim absent. Wings absent. Raphe curving of fruit or fruit segments; straight; not plicate; not visible. Hilum visible; with faboid split; with the not twisted; asymmetrical; rectangular (with or without lips of the faboid split the same color as the rest of the apical beak); when asymmetrical with 1 straight and 1 hilum; punctiform; between cotyledon and radicle lobe; curved suture or both sutures parallelly curved; widest flush; not within corona, halo, or rim. Lens discernible; near middle or D-shaped; not inflated; compressed; less than 0.5 mm in length; with margins curved; with beak; straight, declined, or coiled; with solid beak circular; not in groove of raphe; adjacent to hilum; 0.5 the same color and texture as fruit; long tapered to mm from hilum; mounded; dissimilar color from testa; tapered to short tapered at apex; apex aligned with darker than testa; black; not within corona, halo, or rim. longitudinal axis of fruit; short tapered or truncate at Endosperm present or absent; thin; 1/2 covering entire base; base aligned with longitudinal axis of fruit; with embryo; adnate to testa. Cotyledons smooth; both outer the apex and base uniform in texture; chartaceous to faces convex; both the same thickness; both more or coriaceous or fragile, thinner than chartaceous like less of equal length; not folded; margin not entire 180 Trifolium (23.07); seed chambers externally visible; degrees from base of radicle; similar at apex; partially with the raised seed chambers not torulose. Fruit concealing radicle; split over radicle; with lobes; with margin constricted or not constricted; constricted along lobes not touching; with the interface division terminat- both margins or constricted on 1 margin and slightly ing at base of radicle; without margins recessed; constricted on the other margin; without sulcus; plain. yellow; inner face flat; glabrous around base of radicle. Fruit wings absent. Fruit nonstipitate. Fruit indehiscent. Embryonic axis deflexed; oblique to length of seed. Replum invisible. Loment indehiscent. Loment seg- Radicle bulbose; lobe tip curved; deflexed and parallel ments (articles) inconspicuous; 2Ð4.5 mm long; widest to cotyledon width; centered between cotyledons; less across seed area; with upper 1 different shape than than 1/2 length of cotyledons. Plumule rudimentary; middle one, lower 1 different shape than middle one, or glabrous. upper 1 different shape than lower 1; D-shaped. Epicarp dull; monochrome; with surface texture uniform or not Distribution: United States, West Indies, Mexico, Central uniform, with patches of different texture not restricted America, South America (to northern Argentina and to the base and apex; glabrous, pubescent and indurate, Galapagos Islands), central and southern Africa, or pubescent but soon deciduous; with 1 type of Madagascar, southern India, Sri Lanka, and adventive pubescence; pilose, puberulent, or villous; with in Indonesia to Australia. pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; Notes: Mohlenbrock (1957) revised Stylosanthes and eglandular; without spines; not smooth; with elevated amended his revision (Mohlenbrock 1963). Kirkbride features; longitudinally veined relative to fruit length or and Kirkbride (1987) established the correct names for reticulately veined (with 2 prominent longitudinal the two sections in the genus, and our studied species veins); not tuberculate or tuberculate (minutely in S. were obtained nearly equally from both sections. guianensis); not exfoliating; without cracks. Mesocarp Mohlenbrock (1963) provided a key and fruit illustra- absent. Endocarp dull; monochrome; tan; smooth; tions of the 30 species that he recognized. Mohlenbrock septate; with septa thicker than paper, firm; with the (1957) noted that the lower (of two) articles is usually septa eglandular; coriaceous to chartaceous; not densely pilose to glabrescent and aborted to fertile while the upper article is glabrous or puberulent to

804 sericeous to minutely tuberculate. The relative beak- upper article lengths are diagnostic. T’Mannetje (1984) reported on the species of this genus and has excellent fruit drawings. Burkart (1952) and Sousa Costa and Ferreira (1984) contain excellent fruit and seed drawings. Reis and Martins (1989a,b) presented interesting data on seed germination of apical and basal section seeds and on the distributional potential of the upper one-half of the legume versus the lower one-half of the legume. Some species of Stylosanthes have agronomic potential, especially in Australia and South America (Stace and Edye 1984).

805 Stylosanthes: S. guianensis (J.B.C.F. Aublet) O.P. Swartz (DÐF), S. spp. (AÐC). A, Articles ( 4.7); B, fruit beaks (coiled, hooked, and straight) ( 10); C, seeds ( 6.8); DÐE, testa ( 50, 1000); F, embryos ( 8).

807 Genus: Arachis C. Linnaeus absent or present (according to Kubicek (1970) and see Notes); trace; 1/2 covering entire embryo; adnate to Phylogenetic Number: 14.26. testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; Tribe: Aeschynomeneae. not folded; margin entire 180 degrees from base of radicle; similar at apex; partially concealing radicle; Subtribe: Stylosanthinae. split over radicle; with lobes; with lobes not touching; with the interface division terminating at base of Species Studied—Species in Genus: 18 spp.—71 spp. radicle; without margins recessed; tan; inner face flat; glabrous around base of radicle. Embryonic axis Fruit a legume; unilocular; 0.8Ð6 0.5Ð2.1 0.5Ð2.1 cm; straight; parallel to length of seed. Radicle bulbose; with deciduous corolla; with deciduous calyx; without lobe tip straight or curved; straight with embryonic orifice formed by curving of fruit or fruit segments; axis; centered between cotyledons; less than 1/2 length straight; not plicate; not twisted; asymmetrical or of cotyledons. Plumule well developed; glabrous. symmetrical; oblong, ovate, or irregular; when asymmetrical with both sutures unequally curved; not Distribution: Eastern South America; introduced and inflated; terete; without beak; rounded at apex; apex cultivated elsewhere. oblique or right-angled with longitudinal axis of fruit; rounded at base; base oblique with longitudinal axis of Notes: Hoehne (1940) and Hermann (1954) monographed fruit; with the apex and base uniform in texture; the genus, and Resslar (1980) reviewed the nomencla- coriaceous; seed chambers externally visible or ture. Rudd (1981a) used the same number of recog- invisible; with the raised seed chambers not torulose. nized species as Resslar, and both noted that possibly Fruit margin constricted or not constricted; constricted 40 (Rudd) to 70 (Resslar) species were undescribed. along both margins; without sulcus; plain. Fruit Wynne and Halward (1989) recognized 33 species, nonstipitate. Fruit indehiscent. Replum invisible. including several nomum nuda species names. Epicarp dull; monochrome; tan; glabrous; eglandular; Krapovickas and Gregory (1994) monographed Arachis without spines; not smooth; with elevated features; and recognized 69 species in the genus. Several new reticulately veined; not tuberculate; not exfoliating; species of Arachis have been discovered since publica- without cracks. Mesocarp thick; surface uniformly tion of Krapovickas and Gregory’s monograph (José veined; 1-layered; without balsamic vesicles; without F.M. Valls, personal communication, 1994). Kubicek fibers; firm-walled open empty cells; coriaceous. (1970) stated that “contrary to reports by other investi- Endocarp dull; monochrome; white; cobwebby or gators the endosperm appeared to be present as a single scurfy; nonseptate; chartaceous; not exfoliating; layer which covers the cotyledons in mature seeds.” remaining fused to epicarp; entire. Seeds 1Ð3; length Gregory and Gregory (1979) studied interspecific parallel with fruit length; touching; in 1 series. Funicu- hybrids. Bentham (1841) compared the structures and lus less than 0.5 mm long; of 1 length only; thick; affinities of Arachis and Voandzeia (now Vigna 10.65). straight. Aril absent. Charles F. Simpson, USDA/ARS, made a major contribution of seeds and fruits to the U.S. National Seed 7Ð25 4Ð15 4Ð15 mm; overgrown, 1 seed filling Seed Herbarium (BARC). entire fruit cavity or not overgrown; angular or not angular; asymmetrical; elliptic, oblong, or ovate; terete; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or bichrome (reddish-brown and white); reddish brown or white; glabrous; smooth or not smooth; with elevated features; veined; chartaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum fully concealed; concealed by funicular remnant; without faboid split; larger than punctiform; 2 mm long; with curved outline; Arachis: A. hypogaea C. Linnaeus (CÐE), A. spp. (AÐB). A, elliptic; subapical to radicle tip; flush; not within Fruits and valve ( 1.1); B, seeds ( 2); CÐD, testa corona, halo, or rim. Lens not discernible. Endosperm ( 50, 1000); E, embryos ( 2.5).

809 Adesmieae (15.01) to endocarp; glossy; not modified by a bloom; colored; monochrome, mottled, or streaked; with frequent Genus: Adesmia A.-P. de Candolle mottles; with frequent streaks; black, brown (to reddish), or tan; with brown overlay; glabrous; smooth; Phylogenetic Number: 15.01. osseous to coriaceous. Fracture lines absent. Rim absent. Raphe not visible. Hilum fully concealed or Tribe: Adesmieae. partially concealed; concealed by aril, aril remnant, radicle lobe, or wing; with faboid split; with the lips of Species Studied—Species in Genus: 35 spp.—ca. 230 spp. the faboid split lighter colored than the rest of the hilum and therefore conspicuous; punctiform; between Fruit a loment; 1.2Ð3 ϫ 2.5Ð7 ϫ 0.05Ð0.2 cm; with decidu- cotyledon and radicle lobe; recessed; within rim. Hilum ous corolla; with deciduous calyx; straight or curved (or rim color darker than testa. Lens discernible; equal to or slightly curved); not plicate; not twisted; asymmetrical; greater than 0.5 mm in length; 1Ð2 mm long; with when asymmetrical with 1 straight and 1 curved suture; margins curved; circular (as large as hilum); not in narrowing in several places, resembling Desmodium groove of raphe; adjacent to hilum; mounded; dissimi- fruit; not inflated; flattened or compressed; without lar color from testa; darker than testa; black; not within beak; rounded at apex; apex aligned with longitudinal corona, halo, or rim. Endosperm thin; 1/2 covering axis of fruit; rounded at base; base aligned with entire embryo; adnate to testa. Cotyledons smooth; longitudinal axis of fruit; with the apex and base outer face of 1 cotyledon flat and other cotyledon uniform in texture; membranous; seed chambers convex; 1 thicker than the other; both more or less of externally visible; with the raised seed chambers not equal length; with both folded or not folded; not torulose. Fruit margin constricted only on 1 margin; sufficiently folded for inner face to touch itself; margin without sulcus; embellished; with prickles. Fruit wings entire 180 degrees from base of radicle; similar at apex; absent. Fruit substipitate. Fruit with all layers dehiscing not concealing radicle (nearly); split over radicle; with (occasionally: Burkart (1967, p. 491)) or indehiscent; lobes; with lobes not touching; with the interface splitting along sutures. Dehiscence of valves passive. division terminating at base of radicle; without margins Replum invisible. Loment segments (articles) incon- recessed; white or yellow; inner face flat; glabrous spicuous; 4Ð12 mm long; widest across seed area; with around base of radicle. Embryonic axis deflexed; all essentially similar in shape; triangular or D-shaped. parallel to length of seed. Radicle linear; lobe tip Epicarp dull; monochrome; black, brown (to reddish straight; deflexed and parallel to cotyledon length; brown), or tan; glabrous or pubescent but soon decidu- centered between cotyledons; 1/2 to nearly length of ous; with 1 type of pubescence; pilose, tomentose, or cotyledons, equaling length of cotyledons, or exceeding villous; with pubescence golden, gray-brown, red, gray, length of cotyledons. Plumule rudimentary; glabrous. or tan; with simple or complex hairs; with plumose or bristle-like hairs (up to 1 cm long); stiff or pliable; with Distribution: Montane and temperate South America (Peru hair bases plain; straight; straight at apex; eglandular; and southern Brazil to Tierra del Fuego). without spines; not smooth; with elevated features; reticulately veined; not tuberculate; glandular dotted; Notes: Polhill (1981g) evaluated the phylogenetic position not exfoliating; without cracks. Mesocarp absent. of Adesmieae. For additional seed and fruit drawings, Endocarp dull; monochrome; tan; smooth; nonseptate; see Burkart (1966, 1967), the latter a synopsis of the chartaceous; not exfoliating; remaining fused to genus. In his synopsis, Burkart compared the seeds of epicarp. Seeds 1Ð12; length parallel with fruit length; A. bicolor (J.L.M. Poiret) A.-P. de Candolle to those of neither overlapping nor touching; in 1 series. Funiculus Trifolium repens C. Linnaeus (Trifolieae, 21.06). less than 0.5 mm long; of 1 length only. Aril dry; rim- Ulibarri (1987) monographed the series Microphyllae aril; brown. and included 33 species. Miotto and Leitão filho (1993) treated the Brazilian species of Adesmia. Most of our Seed 1.5Ð3.5 ϫ 1.5Ð3.5 ϫ 1Ð2.4 mm; overgrown, 1 seed fruit samples were composed of individual indehiscent filling entire fruit cavity; not angular (though at first articles, and our data for entire fruits were derived from glance appearing to be angular) or angular; asymmetri- the two papers of Burkart. cal; mitaform (or nearly so); mounded on 1 side and straight on other side; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering

810 811 Adesmia: A. aromatica A. Burkart (H), A. capitellata A. Burkart (G), A. incana E. de Vogel (CÐE), A. lihuelensis A. Burkart (F), A. spp. (AÐB). A, Fruits (ϫ 1.9); B, seeds (ϫ 4.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10); F, bristlelike fruit hair (ϫ 16) redrawn from Burkart (1966); G, simple fruit hairs (ϫ 30) redrawn from Burkart (1966); H, plumose fruit hair (ϫ 15) redrawn from Burkart (1966).

813 Galegeae (16.01Ð16.22) frequent streaks; reddish brown or tan (reddish); with black overlay; glabrous; not smooth; with elevated Genus: Clianthus D.C. Solander ex J. Lindley features; reticulate or wrinkled; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not Phylogenetic Number: 16.01. visible. Hilum visible or fully concealed; concealed by radicle lobe; with faboid split; with the lips of the Tribe: Galegeae. faboid split the same color as the rest of the hilum; punctiform; marginal according to radicle tip; flush; not Subtribe: Coluteinae. within corona, halo, or rim. Lens discernible or not discernible; less than 0.5 mm or equal to or greater than Species Studied—Species in Genus: 1 sp.—1 sp. 0.5 mm in length; up to 0.5 mm long; with margins straight or curved; linear or circular; not in groove of Fruit a legume; unilocular; up to 8 ϫ 0.5Ð1 (assumed) ϫ raphe; adjacent to hilum; up to 0.5 mm from hilum; 0.5Ð1 (assumed) cm; with persistent calyx; with calyx mounded; same color as or dissimilar color from testa; shorter than fruit; without orifice formed by curving of darker than testa; black; not within corona, halo, or rim. fruit or fruit segments; slightly curved; not plicate; not Endosperm thick; covering entire embryo; adnate to twisted; asymmetrical; oblong; when asymmetrical with testa or embryo. Cotyledons smooth; both outer faces both sutures nearly straight; somewhat inflated; terete; convex; both the same thickness; both more or less of with beak; straight; with solid beak the same color and equal length; not folded; margin entire 180 degrees texture as fruit; long tapered at apex; apex aligned with from base of radicle; similar at apex; not concealing longitudinal axis of fruit; long tapered at base; base radicle; entire over radicle; without lobes; with the aligned or oblique (slightly) with longitudinal axis of interface division terminating at base of radicle; without fruit; with the apex and base uniform in texture; margins recessed; yellow or tan (reddish); inner face coriaceous; seed chambers externally invisible. Fruit flat; glabrous around base of radicle. Embryonic axis margin not constricted; without sulcus; plain. Fruit deflexed; oblique to length of seed; without a joint wings absent. Fruit stipitate; with the stipe up to 10 mm evident between the radicle and the cotyledons. Radicle long. Fruit with all layers dehiscing; splitting along differentiated from cotyledon; bulbose; lobe tip straight; suture. Dehiscence of valves along 1 suture; apical and deflexed and parallel to cotyledon length; centered or down; passive. Replum visible. Epicarp dull; mono- not centered between cotyledons (radicle outside 1 chrome; dirty dark brown; glabrous; eglandular; cotyledon and inside other, therefore junctions for each without spines; not smooth; with elevated features; cotyledon different); less than 1/2 or 1/2 to nearly transversely veined relative to fruit length and reticu- length of cotyledons. Plumule rudimentary; glabrous. lately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; Distribution: Northeastern New Zealand. without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; brown; hairy; Notes: Traditionally this tribe has been called Galegeae. nonseptate; chartaceous; not exfoliating; remaining Reveal (1997) reported that the name Astragaleae was fused to mesocarp and epicarp; entire. Seeds many; published before the name Galegeae. In accordance length parallel with fruit length; neither overlapping nor with the International Code of Botanical Nomenclature touching; in 1 series. Funiculus measured; 3Ð3.5 mm (Greuter et al. 1994), the oldest name for a taxon must long; of 1 length only; filiform; more or less straight. be used, so Reveal suggested that this tribe should be Aril faintly present; dry; rim-aril; white. called Astragaleae. In 1999, however, Reveal (1999) reversed himself, so this tribe remains the Galegeae. Seed 3.5Ð4 ϫ 3Ð3.5 ϫ 1Ð1.3 mm; not overgrown; not Welsh (1960) reported on the Galegeae of the North- angular; asymmetrical; reniform; compressed; with Central United States. Sanderson and Liston (1995) surface smooth; with visible radicle and cotyledon carried out cladistic analyses of Galegeae genera using lobes; without external groove between radicle and molecular data. They concluded that Galegeae is cotyledon lobes; without hilar sinus; without umbo on paraphyletic, having given rise to the tribes Cicereae seed faces. Testa not adhering or partially adhering to (20), Hedysareae (18), Trifolieae (21), and Fabeae (19), endocarp (seeds often with tuffs of endocarp hairs); and therefore requiring a re-evaluation of the circum- dull; not modified by a bloom; colored; monochrome or scription of Galegeae. Heenan (1995, 1998c), using mottled and streaked; with frequent mottles; with

814 unpublished nuclear ribosomal DNA ITS data, concluded that “Carmichaelia (17.05) is nested within [the] ‘Astragalean clade’ of Galegeae” and is the sister group of Clianthus. He therefore supported the proposal of Sanderson and Wojciechowski (1996) that Carmichaelieae should not be recognized at the tribal level, but instead should be included in Galegeae. Because the seeds of Clianthus puniceus are so unlike those of C. formsus (G. Don) N.C. Ford & J.W. Vickery, we are pleased with the transfer of the latter species to Swainsona (16.02). Polhill (1981h) noted that C. puniceus “is virtually extinct.” We were unable to study an entire fruit of this genus.

815 Clianthus: C. puniceus (G. Don) D.C. Solander ex J. Lindley (AÐE). A, Broken fruit (ϫ 2.2); B, seeds (ϫ 10); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

817 Genus: Swainsona R.A. Salisbury with external groove between radicle and cotyledon lobes same color as testa; without or with deep hilar Phylogenetic Number: 16.02. sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; Tribe: Galegeae. monochrome; reddish brown; glabrous; not smooth or smooth; with elevated features; reticulate; coriaceous. Subtribe: Coluteinae. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum partially or fully concealed; con- Species Studied—Species in Genus: 15 spp.—85 spp. cealed by radicle lobe; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; Fruit a legume; unilocular or bilocular (see also Astragalus punctiform; between cotyledon and radicle lobe; flush; (16.15)); at least 1Ð5 ϫ 0.3Ð2.5 ϫ 0.1Ð0.35 cm; with within halo or not within corona, halo, or rim. Hilum persistent calyx; with calyx shorter than fruit; without halo color lighter than testa. Lens discernible; less than orifice formed by curving of fruit or fruit segments; 0.5 mm or equal to or greater than 0.5 mm in length; straight or curved (slightly); not plicate; not twisted; 0.5 mm long; with margins straight; linear; not in asymmetrical or symmetrical; linear, oblong, elliptic, or groove of raphe; confluent with hilum; mounded; falcate; when asymmetrical with both sutures parallelly dissimilar color from testa; darker than testa; black; not curved or nearly straight; not inflated or inflated; within corona, halo, or rim or within halo. Lens halo flattened; with beak; straight or declined; with solid color lighter than testa. Endosperm thick; covering beak the same color and texture as fruit; short tapered entire embryo; adnate to embryo. Cotyledons smooth; or rounded at apex; apex aligned, oblique, or right- outer face of 1 cotyledon flat and other cotyledon angled with longitudinal axis of fruit; short tapered at convex; both the same thickness; both more or less of base; base right angled or oblique with longitudinal axis equal length; not folded; margin entire 180 degrees of fruit; with the apex and base uniform in texture; from base of radicle; similar at apex; not concealing membranous, coriaceous, or ligneous; seed chambers radicle; entire over radicle; without lobes; with the externally invisible. Fruit margin not constricted; interface division terminating at base of radicle; without without sulcus; plain. Fruit wings absent. Fruit stipitate, margins recessed; tan; inner face flat; glabrous around substipitate, or nonstipitate; with the stipe at least up to base of radicle. Embryonic axis deflexed; oblique to 10 mm long. Fruit indehiscent or with all layers length of seed; without a joint evident between the dehiscing (to tardily dehiscent); splitting along suture. radicle and the cotyledons. Radicle bulbose; lobe tip Dehiscence of valves along 1 suture; medial and up and straight or hooked; deflexed and parallel to cotyledon down; passive. Replum invisible. Epicarp dull; mono- length; centered between cotyledons; 1/2 to nearly chrome; brown; glabrous or pubescent and indurate; length of cotyledons. Plumule rudimentary; glabrous. with 1 type of pubescence; villous; with pubescence gray; with pubescence uniformly distributed; with Distribution: Australia (most species) and southern New simple hairs; pliable; with hair bases plain; eglandular; Zealand (S. novae-zelandiae J.D. Hooker). without spines; not smooth; with elevated features; reticulately veined and transversely veined relative to Notes: Thompson (1993) monographed Swainsona and fruit length; not tuberculate or tuberculate (bases of recognized 85 species, the count that we used. Although hairs); wrinkled; not exfoliating; without cracks. fruits are seldom collected for most species, the fruits Mesocarp absent. Endocarp dull; monochrome; tan; have great phylogenetic and diagnostic significance for smooth; nonseptate; chartaceous; not exfoliating; the species. Thompson regarded the primitive and most remaining fused to epicarp; entire. Seeds few to unmodified fruits to be greatly inflated with seeds that numerous; length parallel (assumed) with fruit length; are “erratic in their maturing into viable seeds.” She assumed overlapping; in 2 or more series. Funiculus also noted that “seeds have proved difficult to study, measured; up to at least 2 mm long; of 1 length only; few being available in the mature state and those filiform; S-curved. Aril dry; rim-aril; white. seeming to be somewhat inconsistent in size, shape, surface sculpturing and colour.” Species of Swainsona, Seed 3.5Ð4 ϫ 3Ð3.5 ϫ 1 mm; not overgrown; not angular; containing swainsonine, are a well known cause of asymmetrical; reniform; compressed; with surface livestock poisoning. Swainsona in Australia has many smooth; with visible radicle and cotyledon lobes; with parallels with Astragalus (16.15) in North America. external groove between radicle and cotyledon lobes;

818 Thompson transferred Clianthus (16.01) formosus (G. Don) N.C. Ford & J.W. Vickery to Swainsona. Heenan (1998c) carried out phylogenetic analyses of the Carmichaelia (17.05) complex, Clianthus, and 13 species of Swainsona, including S. novae-zelandiae J.D. Hooker. He concluded that “Swainsona is polyphyletic if S. novae-zelandiae is included,” and recommended the segregation of S. novae-zelandiae as a monotypic genus. The genus Montigena P.B. Heenan (Heenan 1998b) was established with the single New Zealand species M. novae-zelandiae (J.D. Hooker) P.B. Heenan. We have chosen to include M. novae-zelandiae in Swainsona pending an expression of a consensus opinion on the status of Montigena by the taxonomic community.

819 Swainsona: S. maccullochiana F.H. Mueller (CÐE), S. spp. (AÐB). A, Fruits and valve (ϫ 1.3); B, seeds (ϫ 6.4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

821 Genus: Sutherlandia R. Brown than 0.5 mm in length; 0.5 mm long; with margins curved; more or less circular; not in groove of raphe; Phylogenetic Number: 16.03. confluent with hilum; barely mounded; same color as testa; not within corona, halo, or rim. Endosperm thick; Tribe: Galegeae. covering entire embryo; adnate to embryo. Cotyledons smooth; both outer faces convex; both the same Subtribe: Coluteinae. thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; similar Species Studied—Species in Genus: 5 spp.—6 spp. at apex; not concealing radicle; entire over radicle; without lobes; with the interface division terminating at Fruit a legume; unilocular; 4Ð6 ϫ 2Ð3.5 cm; with persistent base of radicle; without margins recessed; tan; inner calyx; with calyx shorter than fruit; without orifice face flat; glabrous around base of radicle. Embryonic formed by curving of fruit or fruit segments; straight; axis deflexed; oblique to length of seed; without a joint not plicate; not twisted; asymmetrical; oblong or evident between the radicle and the cotyledons. Radicle circular; when asymmetrical with both sutures bulbose; lobe tip straight; deflexed and parallel to parallelly curved; inflated; without or with beak; cotyledon length; centered between cotyledons; 1/2 to hooked; with solid beak the same color and texture as nearly length of cotyledons. Plumule rudimentary; fruit; short tapered at apex; apex oblique or right-angled glabrous. with longitudinal axis of fruit; tapered at base; base oblique or right angled with longitudinal axis of fruit; Distribution: South Africa, extending to Nambia and with the apex and base uniform in texture; chartaceous; Botswana; naturalized on Mexican, Bolivian, and seed chambers externally invisible. Fruit margin not Argentian plateaux. constricted; without sulcus; plain. Fruit wings absent. Fruit substipitate. Fruit indehiscent. Replum invisible. Notes: Schrire and Andrews (1992) briefly reviewed the Epicarp dull or glossy; monochrome (and transparent); systematics of the cultivated species, and their species tan to purplish tan; glabrous; eglandular; without number is used. The fruit wall of Sutherlandia is very spines; not smooth; with elevated features; reticulately thin and transparent. Using a dissecting microscope, veined; not tuberculate; wrinkled; not exfoliating; only a single layer could be discerned, and it was without cracks. Mesocarp absent. Endocarp absent. presumed to be the epicarp. It is an unusual condition to Seeds 11Ð17; length transverse to fruit length; neither have both the mesocarp and endocarp absent. overlapping nor touching; in 2 or more series. Funicu- lus measured; up to 3 mm long; of 1 length only; nearly thick; straight or curved. Aril dry; rim-aril; tan.

Seed 2Ð3.7 ϫ 1.7Ð3 ϫ 0.7Ð1.3 mm; not overgrown; not angular; asymmetrical; mitaform or reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; with shallow hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; black or brown (blackish to reddish); glabrous; not smooth; with recessed features; pitted with small separate pits; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; recessed; not within corona, halo, or rim or within halo. Sutherlandia: S. frutescens (C. Linnaeus) R. Brown (BÐE), Hilum halo color lighter than testa. Lens discernible or S. spp. (A). A, Fruits (ϫ 1.2); B, seeds (ϫ 6.6); CÐD, not discernible; less than 0.5 mm or equal to or greater testa (ϫ 50, ϫ 1000); E, embryos (ϫ 9).

823 Genus: Lessertia A.-P. de Candolle Seed 2.5Ð4.5 ϫ 2Ð3.5 ϫ 1Ð1.3 mm; not overgrown; not angular; asymmetrical; mitaform or reniform; com- Phylogenetic Number: 16.04. pressed; with surface smooth; with visible radicle and cotyledon lobes; with or without external groove Tribe: Galegeae. between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color Subtribe: Coluteinae. as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modi- Species Studied—Species in Genus: 10 spp.—ca. 50 spp. fied by a bloom; colored; monochrome; reddish to greenish brown or black (reddish); glabrous; smooth or Fruit a legume; unilocular; 0.9Ð4 ϫ 0.5Ð2 ϫ 0.05Ð0.12 cm; not smooth; with elevated features; wrinkled; coria- with persistent calyx; with calyx shorter than fruit; ceous. Fracture lines absent. Rim absent. Wings absent. without orifice formed by curving of fruit or fruit Raphe not visible or visible; from hilum to lens; not segments; straight; not plicate; not twisted; symmetrical bifurcating; color of or darker than testa; reddish or asymmetrical; oblong, ovate, elliptic, linear brown; flush. Hilum visible; with faboid split; with the (broadly), or circular (somewhat); when asymmetrical lips of the faboid split the same color as the rest of the with 1 straight and 1 curved suture, both sutures hilum; punctiform; between cotyledon and radicle lobe; unequally curved, or both sutures parallelly curved; recessed; within halo. Hilum halo color lighter than widest near middle or D-shaped; not inflated or testa. Lens discernible or not discernible; less than 0.5 inflated; compressed, flattened, or terete; with beak; mm in length; with margins curved; circular; not in straight; with solid beak the same color and texture as groove of raphe; adjacent to hilum; 0.3 mm from hilum; fruit; short tapered or rounded at apex; apex aligned or mounded; same or similar color as testa; darker than oblique with longitudinal axis of fruit; short tapered or testa; reddish brown; not within corona, halo, or rim. rounded at base; base aligned with longitudinal axis of Endosperm thick; covering entire embryo; adnate to fruit; with the apex and base uniform in texture; embryo. Cotyledons smooth; both outer faces convex; chartaceous; seed chambers externally visible; with the both the same thickness; both more or less of equal raised seed chambers torulose. Fruit margin not length; not folded; margin entire 180 degrees from base constricted; without sulcus; plain. Fruit wings absent of radicle; similar at apex; partially concealing radicle; (though whole fruit winglike). Fruit substipitate. Fruit entire over radicle; without lobes; with the interface indehiscent or with all layers dehiscing (gaping at division terminating at base of radicle; without margins length at apex or subdehiscent); splitting along suture recessed; yellow or white; inner face flat; glabrous (at least apex). Dehiscence of valves along 1 suture; around base of radicle. Embryonic axis deflexed; apical and down (apex only) or medial and up and oblique to length of seed; with a joint evident between down (when gaping); passive. Replum invisible. the radicle and the cotyledons. Radicle bulbose; lobe tip Epicarp dull (and usually transparent); monochrome or straight; deflexed and parallel to cotyledon length; multicolored; mottled and streaked; tan to reddish tan; centered between cotyledons; less than 1/2 length of with black or purple overlay; glabrous or pubescent and cotyledons. Plumule rudimentary; glabrous. indurate; with hairs erect or appressed; with 1 type of pubescence; puberulent; with pubescence gray; with Distribution: Eastern tropical Africa to South Africa. pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; glandular; with glandular dots (reddish-brown); without spines; not smooth; with elevated features; reticulately veined; not tuberculate; wrinkled; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð9; length oblique or parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; up to 3 mm long; of 1 length only; filiform; straight, curved, or Lessertia: L. benguellensis J.G. Baker (CÐE), L. spp. (AÐB). hooked and contorted (somewhat near apex). Aril dry; A, Fruits and valves (ϫ 1.3); B, seeds (ϫ 5.3); CÐD, rim-aril; white. testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

825 Genus: Colutea C. Linnaeus adhering to endocarp; dull; not modified by a bloom; colored; monochrome or streaked (faintly and occasion- Phylogenetic Number: 16.05. ally); with infrequent streaks; dark reddish or blackish brown or black (nearly); with tan (greenish) overlay; Tribe: Galegeae. glabrous; smooth or not smooth; with recessed features; pitted with small separate pits; coriaceous. Fracture Subtribe: Coluteinae. lines absent. Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by aril; Species Studied—Species in Genus: 11 spp.—28 spp. with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than puncti- Fruit a legume; unilocular; 3Ð7 ϫ 2Ð3 ϫ 2Ð3 cm; with form; 0.5 mm long; with curved outline; circular; deciduous calyx; without orifice formed by curving of marginal according to radicle tip or between cotyledon fruit or fruit segments; straight; not plicate; not twisted; and radicle lobe; recessed; within rim. Hilum rim color asymmetrical or symmetrical; elliptic; when asym- lighter than testa. Lens not discernible or discernible metrical with both sutures parallelly curved; inflated; (faint); equal to or greater than 0.5 mm in length; 1 mm terete; without or with beak; straight or declined; with long; with margins straight or curved; elliptic or solid beak the same color and texture as fruit; long oblong; not in groove of raphe; adjacent to hilum; 0.2 tapered, tapered, or short tapered at apex; apex aligned mm from hilum; mounded; similar color as testa; lighter with longitudinal axis of fruit; short tapered or rounded than testa and darker than testa; reddish brown or tan at base; base aligned or oblique with longitudinal axis (reddish); not within corona, halo, or rim. Endosperm of fruit; with the apex and base uniform in texture; present (clear to reddish-tan); thick; covering entire membranous or fragile (more or less), thinner than embryo; adnate to embryo. Cotyledons smooth; both chartaceous like Trifolium (21.06); seed chambers outer faces convex; both the same thickness; both more externally invisible. Fruit margin not constricted; or less of equal length; not folded; margin entire 180 without sulcus; plain. Fruit wings absent. Fruit stipitate, degrees from base of radicle; similar at apex; not substipitate, or nonstipitate; with the stipe up to 10 mm concealing radicle; entire over radicle; without lobes; long. Fruit indehiscent or with all layers dehiscing (at with the interface division terminating at base of apex); splitting along sutures (at apex). Dehiscence of radicle; without margins recessed; light reddish tan or valves passive. Replum invisible. Epicarp dull; yellow; inner face concave; glabrous around base of monochrome or multicolored; mottled and streaked; radicle. Embryonic axis deflexed; oblique to length of tan; with purple overlay; glabrous or pubescent but seed; without a joint evident between the radicle and soon deciduous; with 1 type of pubescence; with the cotyledons. Radicle bulbose; deflexed and parallel pubescence uniformly distributed; with simple hairs; to cotyledon length; centered between cotyledons; less pliable; eglandular; without spines; not smooth; with than 1/2 or 1/2 to nearly length of cotyledons. Plumule elevated features; reticulately veined; not tuberculate; rudimentary; glabrous. wrinkled (on drying); not exfoliating; without cracks. Mesocarp present or absent; thin; surface not veined; 1- Distribution: Mediterranean region to eastern and northeast- layered; without balsamic vesicles; without fibers; ern Africa and China and the Himalayas. solid; chartaceous. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp or to epicarp; entire. Seeds 12Ð15; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 2 mm long; of 1 length only; filiform or thick; straight or curved. Aril dry; tongue-aril; tan.

Seed 2Ð4.5 ϫ 2Ð4 ϫ 1Ð1.8 mm; not overgrown; not angular; asymmetrical; oblong or reniform (sub-); compressed; with surface smooth; with or without visible radicle and cotyledon lobes; without external Colutea: C. orientalis P. Miller (CÐE), C. spp. (AÐB). A, groove between radicle and cotyledon lobes; without Fruits (closed and dehisced) (ϫ 0.9); B, seeds (ϫ 3.8); hilar sinus; without umbo on seed faces. Testa not CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 7).

827 Genus: Oreophysa (A.A. von Bunge ex P.E. Boissier) J.F.N. faboid split the same color as the rest of the hilum; Bornmüeller punctiform; between cotyledon and radicle lobe; flush; not within corona, halo, or rim. Lens discernible; equal Phylogenetic Number: 16.06. to or greater than 0.5 mm in length; 0.7 mm long; with margins straight; linear; not in groove of raphe; Tribe: Galegeae. adjacent to hilum; 0.3 mm from hilum; flush; dissimilar color from testa; lighter than testa; black; not within Subtribe: Coluteinae. corona, halo, or rim. Endosperm thin; covering entire embryo; adnate to testa. Cotyledons smooth; both outer Species Studied—Species in Genus: 1 sp.—1 sp. faces convex; both the same thickness; both more or less of equal length; margin entire 180 degrees from Fruit a legume; unilocular; 2.7Ð4 ϫ 2 ϫ 2 (assumed) cm; base of radicle; similar at apex; not concealing radicle; with persistent calyx; with calyx shorter than fruit; entire over radicle; without lobes; with the interface without orifice formed by curving of fruit or fruit division terminating at base of radicle; without margins segments; straight; not plicate; not twisted; asymmetri- recessed; green; inner face flat; glabrous around base of cal; ovate, oblong, or obovate; when asymmetrical with radicle. Embryonic axis deflexed; oblique to length of 1 straight and 1 curved suture or both sutures parallelly seed; with a joint evident between the radicle and the curved; widest near middle or D-shaped; inflated; cotyledons. Radicle linear; lobe tip straight; deflexed terete; without beak; rounded at apex; apex aligned or and parallel to cotyledon length; centered between oblique with longitudinal axis of fruit; rounded at base; cotyledons; less than 1/2 length of cotyledons. Plumule base aligned with longitudinal axis of fruit; with the rudimentary; glabrous. apex and base uniform in texture; membranous or chartaceous; seed chambers externally invisible. Fruit Distribution: Northern Iran. margin not constricted; without sulcus; plain. Fruit wings absent. Fruit substipitate. Fruit with all layers Notes: Jaubert and Spach (1843, plate 64) provides excellent dehiscing (apex); splitting along sutures (apical only). fruit and seed drawings. Dehiscence of valves along both sutures; passive. Replum invisible. Epicarp glossy; monochrome or multicolored; mottled and streaked; tan; with purple overlay (faintly); glabrous; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; wrinkled; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 3Ð4 (assumed); length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 6 ϫ 6 ϫ 2 mm; not overgrown; not angular; asymmetrical; mitaform; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Oreophysa: O. microphylla (H.F. Jaubert & É. Spach) K. Hilum visible; with faboid split; with the lips of the Browicz (AÐE). A, Fruit (ϫ 2.1); B, seed (ϫ 8.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

829 Genus: Sphaerophysa A.-P. de Candolle the rest of the hilum; punctiform; between cotyledon and radicle lobe; flush; within halo. Hilum halo color Phylogenetic Number: 16.07. lighter than testa. Lens discernible; less than 0.5 mm or equal to or greater than 0.5 mm in length; 0.5 mm long; Tribe: Galegeae. with margins curved; elliptic; not in groove of raphe; confluent with hilum; mounded; similar color as testa; Subtribe: Coluteinae. darker than testa; greenish brown; within halo. Lens halo color lighter than testa. Endosperm thin; covering Species Studied—Species in Genus: 1 sp.—2 spp. entire embryo; adnate to embryo. Cotyledons smooth; both outer faces convex; both the same thickness; both Fruit a legume; unilocular; 1.3Ð2.4 ϫ 0.9Ð2 ϫ 0.5Ð1.2 cm; more or less of equal length; not folded; margin entire with persistent calyx; with calyx shorter than fruit; 180 degrees from base of radicle; similar at apex; not without orifice formed by curving of fruit or fruit concealing radicle; entire over radicle; without lobes; segments; straight; not plicate; not twisted; asymmetri- with the interface division terminating at base of cal; oblong; when asymmetrical with both sutures radicle; without margins recessed; yellow; inner face nearly straight; inflated; terete; with beak (fragile) or flat; glabrous around base of radicle. Embryonic axis without; straight; with solid beak the same color and deflexed; oblique to length of seed; without a joint texture as fruit; rounded at apex; apex oblique with evident between the radicle and the cotyledons. Radicle longitudinal axis of fruit; rounded at base; base aligned bulbose; lobe tip straight; deflexed and parallel to with longitudinal axis of fruit; with the apex and base cotyledon length; centered between cotyledons; 1/2 to uniform in texture; membranous (but firm); seed nearly length of cotyledons. Plumule rudimentary; chambers externally invisible. Fruit margin not con- glabrous. stricted; with sulcus; plain. Fruit wings absent. Fruit stipitate or substipitate; with the stipe up to 0.6 mm Distribution: Turkey, Syria, Caucasus, Siberia, central Asia, long. Fruit indehiscent or with all layers dehiscing northern Mongolia, and northern China; introduced in (scarcely). Dehiscence of valves passive. Replum United States. invisible. Epicarp dull; monochrome; tan; glabrous; without spines; not smooth; with elevated features; Notes: Polhill (1981h) noted that Sphaerophysa is scarcely reticulately veined and transversely veined relative to distinct from Smirnowia (16.08). The fruit and seed of fruit length; not tuberculate; wrinkled; not exfoliating; S. kotschyana P.E. Boissier are unknown. without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; spongy; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds at least 40; length oblique or transverse to fruit length; overlapping or touching; in 2 or more series. Funiculus measured; 1Ð2 mm long; of 1 length only; filiform (more so than thick); curved, straight, and hooked. Aril dry; rim-aril; white.

Seed 2Ð2.5 ϫ 2Ð2.5 ϫ 1Ð1.5 mm; not overgrown; not angular; asymmetrical; reniform or mitaform; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; brown to greenish or reddish brown, tan (greenish), or green; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings Sphaerophysa: S. salsula (P.S. von Pallas) A.-P. de absent. Raphe not visible. Hilum visible; with faboid Candolle (AÐE). A, Fruits (ϫ 1.2); B, seeds (ϫ 6.5); split; with the lips of the faboid split the same color as CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

831 Genus: Smirnowia A.A. von Bunge monochrome; reddish brown, tan, or orange; glabrous; smooth; coriaceous. Fracture lines absent. Rim present. Phylogenetic Number: 16.08. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same Tribe: Galegeae. color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; recessed; not within corona, Subtribe: Coluteinae. halo, or rim. Lens discernible; equal to or greater than 0.5 mm in length; 1Ð1.5 mm long; with margins straight Species Studied—Species in Genus: 1 sp.—1 sp. or curved; oblong; not in groove of raphe; confluent with hilum; slightly mounded; similar color as testa; Fruit a legume; unilocular; 3.5 ϫ 2Ð2.5 ϫ 2Ð2.5 cm darker than testa; reddish brown or tan (reddish); not (assumed because all fruits were seen from herbarium within corona, halo, or rim. Endosperm thick; covering sheets); with persistent calyx; with calyx shorter than entire embryo; adnate to embryo. Cotyledons smooth; fruit; without orifice formed by curving of fruit or fruit both outer faces convex; both the same thickness; both segments; straight; not plicate; not twisted; asymmetri- more or less of equal length; not folded; margin entire cal; circular; when asymmetrical with both sutures 180 degrees from base of radicle; similar at apex; not parallelly curved or 1 straight and 1 curved suture; concealing radicle; entire over radicle; without lobes; widest near middle or D-shaped; inflated; terete; with the interface division terminating at base of without beak; short tapered at apex; apex aligned or radicle; yellow; inner face flat; glabrous around base of oblique with longitudinal axis of fruit; rounded at base; radicle. Embryonic axis deflexed; oblique to length of base aligned, oblique, or right angled with longitudinal seed; without a joint evident between the radicle and axis of fruit; with the apex and base uniform in texture; the cotyledons. Radicle bulbose; lobe tip straight; chartaceous; seed chambers externally invisible. Fruit deflexed and parallel to cotyledon length; centered margin not constricted; with sulcus; plain. Fruit wings between cotyledons; less than 1/2 length of cotyledons. absent. Fruit nonstipitate. Fruit indehiscent. Replum Plumule rudimentary; glabrous. invisible. Epicarp dull; monochrome; tan to slightly reddish tan; pubescent and indurate; with 1 type of Distribution: Turkestan. pubescence; puberulent; with pubescence gray; with pubescence uniformly distributed; with simple hairs; Notes: Polhill (1981h) noted that Smirnowia is “scarcely pliable; with hair bases plain; eglandular; without distinct from Sphaerophysa” (16.07). spines; not smooth; with elevated features; reticulately veined and transversely veined relative to fruit length; not tuberculate; wrinkled; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; chartaceous. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds at least 5; length transverse to fruit length; overlapping or neither overlapping nor touching; in 2 or more series. Funicu- lus measured; 2 mm long; of 1 length only; thick; curved or hooked. Aril dry; rim-aril; tan.

Seed 4Ð6 ϫ 3.5Ð5 ϫ 1.5Ð2.5 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; with umbo on seed faces. Testa not adhering to Smirnowia: S. turkestana A.A. von Bunge (AÐE). A, Fruit endocarp; dull; not modified by a bloom; colored; (ϫ 1.3); B, seeds (ϫ 3.2); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

833 Genus: Eremosparton F.E.L. von Fischer & B. Meyer Wings absent. Raphe not visible. Hilum partially or fully concealed; concealed by funicular remnant; with Phylogenetic Number: 16.09. faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; between Tribe: Galegeae. cotyledon and radicle lobe; recessed; within halo. Hilum halo color lighter than testa. Lens faintly Subtribe: Coluteinae. discernible; less than 0.5 mm in length; with margins curved; elliptic; not in groove of raphe; adjacent to or Species Studied—Species in Genus: 3 spp.—3 spp. confluent with hilum; 0.5 mm from hilum; mounded; similar color as testa; within halo. Lens halo color Fruit a legume; unilocular; 1Ð1.5 ϫ 0.8Ð1 ϫ 0.14Ð0.26 cm; lighter than testa. Endosperm thin; covering entire with deciduous or persistent calyx; with calyx shorter embryo; adnate to testa. Cotyledons smooth; both outer than fruit; without orifice formed by curving of fruit or faces convex; both the same thickness; both more or fruit segments; straight; not plicate; not twisted; less of equal length; not folded; margin entire 180 asymmetrical; ovate or falcate (broadly); when asym- degrees from base of radicle; similar at apex; not metrical with 1 straight and 1 curved suture, both concealing radicle; entire over radicle; without lobes; sutures unequally curved, or both sutures parallelly with the interface division terminating at base of curved; widest near middle or D-shaped; inflated or not radicle; without margins recessed; greenish brown or inflated; compressed; with beak; with solid beak the green; inner face flat; glabrous around base of radicle. same color and texture as fruit; short tapered at apex; Embryonic axis deflexed; oblique to length of seed; apex oblique with longitudinal axis of fruit; short with a joint evident between the radicle and the tapered at base; base right angled with longitudinal axis cotyledons. Radicle bulbose; lobe tip straight; deflexed of fruit; with the apex and base uniform in texture; and parallel to cotyledon length; centered between membranous; seed chambers externally visible. Fruit cotyledons; 1/2 to nearly length of cotyledons. Plumule margin not constricted; without sulcus; plain. Fruit rudimentary; glabrous. wings absent (though whole fruit is winglike). Fruit substipitate. Fruit with all layers dehiscing; splitting Distribution: Southeastern Russia and central Asia. along suture. Dehiscence of valves along 1 suture; passive. Replum invisible. Epicarp dull; monochrome; tan; pubescent and indurate; with 1 type of pubescence; villous; with pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; nonseptate; coriaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1 or 2; length parallel with fruit length; overlapping or touching; in 1 series. Funiculus measured; up to 1 mm long; of 1 length only; thick; straight or curved. Aril dry; rim-aril; white.

Seed 3.5Ð4.5 ϫ 3Ð3.5 ϫ 0.7Ð1 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; Eremosparton: E. aphyllum F.E.L. von Fischer & B. Meyer monochrome; reddish to greenish brown; glabrous; (CÐE), E. spp. (AÐB). A, Fruits (ϫ 1.4); B, seeds smooth; chartaceous. Fracture lines absent. Rim absent. (ϫ 6.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

835 Genus: Halimodendron F.E.L. von Fischer ex A.-P. de Raphe not visible. Hilum fully concealed; concealed by Candolle funicular remnant; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; Phylogenetic Number: 16.10. larger than punctiform; 0.4 mm long; with curved outline; circular; marginal according to radicle tip or Tribe: Galegeae. between cotyledon and radicle lobe; recessed; within rim. Hilum rim color darker than testa. Lens discern- Subtribe: Astragalinae. ible; less than 0.5 mm or equal to or greater than 0.5 mm in length; 0.5 mm long; with margins straight or Species Studied—Species in Genus: 1 sp.—1 sp. curved; triangular or irregular; not in groove of raphe; confluent with hilum; barely mounded; dissimilar color Fruit a legume; unilocular; 2Ð4 ϫ 1Ð1.5 ϫ 0.7Ð1 cm; with from testa; darker than testa; black; not within corona, deciduous calyx; without orifice formed by curving of halo, or rim. Endosperm thin; covering entire embryo; fruit or fruit segments; straight; not plicate; not twisted; adnate to testa. Cotyledons smooth; both outer faces symmetrical; oblong or obovate; inflated; compressed; convex; both the same thickness; both more or less of with beak; straight; with solid beak the same color and equal length; not folded; margin entire 180 degrees texture as fruit; short tapered or rounded at apex; apex from base of radicle; similar at apex; not concealing aligned with longitudinal axis of fruit; short tapered at radicle; entire over radicle; without lobes; with the base; base aligned with longitudinal axis of fruit; with interface division terminating at base of radicle; without the apex and base uniform in texture; coriaceous; seed margins recessed; white or tan (pale); inner face flat. chambers externally invisible. Fruit margin not con- Embryonic axis deflexed; oblique to length of seed; stricted; with sulcus (on dorsal suture); plain. Fruit without a joint evident between the radicle and the wings absent. Fruit substipitate. Fruit indehiscent or cotyledons. Radicle bulbose; deflexed and parallel to with all layers dehiscing (tardily); splitting along suture. cotyledon length; centered between cotyledons; less Dehiscence of valves along 1 suture; passive. Replum than 1/2 length of cotyledons. Plumule rudimentary or invisible. Epicarp dull; monochrome or multicolored; moderately developed; glabrous. mottled; dark reddish brown, tan, or black; with black or tan overlay; glabrous; eglandular; without spines; not Distribution: Turkey, Iran, Georgia to Siberia, and Afghani- smooth; with elevated features; veined or not veined; stan. reticulately veined; not tuberculate; faintly wrinkled; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; mottled; tan; with mottling (dark); with brown (reddish) overlay; smooth; nonseptate; coriaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 4Ð5; length parallel with fruit length; overlapping; in 2 or more series. Funiculus measured; up to 1.3 mm long; of 1 length only; thick; straight. Aril dry; rim- aril; white.

Seed 3Ð3.5 ϫ 2.4Ð2.8 ϫ 1.4Ð2.3 mm; not overgrown; not angular; asymmetrical; oblong or reniform; compressed; with surface smooth; with or without visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome or mottled; with frequent mottles; reddish to greenish brown, orange, tan, or Halimodendron: H. halodendron (P.S. von Pallas) A. Voss green; with black overlay; glabrous; smooth; coria- (AÐE). A, Fruits (ϫ 1.4); B, seeds (ϫ 5.3); CÐD, testa ceous. Fracture lines absent. Rim absent. Wings absent. (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

837 Genus: Caragana P.C. Fabricius with small separate pits; chartaceous or coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe Phylogenetic Number: 16.11. visible or not visible; from hilum to lens (at base of seed) or from lens to base of seed and terminating; not Tribe: Galegeae. bifurcating; color of testa (barely darker); flush. Hilum partially concealed; concealed by funicular remnant; Subtribe: Astragalinae. with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform or larger Species Studied—Species in Genus: 21 spp.—ca. 80 spp. than punctiform; 0.3Ð0.5 mm long; with curved outline; circular; marginal according to radicle tip or between Fruit a legume; unilocular; 2Ð6 ϫ 0.25Ð0.6 ϫ 0.15Ð0.4 cm; cotyledon and radicle lobe; flush; not within corona, with persistent calyx; with calyx shorter than fruit; halo, or rim. Lens discernible or not discernible; equal without orifice formed by curving of fruit or fruit to or greater than 0.5 mm or less than 0.5 mm in length; segments; straight; not plicate; not twisted; asymmetri- 0.2Ð1 mm long; with margins straight or curved; cal; linear; when asymmetrical with both sutures nearly wedge-shaped, circular, or elliptic; not in groove of straight; not inflated or inflated; compressed; without or raphe; adjacent to or confluent with hilum; up to 3 mm with beak; declined; with solid beak the same color and from hilum; mounded or recessed; similar color as texture as fruit; short tapered at apex; apex oblique with testa; barely darker than testa; reddish brown; not longitudinal axis of fruit; short tapered at base; base within corona, halo, or rim. Endosperm thin; covering oblique with longitudinal axis of fruit; with the apex entire embryo; adnate to testa. Cotyledons smooth; both and base uniform in texture; coriaceous; seed chambers outer faces convex; both the same thickness; both more externally invisible. Fruit margin not constricted; or less of equal length or 1 longer than other; not folded without sulcus; plain. Fruit wings absent. Fruit (though occasionally 1 cotyledon larger than other); nonstipitate. Fruit with all layers dehiscing; splitting margin entire 180 degrees from base of radicle; similar along sutures. Dehiscence of valves along both sutures; at apex; not concealing radicle; entire over radicle; apical and down; active; with valves twisting. Replum without lobes; with the interface division terminating at invisible. Epicarp dull; monochrome; dark reddish base of radicle; without margins recessed; reddish tan; brown; glabrous; glandular or eglandular; with glandu- inner face flat; glabrous around base of radicle. Embry- lar dots; without spines; not smooth; with elevated onic axis deflexed; oblique to length of seed; without a features; reticulately veined; not tuberculate; somewhat joint evident between the radicle and the cotyledons. wrinkled; not exfoliating; without cracks. Mesocarp Radicle linear; lobe tip straight; deflexed and parallel to thin; surface not veined; 1-layered; without balsamic cotyledon length; centered between cotyledons; less vesicles; without fibers; solid; coriaceous. Endocarp than 1/2 length of cotyledons. Plumule rudimentary or dull; monochrome; reddish tan; smooth; nonseptate; moderately developed; glabrous. chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 2Ð5; length parallel Distribution: Eastern Europe, Asia, and naturalized in North with fruit length; neither overlapping nor touching; in 1 America. series. Funiculus measured; up to 1 mm long; of 1 length only; thick; straight. Aril dry; rim-aril; white. Notes: Yakovlev and Sviazeva (1984, 1985a, 1987) have treated some of the Russian species, and Gorbunova Seed 2.5Ð10 ϫ 2Ð4.5 ϫ 2Ð4.5 mm; not overgrown; not (1984) established a sectional and subsectional se- angular or angular (barely); asymmetrical; oblong, quence for the genus. linear, circular, or reniform; terete or compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull or glossy; not modified by a bloom; colored; monochrome or mottled and streaked; with frequent mottles; with frequent streaks; dark reddish to orangish brown or Caragana: C. densa V.L. Komarov (CÐE), C. spp. (AÐB). A, orange; with brown (dark reddish) overlay; glabrous; Valves and dehisced fruits (ϫ 2.1); B, seeds (ϫ 3.4); smooth or not smooth; with recessed features; pitted CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

839 Genus: Calophaca F.E.L. von Fischer ex A.-P. de Candolle pressed; with surface smooth; with or without visible radicle and cotyledon lobes; without external groove Phylogenetic Number: 16.12. between radicle and cotyledon lobes; with deep hilar sinus; without umbo on seed faces. Testa not adhering Tribe: Galegeae. to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth or not Subtribe: Astragalinae. smooth; with elevated features; warty or wrinkled; coriaceous. Fracture lines absent. Rim absent. Wings Species Studied—Species in Genus: 3 spp.—5 spp. absent. Raphe not visible. Hilum partially or fully concealed; concealed by funicular remnant, radicle Fruit a legume; unilocular; 1Ð3 ϫ 0.3Ð0.5 ϫ 0.4Ð0.5 cm; lobe, or wing; with faboid split; with the lips of the with persistent calyx; with calyx shorter than fruit; faboid split the same color as the rest of the hilum; without orifice formed by curving of fruit or fruit larger than punctiform or punctiform; up to 0.7 mm segments; straight; not plicate; not twisted; asymmetri- long; with curved outline; elliptic; marginal according cal; linear; when asymmetrical with both sutures nearly to radicle tip or between cotyledon and radicle lobe; straight; inflated (when mature); terete (when mature); flush; not within corona, halo, or rim. Lens discernible with beak; straight; with solid beak the same color and or not discernible; equal to or greater than 0.5 mm in texture as fruit; short tapered at apex; apex aligned or length; 1 mm long; with margins straight; linear; not in oblique with longitudinal axis of fruit; tapered at base; groove of raphe; confluent with hilum; flush; dissimilar base aligned with longitudinal axis of fruit; with the color from testa; darker than testa; black; not within apex and base uniform in texture; coriaceous; seed corona, halo, or rim. Endosperm thin; covering entire chambers externally invisible. Fruit margin not con- embryo; adnate to testa or embryo. Cotyledons smooth; stricted; without sulcus; plain or embellished; with both outer faces convex; both the same thickness; both fringe (plain and glandular hairs). Fruit wings absent. more or less of equal length; not folded; margin entire Fruit nonstipitate. Fruit with all layers dehiscing; 180 degrees from base of radicle; similar at apex; not splitting along sutures. Dehiscence of valves along both concealing radicle; entire over radicle; without lobes; sutures; apical and down; active; with valves twisting. with the interface division terminating at base of Replum invisible. Epicarp dull; monochrome; dark radicle; without margins recessed; reddish to dark reddish brown; pubescent and indurate; with 1 or 2 reddish brown; inner face flat; glabrous around base of types of pubescence; puberulent or villous (restricted to radicle. Embryonic axis deflexed; oblique to length of sutures or throughout); with pubescence reddish brown seed; without a joint evident between the radicle and or gray; with long and short gray plain-tipped hairs; the cotyledons. Radicle bulbose; lobe tip curved or with pubescence uniformly distributed; with simple and hooked; deflexed and parallel to cotyledon length; glandular hairs or glandular hairs; pliable; with hair centered between cotyledons; less than 1/2 or 1/2 to bases plain; glandular; with glandular hairs (reddish- nearly length of cotyledons. Plumule rudimentary; brown) or dots (reddish-brown); without spines; not glabrous. smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp Distribution: Central Asia (4 spp.) and Russia (C. thick; surface not veined; 2-layered; without balsamic wolgarica). vesicles; without fibers; with vitreous layer over solid layer; coriaceous. Endocarp dull; monochrome; tan; cobwebby and spongy (but thinly filled); subseptate or nonseptate; with septa thin (tissue paper-like), flexible; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð8; length oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 1Ð2 mm long; of 1 length only; somewhat filiform; straight or curved. Aril absent. Calophaca: C. wolgarica (C. Linnaeus f.) F.E.L. von Fischer ex A.-P. de Candolle (CÐE), C. spp. (AÐB). A, Seed 4Ð8 ϫ 0.4Ð5 ϫ 0.1Ð3.3 mm; not overgrown; not Fruits (dehisced and closed) (ϫ 1.4); B, seeds (ϫ 5.8); angular; asymmetrical; subreniform or oblong; com- CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

841 Genus: Chesneya J. Lindley ex S.F.L. Endlicher seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled Phylogenetic Number: 16.14. and streaked; with frequent mottles; with frequent streaks; brown to reddish brown or tan; with black Tribe: Galegeae. overlay; glabrous; smooth or not smooth; with elevated or recessed features; wrinkled, warty, shagreen, or Subtribe: Astragalinae. reticulate; pitted with small separate pits or large depressions on each face (2); chartaceous. Fracture Species Studied—Species in Genus: 9 spp.—20 spp. lines absent. Rim absent. Wings absent. Raphe from hilum to lens; not bifurcating; darker than testa; Fruit a legume; unilocular; 2.5Ð8 ϫ 0.5Ð1.2 ϫ 0.2Ð0.5 cm; blackish brown or black; flush. Hilum fully concealed; with persistent calyx; with calyx shorter than fruit; concealed by radicle lobe or wing; with faboid split; without orifice formed by curving of fruit or fruit with the lips of the faboid split the same color as the segments; straight; not plicate; not twisted; asymmetri- rest of the hilum; punctiform; between cotyledon and cal; lanceolate, linear (to lanceolate), or oblong; when radicle lobe; flush; not within corona, halo, or rim. Lens asymmetrical with both sutures nearly straight; not barely discernible or not discernible; less than 0.5 mm inflated or inflated; compressed; with beak; straight; in length; with margins curved; more or less circular; with solid beak the same color and texture as fruit; not in groove of raphe; adjacent to hilum; 1.5 mm from tapered at apex; apex aligned with longitudinal axis of hilum; flush; dissimilar color from testa; darker than fruit; rounded at base; base aligned with longitudinal testa; blackish brown; not within corona, halo, or rim. axis of fruit; with the apex and base uniform in texture; Endosperm thick; covering entire embryo; adnate to coriaceous; seed chambers externally faintly visible; testa. Cotyledons smooth; both outer faces convex; both with the raised seed chambers not torulose. Fruit the same thickness; both more or less of equal length; margin not constricted; without sulcus; plain. Fruit not folded; margin entire 180 degrees from base of wings absent. Fruit nonstipitate. Fruit with all layers radicle; similar at apex; partially concealing radicle; dehiscing; splitting along sutures. Dehiscence of valves entire over radicle; without lobes; with the interface along both sutures; apical and down; active or passive; division terminating at base of radicle; without margins with valves twisting. Replum invisible. Epicarp dull; recessed; reddish brown; inner face flat; glabrous monochrome; reddish brown; pubescent and indurate; around base of radicle. Embryonic axis deflexed; with hairs erect or appressed; with 1 type of pubes- oblique to length of seed; without a joint evident cence; puberulent or villous; with pubescence gray; between the radicle and the cotyledons. Radicle linear; with pubescence uniformly distributed; with simple lobe tip hooked; deflexed and parallel to cotyledon hairs; pliable; with hair bases plain; eglandular; without length; centered between cotyledons; less than 1/2 or spines; not smooth; with elevated features; reticulately 1/2 to nearly length of cotyedons. Plumule rudimentary; veined; not tuberculate; not exfoliating; without cracks. glabrous. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Distribution: Central and southwestern Asia to Mongolia. Endocarp dull; monochrome; brown; smooth and scurfy or cobwebby; septate or nonseptate; with septa thin (tissue paper-like), flexible; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 8Ð12; length parallel with fruit length; touching or neither overlapping nor touching; in 1 series. Funiculus measured; 0.5Ð 4 mm long; of 1 length only; filiform (or nearly so); straight or curved (slightly). Aril absent.

Seed 2.5Ð6 ϫ 2Ð4 ϫ 1Ð1.2 mm; not overgrown; not angular or angular; asymmetrical; mitaform (and bent on faces), reniform (and bent on faces), or irregular; compressed; Chesneya: C. rytidosperma H.F. Jaubert & É. Spach (CÐE), with surface smooth; without visible radicle and C. spp. (AÐB). A, Fruits (dehisced) (ϫ 1.1); B, seeds cotyledon lobes; without hilar sinus; without umbo on (ϫ 5.7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

843 Genus: Astragalus C. Linnaeus green, or purple; with purple, red, brown (purplish), or green overlay; with mottling over seed chambers; Phylogenetic Number: 16.15. pubescent and indurate or glabrous; with hairs erect or appressed; with 1 or 2 types of pubescence; pilose, Tribe: Galegeae. puberulent, tomentose, velutinous, villous, or sericeous; with pubescence gray or black (to rusty); with gray and Subtribe: Astragalinae. black hairs intermixed; with pubescence uniformly distributed; with simple or complex hairs; T-shaped Species Studied—Species in Genus: 195 spp.—ca. 2,000 hairs (malpighiaceous); pliable or stiff; with hair bases spp. plain or swollen (somewhat); antrorse; straight or hooked at apex; eglandular or glandular; with glandular Fruit a legume; unilocular or bilocular (including dots (reddish); without spines; not smooth or smooth; semibilocular); 0.2Ð20 ϫ 0.15Ð6 ϫ 0.1Ð3 cm; with with elevated features; veined (venation not ribbed to persistent calyx; with calyx shorter than, equal in length ribbed); reticulately veined or transversely veined to, or longer than fruit (and calyx inflated or not); relative to fruit length; not tuberculate; rugose (to without or with orifice formed by curving of fruit or reticulate), verrucose-rugose, or wrinkled (irregularly to fruit segments; straight, curved (or slightly curved), 0.5- regularly or cross to lengthwise); not exfoliating; coiled, 1-coiled, or S-curved; not plicate; not twisted; without cracks. Mesocarp present or absent (or nearly symmetrical or asymmetrical; linear, oblong, elliptic, so); thin or trace; surface not veined; 1-layered; without circular, didymous, ovate, C-shaped, or falcate; when balsamic vesicles; without fibers; solid; ligneous (or asymmetrical with both sutures parallelly curved, both subligneous), coriaceous, or chartaceous. Endocarp sutures unequally curved, or 1 straight and 1 curved dull; monochrome; tan; smooth, cobwebby, or fibrous; suture; widest near middle or D-shaped; not inflated or nonseptate (longitudinal separation between seed inflated; compressed, terete, subtriangular (to triangu- chambers making 2 loculi); chartaceous; not exfoliat- lar), or quadrangular; without or with beak; straight, ing; remaining fused to mesocarp and epicarp; remain- declined, coiled, or hooked; with solid beak the same ing fused to epicarp; entire. Seeds 1Ð20; length parallel color and texture as fruit; short tapered, tapered, long with or transverse to fruit length; neither overlapping tapered, or rounded at apex; apex aligned or oblique nor touching, touching, or overlapping; in 2 or more or with longitudinal axis of fruit; short tapered, tapered, 1 series. Funiculus less than 0.5 mm long or measured; long tapered, rounded, or truncate at base; base aligned up to 15 mm long; of 1 length only (and with or without with longitudinal axis of fruit; with the apex and base hairs); filiform, thick, or triangular; straight, curved, or uniform in texture; coriaceous, leathery, membranous, hooked. Aril dry; rim-aril; greenish brown, tan, white, chartaceous, ligneous, drupaceous, or fleshy (when or green. fresh); seed chambers externally invisible or visible; with the raised seed chambers not torulose. Fruit Seed 1Ð7 ϫ 0.8Ð5 ϫ 0.1Ð0.6 mm; not overgrown or rarely margin not constricted or constricted (one species, A. overgrown, 1 seed filling entire fruit cavity; not angular shelkovnikovii A.A. Grossheim); without or with sulcus or angular (dented or not on faces); asymmetrical or (intrusion of dorsal or dorsal and ventral sutures); plain symmetrical (except hilum); mitaform, oblong, reni- or embellished; with fringe (of spines). Fruit wings form, rectangular, rhombic, triangular, pyriform, absent or present (narrow). Fruit nonstipitate (or sessile elliptic, D-shaped, cordate, or irregular; compressed; on stipelike gynophore), substipitate, or stipitate; with with surface smooth; without or with visible radicle and the stipe up to 35 mm long. Fruit with all layers cotyledon lobes; without or with external groove dehiscing (valves may completely separate but because between radicle and cotyledon lobes; with external of the longitudinal walls which separate the 2 valves in groove between radicle and cotyledon lobes same color bilocular legume, each half functioning as an “indehis- as testa; with deep or shallow hilar sinus or without cent fruit”) or indehiscent (fragile and usually inflated hilar sinus; without umbo on seed faces. Testa not and easily fracturing); splitting along sutures. Dehis- adhering to endocarp; dull or glossy; not modified by a cence of valves along 1 suture or both sutures; apical bloom; colored; monochrome or mottled and streaked; and down or basal and up; active or passive; with with frequent or infrequent mottles; with frequent or valves reflexing (inflexed). Replum invisible. Epicarp infrequent streaks; reddish to yellowish brown to dull; monochrome or multicolored; mottled; brown to brown, tan (to greenish tan), cream, green (yellowish), reddish or blackish brown, tan (to purplish tan), black, olive, orange, yellow, purple, or black; with black,

844 purple, or red overlay; glabrous; smooth (sometimes 1947); highlands of tropical Africa, Gillett (1963a); with a fine reticulate-colored pattern on smooth Iran, Maassoumi (1986, 1989); Russia, Gontscharov surface) or not smooth; with elevated or recessed and Borisova (1946); Pakistan and Himalayas, Ali features; rugose or wrinkled; pitted with small separate (1961). In addition Maassoumi, Podlech, and others pits or concaved; coriaceous or chartaceous. Fracture have issued a series of more than 30 papers for the Near lines absent. Rim absent. Wings absent. Raphe not East. Several sections of Astragalus have been visible or visible; from hilum to lens; not bifurcating; monographed: Section Acanthophace A.A. von Bunge darker than testa; reddish brown to brown; flush. Hilum and section Aegacantha A.A. von Bunge (Deml 1972); visible or partially concealed; concealed by wing; with section Alopecuroidei A.-P. de Candolle (Becht 1978); faboid split; with the lips of the faboid split the same section Caprini A.-P. de Candolle (Podlech 1988); color as the rest of the hilum; punctiform or larger than sections Chlorostachys A.A. von Bunge, Phyllolobium punctiform; up to 0.5 mm long; with curved outline; A.A. von Bunge, and Skythropos N.D. Simpson circular; between cotyledon and radicle lobe; recessed; (Wenninger 1991); section Chronopus A.A. von Bunge not within corona, halo, or rim or within halo. Hilum (Ott 1978); section Dasyphyllium A.A. von Bunge halo color lighter than testa. Lens discernible or not (Aytaç 1997); section Eremophysa A.A. von Bunge and discernible; less than 0.5 mm or equal to or greater than section Eremophysopsis N.F. Gontscharov (Podlech 0.5 mm in length; up to 1 mm long; with margins 1993); section Herpocaulos A.A. von Bunge (Podlech straight or curved; oblong, linear, elliptic, or 2 circular 1984); section Laxiflori C. Agerer-Kirchhoff (Agerer- mounds separated by groove; not in groove of raphe; Kirchhoff and Agerer 1977); section Megalocystis A.A. adjacent to or confluent with hilum; 0.2 mm from von Bunge (Tietz and Zarre 1994); section Platyglottis hilum; mounded or flush; similar or same color as testa; A.A. von Bunge (Podlech 1990); section Sesamei A.-P. darker than testa; brown or tan (greenish); not within de Candolle (Gazer 1993); and section Theiochrus A.A. corona, halo, or rim. Endosperm thick or thin; covering von Bunge (Podlech and Kozik 1983). Sanderson and entire embryo; adnate to embryo or testa. Cotyledons Liston (1995) cladistically analyzed the Galegeae smooth; both outer faces convex; both the same genera using molecular data. They concluded that thickness; both more or less of equal length; not folded; Astragalus is monophyletic with Astracantha (16.16) margin entire 180 degrees from base of radicle; similar nested within it and that Astracantha should be in- at apex; not concealing radicle; entire over radicle; cluded in it, possibly as a subgenus. Zarre and Podlech without lobes; with the interface division terminating at (1997), using morphological and anatomical data, base of radicle; without margins recessed; tan or brown concurred that Astracantha should not be maintained as (reddish); inner face flat; glabrous around base of a genus. They concluded that it should be synonymized radicle. Embryonic axis deflexed; oblique to length of with Astragalus and that it cannot be maintained even seed; without a joint evident between the radicle and at subgeneric level. Polhill (1994b) maintained the cotyledons. Radicle bulbose (some almost 1/2 size Neodielsia H.A.T. Harms (16.18) as a genus, but of cotyledons); lobe tip straight, curved, or hooked; Mabberley (1997) kept it as a synonym of Astragalus. deflexed and parallel to cotyledon length; centered We have chosen to follow Mabberley. Barneby (1964) between cotyledons; less than 1/2 or 1/2 to nearly and others, including Hutchinson (1964), who noted length of cotyledons. Plumule rudimentary or moder- that “an important feature of the Astragalus pod is the ately developed; glabrous. septum or longitudinal wall produced across the cavity from the dorsal suture.” Because we have restricted the Distribution: North America to South America (Patagonia), term “septum” to the transverse wall of a legume that Europe, Russia, temperate northern Africa, tropical separates seeds, we will not use “septum” in the sense Africa (mountains), the Iranian Plateau, northern India, used by Barneby and others. We agree with Barneby China, and Japan. that legumes endowed with a complete internal longitudinal wall are to be termed bilocular (replacing two- Notes: Astragalus has more species than any other legume celled fruits of the literature). A partial walled cavity is genus and more than most other genera, regardless of termed “semibilocular,” and a pod lacking this wall is family. There are a remarkable number of publications “unilocular.” Barneby (1964) has a detailed discussion dealing with the species in various parts of the world, of Astragalus fruits. and a few of the major ones include: North America, Barneby (1964) and Isely (1983, 1984, 1985, 1986); southern temperate South America, Johnston (1938,

845 Astragalus: A. crassicarpus T. Nuttall (DÐF), A. spp. (AÐC). A, Fruits (ϫ 1); BÐC, seeds (ϫ 4.1, ϫ 4.5); DÐE, testa (ϫ 50, ϫ 1000); F, embryos (ϫ 10).

847 Genus: Biserrula C. Linnaeus colored; monochrome or mottled; with frequent mottles; reddish brown, tan, white, or yellow; with Phylogenetic Number: 16.15A. black or purple overlay; glabrous; not smooth; with elevated or recessed features; reticulately wrinkled; Tribe: Galegeae. punctate; chartaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Subtribe: Astragalinae. partially or fully concealed; concealed by radicle lobe or wing; with faboid split; with the lips of the faboid Species Studied—Species in Genus: 1 sp.—1 sp. split the same color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; flush; within rim. Fruit a legume; bilocular; 1.2Ð5 ϫ 0.4Ð1 ϫ 0.3Ð0.7 cm; Hilum rim color of testa. Lens discernible; less than 0.5 with deciduous calyx; without orifice formed by mm in length; with margins straight; diamond-shaped; curving of fruit or fruit segments; straight or curved not in groove of raphe; confluent with hilum; mounded; (slightly); not plicate (but occasionally irregularly dissimilar color from testa; darker than testa; reddish bent); not twisted; symmetrical; linear; not inflated; brown; not within corona, halo, or rim. Endosperm flattened; without beak; short tapered at apex; apex thick; covering entire embryo; adnate to embryo. aligned with longitudinal axis of fruit; short tapered and Cotyledons smooth; both outer faces convex; both the truncate at base; base aligned with longitudinal axis of same thickness; both more or less of equal length; not fruit; with the apex and base uniform in texture; seed folded; margin entire 180 degrees from base of radicle; chambers externally invisible. Fruit margin constricted similar at apex; not concealing radicle; entire over along both margins; without sulcus; embellished or radicle; without lobes; with the interface division plain (B. pelecinus subsp. leiocarpa (A. Richard) J.B. terminating at base of radicle; without margins re- Gillett); with fringe (sinuate dentate with denticulate cessed; reddish brown, tan, yellow, or white; inner face lobes). Fruit wings absent. Fruit nonstipitate. Fruit flat; glabrous around base of radicle. Embryonic axis indehiscent. Replum invisible. Epicarp dull; mono- deflexed; oblique to length of seed; without a joint chrome; brown to reddish brown; glabrous or pubescent evident between the radicle and the cotyledons. Radicle but soon deciduous; with 1 type of pubescence; pilose; bulbose; lobe tip straight or curved (slightly); deflexed with pubescence gray; with pubescence uniformly and parallel to cotyledon length; centered between distributed; with simple hairs; pliable; with hair bases cotyledons; 1/2 to nearly length of cotyledons. Plumule plain; eglandular; without spines; not smooth; with moderately developed; glabrous. elevated features; transversely veined relative to fruit length and reticulately veined; not tuberculate; not Distribution: Mediterranean region and northeastern and exfoliating; without cracks. Mesocarp thin; surface not eastern tropical Africa. veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; Notes: Barneby (1964) included this taxon in Astragalus reddish brown; smooth and cobwebby (near sutures and (16.15) as A. pelecinus (C. Linnaeus) R.C. Barneby. funiculi); nonseptate (longitudinal separation between seed chambers making 2 locules); chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 5Ð21; length parallel with or transverse to fruit length; neither overlapping nor touching; in 2 or more series. Funiculus measured; 0.5Ð0.8 mm long; of 1 length only; filiform; straight. Aril dry; rim-aril; reddish brown.

Seed 1.5Ð2 ϫ 1.5Ð2 ϫ 0.7Ð0.8 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; with deep hilar sinus; without umbo on Biserrula: B. pelecinus C. Linnaeus (AÐE). A, Fruits seed faces. Testa not adhering to endocarp; dull; not (ϫ 2.3); B, seeds (ϫ 8); CÐD, testa (ϫ 50, ϫ 1000); modified by a bloom; clear (except for mottles) or E, embryos (ϫ 15).

849 Genus: Astracantha D. Podlech seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled; Phylogenetic Number: 16.16. reddish brown or tan (reddish to greenish); glabrous; smooth or not smooth; with elevated or recessed Tribe: Galegeae. features; rugose or wrinkled; pitted with small separate pits; coriaceous. Fracture lines absent. Rim absent. Subtribe: Astragalinae. Wings absent. Raphe visible or not visible; from hilum to lens; not bifurcating; darker than testa; reddish Species Studied—Species in Genus: 5 spp.—215 spp. brown; raised. Hilum visible or fully concealed; concealed by funicular remnant; with faboid split; with Fruit a legume; unilocular; 0.3Ð0.6 ϫ 0.15Ð0.35 ϫ 0.15Ð the lips of the faboid split the same color as the rest of 0.35 cm; with persistent calyx; with calyx longer or the hilum; punctiform or larger than punctiform; 0.3Ð shorter than fruit; without orifice formed by curving of 0.4 mm long; with curved outline; more or less circular; fruit or fruit segments; straight; not plicate; not twisted; between cotyledon and radicle lobe; slightly recessed; asymmetrical or symmetrical; circular; when asym- within rim. Hilum rim color of testa. Lens discernible; metrical with both sutures unequally or parallelly less than 0.5 mm in length; with margins straight or curved; inflated; terete; without beak; rounded at apex; curved; triangular or circular; not in groove of raphe; apex aligned with longitudinal axis of fruit; rounded at adjacent to or confluent with hilum; up to 0.3 mm from base; base aligned with longitudinal axis of fruit; with hilum; mounded; similar color as testa; darker than the apex and base uniform in texture; coriaceous; seed testa; reddish brown; not within corona, halo, or rim. chambers externally invisible. Fruit margin not con- Endosperm thin; covering entire embryo; adnate to stricted; without sulcus; plain. Fruit wings absent. Fruit testa. Cotyledons smooth; both outer faces convex; both nonstipitate. Fruit with all layers dehiscing; splitting the same thickness; both more or less of equal length; along sutures. Dehiscence of valves along both sutures; not folded; margin entire 180 degrees from base of apical and down; passive. Replum invisible. Epicarp radicle; similar at apex; not concealing radicle; entire dull; monochrome; black; pubescent and indurate; with over radicle; without lobes; with the interface division 1 type of pubescence; somewhat villous; with pubes- terminating at base of radicle; white; inner face flat; cence dull and dark golden; with pubescence uniformly glabrous around base of radicle. Embryonic axis distributed; with simple hairs; pliable; with hair bases deflexed; parallel to length of seed; without a joint swollen; eglandular; without spines; smooth or not evident between the radicle and the cotyledons. Radicle smooth; with elevated features; not veined; not tubercu- bulbose; lobe tip straight; deflexed and parallel to late; somewhat wrinkled; not exfoliating; without cotyledon length; centered between cotyledons; less cracks. Mesocarp thin; surface not veined; 1-layered; than 1/2 or 1/2 to nearly length of cotyledons. Plumule without balsamic vesicles; without fibers; solid; rudimentary; glabrous. coriaceous. Endocarp glossy; monochrome; brownish black; smooth; nonseptate; chartaceous; not exfoliating; Distribution: Europe and Russia. remaining fused to mesocarp and epicarp; entire. Seeds 1(Ð2); length parallel with or transverse to fruit length; Notes: Podlech (1983) founded the genus, a segregate of neither overlapping nor touching; in 1 series. Funiculus Astragalus (16.15), and the species count is based on less than 0.5 mm long; of 1 length only; thick; assumed his work. Reer and Podlech (1986) expanded on the straight. Aril absent or present; dry; rim-aril; tan. original paper, and this was followed by a seed morphology study with testa micrographs by Engel (1990). Seed 2Ð7 ϫ 2Ð4.5 ϫ 1Ð2 mm; not overgrown; angular or Sanderson and Liston (1995) performed cladistic not angular; asymmetrical; mitaform, oblong, rectangu- analyses for the Galegeae genera using molecular data. lar (more or less), or D-shaped; compressed; with They concluded that Astragalus (16.15) is monophyl- surface smooth; with visible radicle and cotyledon etic with Astracantha nested within it and that lobes; with or without external groove between radicle Astracantha should be returned to it, possibly as a and cotyledon lobes; with external groove between subgenus. Zarre and Podlech (1997), using morphologi- radicle and cotyledon lobes same color as or lighter in cal and anatomical data, concurred that Astracantha color than testa; without hilar sinus; without umbo on

850 should not be maintained as a genus. They concluded that it should be synonymized with Astragalus and that it cannot be maintained even at the subgeneric level. Our number of seed and fruit samples is inadequate, but our morphological data were enhanced by the cited literature.

851 Astracantha: A. echinus (A.-P. de Candolle) D. Podlech (A), A. gossypina (F.E.L. von Fischer) D. Podlech (CÐE), A. spp. (B). A, Fruit (ϫ 2.8); B, seeds (ϫ 6.5); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 1).

853 Genus: Oxytropis A.-P. de Candolle segments. Seeds (3Ð)5Ð11; length parallel with fruit length; neither overlapping nor touching; in 2 or more Phylogenetic Number: 16.17. or 1 series. Funiculus measured; 1 mm long; of 1 length only; filiform; S-curved or curved. Aril dry; rim-aril; Tribe: Galegeae. reddish brown.

Subtribe: Astragalinae. Seed 0.75Ð3.5 ϫ 1Ð1.5 ϫ 0.5Ð1 mm; not overgrown; not angular; asymmetrical; mitaform, reniform, circular, or Species Studied—Species in Genus: 31 spp.—ca. 300 spp. quadrangular; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external Fruit a legume; unilocular; 0.8Ð4 ϫ 0.3Ð0.8 ϫ 0.3 cm; with groove between radicle and cotyledon lobes; with persistent calyx; with calyx shorter or longer than fruit; external groove between radicle and cotyledon lobes without orifice formed by curving of fruit or fruit same color as testa; without hilar sinus; without umbo segments; straight or curved (or slightly curved); not on seed faces. Testa not adhering to endocarp; dull; not plicate; not twisted; symmetrical or asymmetrical; modified by a bloom; colored; monochrome or mottled oblong, linear, circular, falcate, or C-shaped; when and streaked; with frequent mottles; with frequent asymmetrical with both sutures parallelly curved; not streaks; brown to reddish to greenish or pinkish brown, inflated or inflated; compressed or terete; with beak; tan, yellow (greenish), orange, olive, green, or black; straight or declined; with solid beak the same color and with black or purple overlay; glabrous; smooth or not texture as fruit; short tapered, tapered, or long tapered smooth; with elevated features; rugose or shagreen; at apex; apex aligned, oblique, or right-angled with coriaceous. Fracture lines absent. Rim absent. Wings longitudinal axis of fruit; long tapered or short tapered absent. Raphe not visible. Hilum visible; with faboid at base; base aligned or right angled with longitudinal split; with the lips of the faboid split the same color as axis of fruit; with the apex and base uniform in texture; the rest of the hilum; punctiform; between cotyledon chartaceous, membranous, coriaceous, or ligneous; seed and radicle lobe; flush; not within corona, halo, or rim chambers externally invisible. Fruit margin not con- or within halo. Hilum halo color lighter than testa. Lens stricted; with sulcus (seed bearing suture more or less discernible; less than 0.5 mm in length; with margins intruded but legume undivided); plain. Fruit wings curved; elliptic; not in groove of raphe; adjacent to or absent. Fruit nonstipitate, substipitate, or stipitate; with confluent with hilum; 0.2 mm from hilum; mounded; the stipe up to 15 mm long. Fruit with all layers similar color as testa; darker than testa; reddish brown; dehiscing; splitting along sutures. Dehiscence of valves not within corona, halo, or rim. Endosperm thick; along 1 suture or both sutures; apical and down; passive covering entire embryo; adnate to embryo. Cotyledons or active; with valves twisting (loosely). Replum smooth; both outer faces convex; both the same invisible. Epicarp dull; monochrome or multicolored; thickness; both more or less of equal length; not folded; mottled; brown; with purple overlay; pubescent and margin entire 180 degrees from base of radicle; similar indurate or pubescent but soon deciduous; with hairs at apex; not concealing radicle; entire over radicle; appressed or erect; with 1 or 2 types of pubescence; without lobes; with the interface division terminating at puberulent, villous, or velutinous; with pubescence base of radicle; without margins recessed; tan; inner golden or gray and black; with gray and black hairs face flat; glabrous around base of radicle. Embryonic intermixed; with pubescence uniformly distributed; axis deflexed; oblique to length of seed; without a joint with simple hairs; pliable; with hair bases plain; evident between the radicle and the cotyledons. Radicle glandular or eglandular; without spines; not smooth; bulbose; lobe tip straight; deflexed and parallel to with elevated features; reticulately veined or trans- cotyledon width; centered between cotyledons; less versely veined relative to fruit length; not tuberculate; than 1/2 length of cotyledons. Plumule rudimentary; wrinkled; not exfoliating; without cracks. Mesocarp glabrous. present or absent; thin or thick; surface not veined; 1- layered; without balsamic vesicles; without fibers; Distribution: North America (22 spp.) and Eurasia. solid; coriaceous or chartaceous. Endocarp dull or glossy; monochrome; tan; smooth or cowebby; Notes: Barneby (1952) monographed the North American nonseptate; chartaceous or coriaceous; not exfoliating; species and noted that only two species were found remaining fused to mesocarp and epicarp; remaining between 1802 and 1951 that challenged the original fused to epicarp; separating into 1-seeded winged circumscription of Oxytropis. Oxytropis has an “intro-

854 flexion of the pod’s ventral suture, as opposed to the muticous keel of Phaca and Astragalus, with their pods unilocular or bilocular (from the dorsal suture) respectively.” Phaca C. Linnaeus is subsumed into Astragalus (16.15). Fedchenko et al. (1948) reported on the species in Russia, and Ulziykhutag (1979) summarized the 78 Mongolian species in 17 sections. The testa surfaces and transverse sections of O. riparia D.I. Litvinov and O. campestris (C. Linnaeus) A.-P. de Candolle were studied using SEM (Solum and Lockerman 1991). Pandey and Jha (1988) also reported on testa micrographs of three species of Oxytropis, including O. compestris.

855 Oxytropis: O. pilosa (C. Linnaeus) A.-P. de Candolle (CÐE), O. spp. (AÐB). A, Fruits (ϫ 1.5); B, seeds (ϫ 5.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

857 Genus: Gueldenstaedtia F.E.L. von Fischer lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on Phylogenetic Number: 16.19. seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome or Tribe: Galegeae. mottled and streaked; greenish brown; with black overlay; glabrous; smooth or not smooth; with elevated Subtribe: Astragalinae. or recessed features; reticulate; pitted with small separate pits; chartaceous. Fracture lines absent. Rim Species Studied—Species in Genus: 9 spp.—14 spp. absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by radicle lobe or wing; Fruit a legume; unilocular; 1.2Ð3 ϫ 0.25Ð0.4 ϫ 0.13Ð0.15 with faboid split; with the lips of the faboid split the cm; with persistent or deciduous calyx; with calyx same color as the rest of the hilum; punctiform; shorter than fruit; without orifice formed by curving of between cotyledon and radicle lobe; flush; within halo. fruit or fruit segments; straight; not plicate; not twisted; Hilum halo color darker than testa. Lens discernible; asymmetrical; linear or ovate (rarely); when asymmetri- less than 0.5 mm or equal to or greater than 0.5 mm in cal with both sutures nearly straight; not inflated; terete; length; 0.5 mm long; with margins straight or curved; with beak; coiled (to almost so); with solid beak the oblong; not in groove of raphe; confluent with hilum; same color and texture as fruit; short tapered at apex; mounded; dissimilar color from testa; darker than testa; apex aligned with longitudinal axis of fruit; rounded at reddish brown; within halo. Lens halo color darker than base; base aligned with longitudinal axis of fruit; with testa. Endosperm thin; covering entire embryo; adnate the apex and base uniform in texture; coriaceous; seed to embryo or testa. Cotyledons smooth; both outer faces chambers externally invisible. Fruit margin not con- convex; both the same thickness; both more or less of stricted; without sulcus; plain. Fruit wings absent. Fruit equal length; not folded; margin entire 180 degrees nonstipitate. Fruit with all layers dehiscing; splitting from base of radicle; similar at apex; partially conceal- along sutures. Dehiscence of valves along both sutures; ing radicle; entire over radicle; without lobes; with the apical and down; active; with valves twisting. Replum interface division terminating at base of radicle; without invisible. Epicarp dull; monochrome or multicolored; margins recessed; tan or white; inner face flat; glabrous streaked; reddish brown; with black overlay; pubescent around base of radicle. Embryonic axis deflexed; and indurate or pubescent but soon deciduous; with 1 oblique to length of seed; without a joint evident type of pubescence; puberulent, villous, or tomentose; between the radicle and the cotyledons. Radicle with pubescence golden or gray; with pubescence bulbose; lobe tip straight or curved; deflexed and uniformly distributed; with simple hairs; pliable; with parallel to cotyledon length; centered between cotyle- hair bases plain; eglandular; without spines; not dons; less than 1/2 length of cotyledons. Plumule smooth; with elevated features; not veined; not tubercu- rudimentary; glabrous. late; shagreen (bases of deciduous hairs); not exfoliating; without cracks. Mesocarp trace; surface not veined; Distribution: Sino-Himalayan region to Siberia. 1-layered; without balsamic vesicles; without fibers; vitreous (tiny separate dots); chartaceous. Endocarp Notes: Yakovlev (1980) reported on the two species in the dull; monochrome; tan; smooth or fibrous (somewhat former U.S.S.R. Mabberley (1997) cited 14 species in between seeds); nonseptate (though thinly hairy the genus, which is the number used here. between seeds) or subseptate; with septa thin (tissue paper-like), flexible; with septa eglandular; coriaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 3Ð17; length transverse to or parallel with fruit length; touching; in 2 or more series. Funiculus measured; 0.5Ð1 mm long; of 1 length only; filiform; straight or hooked. Aril absent.

Seed 1Ð1.5 ϫ 1Ð1.5 ϫ 0.4Ð0.7 mm; not overgrown; not angular or angular; asymmetrical; mitaform, reniform, Gueldenstaedtia: G. himalaica J.G. Baker (CÐE), G. spp. oblong, or quadrangular (somewhat); compressed; with (AÐB). A, Fruits (ϫ 2.2); B, seeds (ϫ 7.7); CÐD, testa surface smooth; with visible radicle and cotyledon (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

859 Genus: Galega C. Linnaeus absent. Raphe not visible. Hilum visible or partially concealed; concealed by funicular remnant; with faboid Phylogenetic Number: 16.20. split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.4Ð0.5 Tribe: Galegeae. mm long; with curved outline; circular; marginal according to radicle tip; recessed; within halo. Hilum Subtribe: Galeginae. halo color lighter than testa. Lens discernible; equal to or greater than 0.5 mm in length; up to 1 mm long; with Species Studied—Species in Genus: 5 spp.—6 spp. margins straight or curved; oblong or triangular; not in groove of raphe; confluent with or adjacent to hilum; Fruit a legume; unilocular; 3Ð3.5 ϫ 0.2Ð0.9 ϫ 0.12Ð0.13 0.3 mm from hilum; mounded; dissimilar color from cm; with persistent or deciduous calyx; with calyx testa; darker than testa; reddish brown or orange; not shorter than fruit; without orifice formed by curving of within corona, halo, or rim. Endosperm thin; covering fruit or fruit segments; straight or curved (slightly); not entire embryo; adnate to testa. Cotyledons smooth; both plicate; not twisted; symmetrical; linear; not inflated; outer faces convex; both the same thickness; both more compressed; with beak; straight; with solid beak the or less of equal length; not folded; margin entire 180 same color and texture as fruit; tapered at apex; apex degrees from base of radicle; similar at apex; not aligned with longitudinal axis of fruit; tapered at base; concealing radicle; entire over radicle; without lobes; base aligned with longitudinal axis of fruit; with the with the interface division terminating at base of apex and base uniform in texture; coriaceous; seed radicle; without margins recessed; yellow or brown chambers externally invisible or visible; with the raised (reddish); inner face flat; glabrous around base of seed chambers not torulose. Fruit margin not con- radicle. Embryonic axis deflexed; oblique to length of stricted or constricted; constricted along both margins; seed; with a joint evident between the radicle and the without sulcus; plain. Fruit wings absent. Fruit cotyledons. Radicle bulbose; lobe tip straight; deflexed nonstipitate. Fruit with all layers dehiscing; splitting and parallel to cotyledon length; centered between along sutures. Dehiscence of valves along both sutures; cotyledons; less than 1/2 length of cotyledons. Plumule active; with valves twisting. Replum invisible. Epicarp moderately developed or rudimentary; glabrous. dull; monochrome; dark reddish brown; glabrous or pubescent and indurate; with 1 type of pubescence; Distribution: Eastern Africa and Eurasia. puberulent; with pubescence gray; with pubescence uniformly distributed; with glandular hairs; pliable; Notes: Gillett (1963b) studied the Galega species in the with hair bases plain; glandular; with glandular hairs; mountains of eastern tropical Africa. Pandey and Jha without spines; not smooth; with elevated features; (1988) described the testa of G. officinalis using the longitudinally veined relative to fruit length; not SEM. tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; nonseptate; coriaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð6; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 1 mm long; of 1 length only; filiform; curved. Aril dry; rim-aril; reddish brown.

Seed 3.5Ð4.5 ϫ 1.8Ð2 ϫ 1Ð1.5 mm; not overgrown; not angular; asymmetrical; reniform or linear; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; Galega: G. officinalis C. Linnaeus (CÐE), G. spp. (AÐB). A, monochrome; orange or yellow; glabrous; smooth; Fruits (ϫ 1.7); B, seeds (ϫ 7.6); CÐD, testa (ϫ 50, coriaceous. Fracture lines absent. Rim absent. Wings ϫ 1000); E, embryos (ϫ 6).

861 Genus: Alhagi F. Gagnepain thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to Phylogenetic Number: 16.21. mesocarp and epicarp; entire. Seeds 1Ð9; length parallel with fruit length; neither overlapping nor touching; in 1 Tribe: Galegeae. series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril dry; rim-aril; blackish brown Subtribe: Alhagiinae. or black.

Species Studied—Species in Genus: 3 spp.—3 spp. Seed 2Ð3 ϫ 0.8Ð2.3 ϫ 1.3 mm; overgrown, 1 seed filling entire fruit cavity; angular; asymmetrical; reniform or Fruit a legume or loment (or loment segment, only tardily quadrangular; compressed; with surface smooth; with separating: see Notes); unilocular; 0.8Ð2 ϫ 0.2Ð0.4 ϫ visible radicle and cotyledon lobes; with or without 0.2Ð0.4 cm; with persistent calyx; with calyx shorter external groove between radicle and cotyledon lobes; than fruit; without orifice formed by curving of fruit or with external groove between radicle and cotyledon fruit segments; straight or curved (or slightly curved); lobes same color as testa; without hilar sinus; without not plicate; not twisted; asymmetrical; linear or umbo on seed faces. Testa not adhering to endocarp; moniliform; when asymmetrical with both sutures dull; not modified by a bloom; colored; monochrome or unequally curved, 1 straight and 1 curved suture, or mottled; with frequent mottles; brown to blackish both sutures nearly straight; narrowing in several brown, tan, yellow, green, or black; with black (to places, resembling Desmodium (9.09) fruit; not inflated; purplish black) overlay; glabrous; smooth; coriaceous. terete; with beak; straight or declined; with solid beak Fracture lines absent. Rim absent. Wings absent. Raphe the same color and texture as fruit; rounded at apex; not visible. Hilum visible; with faboid split; with the apex aligned or oblique with longitudinal axis of fruit; lips of the faboid split the same color as the rest of the short tapered at base; base aligned with longitudinal hilum; punctiform; between cotyledon and radicle lobe; axis of fruit; with the apex and base uniform in texture; recessed; not within corona, halo, or rim. Lens discern- chartaceous or coriaceous; seed chambers externally ible; less than 0.5 mm or equal to or greater than 0.5 visible; with the raised seed chambers not torulose. mm in length; 0.5 mm long; with margins straight; Fruit margin constricted (some constrictions may be diamond-shaped or irregular; not in groove of raphe; well developed) or not constricted; constricted along confluent with or adjacent to hilum; 0.1 mm from both margins or constricted only on 1 margin; without hilum; mounded; dissimilar color from testa; darker sulcus; plain. Fruit wings absent. Fruit substipitate or than testa; blackish brown or black; not within corona, stipitate (because of aborting of lowest seed chamber); halo, or rim. Endosperm thick; covering entire embryo; with the stipe up to 5 mm long. Fruit indehiscent. adnate to embryo. Cotyledons smooth; both outer faces Replum invisible. Loment indehiscent. Loment seg- convex; both the same thickness; both more or less of ments widest across seed area; oblong. Epicarp dull; equal length; not folded; margin entire 180 degrees monochrome; reddish to blackish brown or tan; from base of radicle; similar at apex; not concealing glabrous, pubescent and indurate (especially between radicle; entire over radicle; without lobes; with the seed chamber), or pubescent but soon deciduous; with interface division terminating at base of radicle; without hairs appressed or erect; with 1 type of pubescence; margins recessed; white; inner face flat; glabrous sericeous (to sparingly); with pubescence gray; with around base of radicle. Embryonic axis deflexed; pubescence uniformly distributed; with simple hairs; oblique to length of seed; without a joint evident pliable; with hair bases plain; glandular or eglandular; between the radicle and the cotyledons. Radicle linear; with glandular dots (reddish-brown); without spines; deflexed and parallel to cotyledon length; centered smooth or not smooth; with elevated features; not between cotyledons; less than 1/2 length of cotyledons. veined; not tuberculate; warty or wrinkled; not exfoliat- Plumule rudimentary; glabrous. ing; without cracks. Mesocarp thick; surface not veined; 2-layered; without balsamic vesicles; without fibers; Distribution: Mediterranean region to Nepal. with solid layer over vitreous layer; coriaceous. Endocarp dull; monochrome or mottled; white; with Notes: Polhill (1981h) noted that Alhagi is usually placed in mottling (dark); with brown (reddish-brown (vitreous the Hedysareae, “but as discovered there the flowers layer of mesocarp)) overlay; smooth; septate; with septa and fruits do not agree well with the current narrow

862 circumscription of that tribe.” Yakovlev (1979) discussed the species A. maurorum as occurring in the former U.S.S.R. Hutchinson (1964) reported “embryo covered by a fleshy membrane,” which is the endosperm adnate to the embryo. The fruit of Alhagi is technically and functionally an indehiscent legume and not a loment, notwithstanding the fact that the fruits may tardily and irregularly fracture at the isthmuses, which occur between seed chambers. The fruit has been described accurately as lomentoid or lomentaceous, and, recognizing this situation, we have scored it both ways. Pandey and Jha (1988) described the testa of A. maurorum using the SEM.

863 Alhagi: A. maurorum F.C. Medikus (CÐE), A. spp. (AÐB). A, Fruits and fruit segment (ϫ 3.9); B, seeds (ϫ 7.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 12).

865 Genus: Glycyrrhiza C. Linnaeus exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð8; length parallel with fruit length; Phylogenetic Number: 16.22. neither overlapping nor touching; in 1 series. Funiculus measured; up to 1 mm long; of 1 length only; thick; Tribe: Galegeae. straight or curved. Aril absent.

Subtribe: Glycyrrhizinae. Seed 2.5Ð7.5 ϫ 2Ð6 ϫ 1.8Ð3.3 mm; not overgrown; not angular; asymmetrical; reniform, oblong, D-shaped, or Species Studied—Species in Genus: 9 spp.—ca. 20 spp. circular; compressed; with surface smooth; with visible radicle and cotyledon lobes; with or without external Fruit a legume; unilocular; 1.2Ð3.5 ϫ 0.5Ð1.2 ϫ 0.2Ð1 cm; groove between radicle and cotyledon lobes; with with persistent or deciduous calyx; with calyx shorter external groove between radicle and cotyledon lobes than fruit; without orifice formed by curving of fruit or same color as testa; without hilar sinus; without umbo fruit segments; slightly curved, curved, or 0.5-coiled; on seed faces. Testa not adhering to endocarp; dull; not not plicate or plicate (somewhat folded: wavy); not modified by a bloom; colored; monochrome or mottled twisted or twisted; symmetrical; elliptic, linear, or and streaked; with frequent mottles; with frequent moniliform; not inflated; compressed or terete; with streaks; yellowish, reddish, or greenish brown or green beak; straight; with solid beak the same color and (yellowish or brownish); with black overlay; glabrous; texture as fruit; short tapered at apex; apex aligned with smooth; coriaceous. Fracture lines reticulate. Rim longitudinal axis of fruit; short tapered at base; base absent. Wings absent. Raphe visible or not visible; from aligned with longitudinal axis of fruit; with the apex hilum through lens to base of seed and terminating; not and base uniform in texture; coriaceous; seed chambers bifurcating; darker than testa; black; flush. Hilum externally visible; with the raised seed chambers visible or partially concealed; concealed by funicular torulose or not torulose. Fruit margin constricted along remnant; with faboid split; with the lips of the faboid both margins; without sulcus; plain or embellished; split the same color as the rest of the hilum; larger than with spines (with straight or hooked apices). Fruit punctiform; 0.8 mm long; with curved outline; circular; wings absent. Fruit stipitate or nonstipitate; with the between cotyledon and radicle lobe; recessed; within stipe up to 30 mm long. Fruit with all layers dehiscing; rim or within halo. Hilum halo color of or darker splitting along suture. Dehiscence of valves along 1 (slightly) than testa. Hilum rim color of or darker suture; apical and down (or just apical); passive. (slightly) than testa. Lens discernible; less than 0.5 mm Replum invisible. Epicarp dull; monochrome; reddish or equal to or greater than 0.5 mm in length; 0.2Ð1 mm brown; glabrous or pubescent and indurate; with 1 type long; with margins curved; circular; not in groove of of pubescence; puberulent; with pubescence golden raphe; adjacent to hilum; 0.5Ð1.5 mm from hilum; (and gland tipped); with pubescence uniformly distrib- mounded; same color as testa; not within corona, halo, uted; with simple hairs; pliable; with hair bases plain; or rim. Endosperm thick; covering entire embryo; glandular; with glandular hairs or dots; with spines adnate to embryo. Cotyledons smooth; both outer faces (with straight or hooked or plain or glandular tips) or convex; both the same thickness; both more or less of without spines; with spines persistent or broken off and equal length; not folded; margin entire 180 degrees their bases evident; with spines same color as the rest of from base of radicle; similar at apex; partially conceal- the fruit; not smooth; with elevated or recessed features; ing radicle; entire over radicle; without lobes; with the veined or not veined; reticulately veined or longitudi- interface division terminating at base of radicle; without nally veined relative to fruit length; not tuberculate; margins recessed; tan; inner face flat; glabrous around tuberculate (bases of spines); pitted (centers of spine base of radicle. Embryonic axis deflexed; oblique to bases); not exfoliating; without cracks. Mesocarp thin; length of seed; without a joint evident between the surface not veined; 1-layered; without balsamic radicle and the cotyledons. Radicle bulbose; lobe tip vesicles; without fibers; vitreous; coriaceous. Endocarp straight; deflexed and parallel to cotyledon width; dull; monochrome or mottled; tan; with brown (reddish) centered between cotyledons; less than 1/2 length of overlay; smooth; septate, subseptate, or nonseptate; cotyledons. Plumule rudimentary; glabrous. with septa thin (tissue paper-like), flexible or thicker than paper, firm; with septa eglandular; coriaceous; not Distribution: North America, temperate South America, Eurasia to Australia.

866 867 Glycyrrhiza: G. lepidota F.T. Pursh (CÐE), G. spp. (AÐB). A, Fruits (ϫ 1.4); B, seeds (ϫ 3.1); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

Genus: Streblorrhiza S.F.L. Endlicher Notes: Hutchinson (1964) established the tribe Carmichaelieae, and Polhill (1981i, 1994a,b) accepted Phylogenetic Number: 17.01. it. Heenan (1995, 1998c), using unpublished nuclear ribosomal DNA ITS data, concluded that Tribe: Carmichaelieae. “Carmichaelia (17.05) is nested within [the] ‘Astragalean clade’ of Galegeae” and is the sister group Species Studied—Species in Genus: 1 sp.—1 sp. of Clianthus (16.01). He therefore supported the proposal of Sanderson and Wojciechowski (1996) that Fruit a legume; unilocular; 4.5 ϫ 1.8 ϫ 0.3 cm; with Carmichaelieae should not be recognized at tribal level deciduous corolla; with deciduous calyx; without but rather should be included in Galegeae (16). Heenan orifice formed by curving of fruit or fruit segments; (1998c) also carried out cladistic analyses of straight; not plicate; not twisted; asymmetrical or Carmichaelia (17.05), Chordospartium (17.03), symmetrical; oblong or falcate (obliquely subfalcate); Corollospartium (17.04), and Notospartium (17.02) when asymmetrical with both sutures parallelly to using morphological and anatomical characters. Heenan unequally curved; not inflated; compressed; without (1998a,c) concluded that “Carmichaelia is paraphyletic beak; tapered at apex; apex right-angled with longitudi- with Chordospartium, Corollospartium, and nal axis of fruit; tapered at base; base aligned with Notospartium excluded,” and reunited them with longitudinal axis of fruit; with the apex and base Carmichaelia. Polhill (1981i) noted that Streblorrhiza uniform in texture; coriaceous; seed chambers exter- is extinct. Hutchinson (1964) stated that the “embryonic nally visible (barely). Fruit margin not constricted; axis is twice folded.” We were able to study only one without sulcus; plain. Fruit stipitate; with the stipe 6 mature pod and one mature seed. mm long. Replum invisible. Epicarp dull; monochrome; dark brown; pubescent but soon deciduous to glabrous (with age); with 1 type of pubescence; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; transversely veined relative to fruit length; not tuberculate; not exfoliating; without cracks. Seeds length parallel with fruit length; neither overlapping nor touching; in 1 series. Aril absent.

Seed 6 ϫ 4.5 ϫ 3 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; brown (dark reddish); glabrous; smooth; coriaceous (assumed). Fracture lines absent. Rim absent. Raphe from hilum to lens; not bifurcating; color of testa; recessed. Hilum fully concealed or partially concealed; concealed by radicle lobe or wing; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 1 mm long; with curved outline; elliptic; marginal according to radicle tip; recessed; not within corona, halo, or rim. Lens discernible; less than 0.5 mm in length; with margins curved; circular; in groove of raphe; adjacent to hilum; 1.5 mm from hilum; recessed; Streblorrhiza: S. speciosa S.F.L. Endlicher (AÐE). A, Fruit dissimilar color from testa; darker than testa; black; not (ϫ 1.2); B, seed (ϫ 3.3); CÐD, testa (ϫ 50, ϫ 1000); E, within corona, halo, or rim. Radicle lobe tip curved. embryos (ϫ 4.7).

871 Genus: Notospartium J.D. Hooker mounded; similar color as testa; darker than testa; brownish red; not within corona, halo, or rim. En- Phylogenetic Number: 17.02. dosperm thick; covering entire embryo; adnate to testa. Cotyledons smooth; both outer faces convex; both the Tribe: Carmichaelieae. same thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; Species Studied—Species in Genus: 3 spp.—3 spp. similar at apex; not concealing radicle; entire over radicle; without lobes; with the interface division Fruit a legume; unilocular; 2.5Ð2.8 ϫ 0.5Ð0.6 ϫ 0.17Ð0.3 terminating at base of radicle; without margins re- cm; with deciduous corolla; with deciduous calyx; cessed; tan; inner face flat; glabrous around base of without orifice formed by curving of fruit or fruit radicle. Embryonic axis deflexed; oblique to length of segments; straight or curved (slightly); not plicate; not seed. Radicle linear; lobe tip straight; with 360-degree twisted; asymmetrical; linear (nearly); when asym- turn; not centered between cotyledons (radicle outside 1 metrical with both sutures parallelly curved to 1 straight cotyledon and inside other, therefore junctions for each and 1 curved suture; not inflated; compressed; without cotyledon different); exceeding length of cotyledons. beak; rounded at apex; apex aligned with longitudinal Plumule rudimentary; glabrous. axis of fruit; tapered at base; base aligned with longitudinal axis of fruit; with the apex and base uniform in Distribution: New Zealand (South Island). texture; coriaceous; seed chambers externally visible; with the raised seed chambers not torulose. Fruit Notes: Heenan (1998a,c) carried out cladistic analyses of margin not constricted; without sulcus; plain. Fruit Carmichaelia (17.05), Chordospartium (17.03), wings absent. Fruit substipitate. Fruit indehiscent. Corollospartium (17.04), and Notospartium using Replum invisible. Epicarp dull; monochrome; tan; morphological and anatomical characters and con- glabrous; eglandular; without spines; not smooth; with cluded that this genus should be merged with elevated features; transversely veined relative to fruit Carmichaelia. length; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; cobwebby; subseptate; with septa thin (tissue paper-like), flexible; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 3Ð5; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 1 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 2.5Ð3.5 ϫ 2Ð2.8 ϫ 1.2Ð1.3 mm; not overgrown; angular or not angular; asymmetrical; mitaform; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled and streaked; with frequent mottles; with frequent streaks; brownish red; with black overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe from hilum to lens; not bifurcating; darker than testa; raised. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; marginal according to radicle tip; recessed; not within corona, halo, or rim. Lens discernible; less than 0.5 mm Notospartium: N. carmichaeliae J.D. Hooker (CÐE), N. spp. in length; with margins curved; punctiform; not in (AÐB). A, Fruits and valve (ϫ 2); B, seeds (ϫ 5.3); groove of raphe; adjacent to hilum; 1 mm from hilum; CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

873 Genus: Chordospartium T.F. Cheeseman red; not within corona, halo, or rim. Endosperm thick; covering entire embryo; adnate to testa. Cotyledons Phylogenetic Number: 17.03. smooth; both outer faces convex; both the same thickness; both more or less of equal length; not folded; Tribe: Carmichaelieae. margin entire 180 degrees from base of radicle; differing at apex (1 concealed by overarching radicle Species Studied—Species in Genus: 1 sp.—1 sp. and other auriculate and concealing radicle); not concealing radicle; 1 cotyledon scooped out to accom- Fruit a legume; bilocular; 0.4Ð0.5 ϫ 0.2Ð0.26 ϫ 0.13Ð0.18 modate plicate radicle and other cotyledon entire; cm; with deciduous corolla; with deciduous calyx; without lobes; with the interface division terminating at without orifice formed by curving of fruit or fruit base of radicle; brownish red; inner face flat; glabrous segments; straight; not plicate; not twisted; asymmetri- around base of radicle. Embryonic axis deflexed; cal; when asymmetrical with 1 straight and 1 curved plicate to length of seed. Radicle linear; lobe tip curved; suture; widest near middle or D-shaped; not inflated; with 360 degree turn; centered between cotyledons; compressed; without beak; short tapered at apex; apex exceeding length of cotyledons. Plumule rudimentary; aligned with longitudinal axis of fruit or right-angled glabrous. with longitudinal axis of fruit; short tapered at base; base aligned with longitudinal axis of fruit; with the Distribution: New Zealand (South Island). apex and base uniform in texture; coriaceous; seed chambers externally visible. Fruit margin not con- Notes: Heenan (1998a,c) carried out cladistic analyses of stricted; without sulcus; plain. Fruit wings absent. Fruit Carmichaelia (17.05), Chordospartium, substipitate. Fruit indehiscent. Replum invisible. Corallospartium (17.04), and Notospartium (17.02) Epicarp dull; multicolored; mottled; brown; with brown using morphological and anatomical characters and overlay; pubescent and indurate; with 1 type of pubes- concluded that this genus should be merged with cence; pilose (crinkled); with pubescence gray; with Carmichaelia. pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; transversely veined relative to fruit length; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp. Seed 1; length parallel with fruit length. Funiculus measured; to 1 mm long; filiform; straight. Aril absent.

Seed 2.5Ð3 ϫ 1.8Ð2 ϫ 1Ð1.2 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; red; glabrous; not smooth; with recessed features; pitted with small separate pits; coriaceous. Fracture lines absent. Rim absent. Raphe from hilum to lens; darker than testa (slightly); flush. Hilum visible; without faboid split; punctiform; marginal according to radicle tip; recessed; not within corona, halo, or rim. Lens discernible; less than 0.5 mm in length; with margins curved; punctiform; not in groove of raphe; adjacent to Chordospartium: C. stevensonii T.F. Cheeseman (AÐE). A, hilum; 0.7 mm from hilum; mounded; same color as Fruits (ϫ 3.9); B, seeds (ϫ 4); CÐD, testa (ϫ 50, testa or dissimilar color from testa; darker than testa; ϫ 1000); E, embryos (ϫ 8).

875 Genus: Corallospartium J.B. Armstrong testa; darker than testa; brown; not within corona, halo, or rim. Endosperm thin; covering entire embryo; adnate Phylogenetic Number: 17.04. to testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal Tribe: Carmichaelieae. length; not folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing radicle; entire Species Studied—Species in Genus: 1 sp.—1 sp. over radicle; without lobes; with the interface division terminating at base of radicle; without margins re- Fruit a legume; unilocular; 0.5Ð0.6 ϫ 0.25Ð0.3 ϫ 0.2Ð0.22 cessed; reddish tan; inner face flat; glabrous around cm; with deciduous corolla; with deciduous calyx; base of radicle. Embryonic axis oblique; perpendicular without orifice formed by curving of fruit or fruit to length of seed. Radicle linear; lobe tip curved; with segments; straight; not plicate; not twisted; asymmetri- 180 degree turn; not centered between cotyledons cal; when asymmetrical with 1 straight and 1 curved (radicle outside 1 cotyledon and inside other, therefore suture; widest near middle or D-shaped; not inflated; junctions for each cotyledon different); exceeding compressed; without beak; short tapered at apex; apex length of cotyledons. Plumule rudimentary; glabrous. aligned with longitudinal axis of fruit; rounded at base; base aligned with longitudinal axis of fruit; with the Distribution: New Zealand (South Island). apex and base uniform in texture; coriaceous; seed chambers externally invisible. Fruit margin not con- Notes: Heenan (1998a,c) carried out cladistic analyses of stricted; without sulcus; plain. Fruit wings absent. Fruit Carmichaelia (17.05), Chordospartium (17.03), nonstipitate. Fruit with all layers dehiscing; splitting Corallospartium, and Notospartium (17.02) using along sutures. Dehiscence of valves along both sutures; morphological and anatomical characters and con- apical and down (assumed); passive. Replum invisible. cluded that Corallospartium should be merged with Epicarp dull; monochrome; gray; pubescent and Carmichaelia. Cheeseman (1906) noted that indurate; with 1 type of pubescence; velutinous; with Corallospartium “is technically separated from pubescence gray; with pubescence uniformly distrib- Carmichaelia by the 2-valved pod without a persistent uted; with simple hairs; pliable; with hair bases plain; replum.” eglandular; without spines; not smooth; with elevated features; transversely veined relative to fruit length; not tuberculate; not exfoliating; without cracks. Mesocarp absent or present (if present, very thin). Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1(Ð2); length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; less than 1 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 1.9Ð2.1 ϫ 1.4Ð1.6 ϫ 1.4Ð1.6 mm; overgrown, 1 seed filling entire fruit cavity; not angular; symmetrical; circular; terete; with visible radicle and cotyledon lobes (barely); without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; mottled and streaked; with frequent mottles; with frequent streaks; brown or green; with green or black (bluish) overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; without faboid split; punctiform; marginal according to radicle tip; recessed. Lens Corallospartium: C. crassicaule (J.D. Hooker) J.B. discernible; less than 0.5 mm in length; with margins Armstrong (AÐE). A, Fruits (with calyx and floral parts) curved; punctiform; not in groove of raphe; adjacent to (ϫ 4.1); B, seeds (ϫ 7.3); CÐD, testa (ϫ 50, ϫ 1000); hilum; 0.5 mm from hilum; flush; dissimilar color from E, embryos (ϫ 20).

877 Genus: Carmichaelia R. Brown infrequent streaks; black (to bluish-black), brown, green (pale), orange (reddish), red, tan, white (to bluish- Phylogenetic Number: 17.05. white), yellow, or blue; with black overlay (to bluish- black); glabrous; smooth; coriaceous. Fracture lines Tribe: Carmichaelieae. absent. Rim absent. Raphe not visible. Hilum visible or fully concealed; concealed by funicular remnant; with Species Studied—Species in Genus: 7 spp.—18 spp. faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; up Fruit a legume; unilocular; 0.5Ð5 ϫ 0.15Ð0.8 ϫ 0.1Ð0.15 to 7 mm long; with straight outline; oblong; marginal cm; with deciduous corolla; with deciduous calyx; according to radicle tip or between cotyledon and without orifice formed by curving of fruit or fruit radicle lobe; recessed; not within corona, halo, or rim. segments; straight; not plicate; not twisted; asymmetri- Lens discernible or not discernible; less than 0.5 mm in cal or symmetrical; subcircular, oblong, ovate, elliptic, length; with margins curved; punctiform; not in groove linear, or falcate; when asymmetrical with both sutures of raphe; adjacent to hilum; 0.5 mm from hilum; parallelly curved; not inflated; compressed to terete; mounded or flush; same color as testa; darker than with beak; straight; with solid beak the same color and testa; black or red (dish); not within corona, halo, or texture as fruit; abruptly long acuminate at apex; apex rim. Endosperm thick; covering entire embryo; adnate aligned with longitudinal axis of fruit; rounded at base; to testa or embryo. Cotyledons smooth; both outer faces base aligned with longitudinal axis of fruit; with the convex; both the same thickness; both more or less of apex and base uniform in texture; ligneous; seed equal length; not folded; margin entire 180 degrees chambers externally visible; with the raised seed from base of radicle; differing at apex (1 concealed by chambers not torulose. Fruit margin not constricted; overarching radicle and other auriculate and concealing without sulcus; embellished; with thickened sutural radicle); not concealing radicle; entire over radicle; areas. Fruit wings absent. Fruit substipitate. Fruit with without lobes; with the interface division terminating at all layers dehiscing or indehiscent; splitting along base of radicle; white or red; inner face flat; glabrous suture. Dehiscence of valves along 1 suture; basal and around base of radicle. Embryonic axis deflexed; up; active; with valves twisting. Replum visible or perpendicular to length of seed. Radicle bulbose; lobe invisible. Epicarp dull; monochrome; black, brown, or tip curved; with 180-degree or 90-degree turn; centered tan; glabrous; without spines; not smooth; with between cotyledons; 1/2 to nearly length of or equaling elevated features; transversely veined relative to fruit length of ctyledons. Plumule rudimentary; glabrous. length; not tuberculate; wrinkled; not exfoliating; without cracks. Mesocarp thin; 1-layered; without Distribution: Seventeen species in New Zealand and one (C. balsamic vesicles; without fibers; solid; ligneous. exsul H.F. Mueller) on Lord Howe Island. Endocarp dull; monochrome; brown; fibrous; subseptate or nonseptate; with septa thin (tissue paper- Notes: Simpson (1945) revised the genus recognizing eight like), flexible; with the septa eglandular; chartaceous; subgenera and 41 species based on whether pods not exfoliating; remaining fused to epicarp. Seeds 1Ð dehisce or not and their dehiscence mechanisms and 12; length parallel with fruit length; overlapping or presented a key to the subgenera essentially using pod neither overlapping nor touching; in 1 series. Funiculus characters. Allan (1961) used pod characters as primary measured; 0.3Ð2.5 mm long; of 1 length only; filiform divisions in his New Zealand species key. Heenan or thick; straight, S-curved, hooked, or plicate. Aril (1995, 1996, 1998a,c) revised the genus for New dry; rim-aril; brown. Zealand and carried out cladistic analyses of Carmichaelia and related genera. He concluded that Seed 2Ð4.5 ϫ 2Ð3 ϫ 1.5Ð3 mm; not overgrown; angular “Carmichaelia is paraphyletic if Chordospartium (because of pressure of adjacent seeds in fruit) or not (17.03), Corallospartium (17.04), and Notospartium angular; symmetrical or asymmetrical; irregular, (17.02) are excluded.” Therefore Heenan (1998a) mitaform, oblong, or reniform; terete or compressed; merged the four genera and created new combinations with or without visible radicle and cotyledon lobes; in Carmichaelia for the species of Chordospartium, without umbo on seed faces. Testa present or absent; Corallospartium, and Notospartium. We have chosen to not adhering to endocarp; glossy; not modified by a use Polhill’s (1981i, 1994b) generic delimitations bloom; colored; monochrome or mottled and streaked; pending the expression of a general consensus of with frequent or infrequent mottles; with frequent or opinion concerning Heenan’s work by taxonomic

878 botanists. Although we studied seeds and fruits of 7 species, the literature cited above provided brief descriptions of seeds and fruits for all species. Heenan’s (1995, 1996, 1998a) species count was used.

879 Carmichaelia: C. uniflora T. Kirk (CÐE), C. spp. (AÐB). A, Fruits (ϫ 4.4); B, seeds (ϫ 4.9); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

881 Hedysareae (18.01Ð18.07) recessed; within halo. Hilum halo color darker than testa. Lens discernible or not discernible; less than 0.5 Genus: Eversmannia A.A. von Bunge mm in length; with margins curved; circular; not in groove of raphe; adjacent to hilum; 0.5 mm from hilum; Phylogenetic Number: 18.01. mounded; similar color as testa; darker than testa; reddish brown; not within corona, halo, or rim. En- Tribe: Hedysareae. dosperm thin; covering entire embryo; adnate to embryo. Cotyledons smooth; both outer faces convex; Species Studied—Species in Genus: 1 sp.—1 sp. both the same thickness; both more or less of equal length; not folded; margin entire 180 degrees from base Fruit a legume (apparently breaking between seed chambers of radicle; similar at apex; not concealing radicle; entire (like a loment)); unilocular; 3 ϫ 0.6Ð0.7 ϫ 0.2 cm; over radicle; without lobes; with the interface division with deciduous corolla; with deciduous calyx; without terminating at base of radicle; without margins re- orifice formed by curving of fruit or fruit segments; cessed; tan; inner face flat; glabrous around base of slightly curved; plicate (loosely) or not plicate; not radicle. Embryonic axis deflexed; oblique to length of twisted; asymmetrical or symmetrical; linear; when seed; without a joint evident between the radicle and asymmetrical with both sutures nearly straight; not the cotyledons. Radicle linear; lobe tip straight; inflated; compressed; without beak; short tapered or deflexed and parallel to cotyledon length; centered rounded at apex; apex aligned with longitudinal axis of between cotyledons; more than 1/2 length of cotyle- fruit; short tapered or rounded at base; base aligned dons. Plumule rudimentary; glabrous. with longitudinal axis of fruit; with the apex and base uniform in texture; coriaceous; seed chambers exter- Distribution: Russia and Iran from the Caspian to Tian Shan nally visible; with the raised seed chambers torulose. (border of former U.S.S.R. and China). Fruit margin constricted along both margins; without sulcus; plain. Fruit wings absent. Fruit nonstipitate. Notes: Polhill (1981j) primarily used fruit characters for his Fruit indehiscent (probably). Replum visible (“articles” key to genera of this tribe. The fruits of E. subspinosa falling free from replum). Epicarp dull; monochrome; (formerly E. hedysaroides A.A. von Bunge) function as reddish brown; glabrous; eglandular; without spines; a loment, though they are not true loments. The fruit not smooth; with elevated features; reticulately veined; tissue between the seed chambers is thin and easily not tuberculate; not exfoliating; without cracks. broken (but by no regular separation) into one-seeded Mesocarp thin; surface not veined; 1-layered; without fruit segments leaving the replum. balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth; nonseptate (seed fits into fruit like a ball in a socket and walls of fruit form pseudosepta); chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð4; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; triangular; straight. Aril absent.

Seed 3Ð3.7 ϫ 3 ϫ 1.3Ð1.5 mm; not overgrown; not angular; asymmetrical; D-shaped, mitaform, or rhombic; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings Eversmannia: E. subspinosa (F.E.L. von Fischer ex A.-P. de absent. Raphe not visible. Hilum fully concealed; Candolle) B.A. Fedtschenko (AÐE). A, Fruits (ϫ 2.4); concealed by funicular remnant; without faboid split; B, seeds (ϫ 5.5); CÐD, testa (ϫ 50, ϫ 1000); E, punctiform; between cotyledon and radicle lobe; embryos (ϫ 15).

883 Genus: Hedysarum C. Linnaeus series. Funiculus less than 0.5 mm long or measured; up to 3 mm long; of 1 length only; filiform; straight. Aril Phylogenetic Number: 18.02. absent.

Tribe: Hedysareae. Seed 2.8Ð4.5 ϫ 2Ð4 ϫ 1Ð2 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with Species Studied—Species in Genus: 50 spp.—ca. 100 spp. surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and Fruit a loment (with 2 to several joints); 2Ð4 ϫ 0.3Ð1 ϫ cotyledon lobes; with external groove between radicle 0.15Ð0.25 cm; with deciduous corolla; with persistent and cotyledon lobes same color as testa; with shallow calyx; with calyx shorter than fruit; without orifice hilar sinus; without umbo on seed faces. Testa not formed by curving of fruit or fruit segments; straight; adhering to endocarp; glossy or dull; not modified by a not plicate; not twisted; symmetrical or asymmetrical; bloom; colored; monochrome; reddish or greenish linear, elliptic, or ovate; when asymmetrical with both brown, tan (to greenish), yellow, or cream; glabrous; sutures parallelly curved; not inflated; compressed or smooth; coriaceous. Fracture lines absent. Rim absent. terete; without beak; short tapered at apex; apex aligned Wings absent. Raphe not visible. Hilum partially with longitudinal axis of fruit; rounded or short tapered concealed; concealed by funicular remnant or concealed at base; base aligned with longitudinal axis of fruit; by wing; with faboid split; with the lips of the faboid with the apex and base uniform in texture; coriaceous, split the same color as the rest of the hilum; punctiform; membranous, or ligneous; seed chambers externally between cotyledon and radicle lobe; flush; within rim. visible; with the raised seed chambers torulose. Fruit Hilum rim color darker than testa or color of testa. Lens margin constricted along both margins; without or with discernible or not discernible (because may be color of sulcus; plain or embellished; with spines (with usually testa); less than 0.5 mm in length; with margins straight curved or occasionally straight apex), prickles, or wing. or with margins curved; circular or oblong; not in Fruit wing present or absent; 1; 0.5Ð3 mm wide; groove of raphe; adjacent to or confluent with hilum; sutural. Fruit nonstipitate or substipitate. Replum 0.4 mm from hilum; mounded; dissimilar color from or invisible. Loment indehiscent. Loment segments same color as testa; darker than testa; black; not within (articles) inconspicuous; 3.5Ð16 mm long; widest corona, halo, or rim. Endosperm thin; covering entire across seed area; with all essentially similar in shape or embryo; adnate to testa. Cotyledons smooth; both outer lower 1 different shape than middle ones; elliptic to faces convex; both the same thickness; both more or oblong, D-shaped, quadrangular, or circular. Epicarp less of equal length; not folded; margin entire 180 dull; monochrome; brown; glabrous, glabrate, or degrees from base of radicle; similar at apex; not pubescent but soon deciduous; with hairs appressed or concealing radicle; entire over radicle; without lobes; erect; with 1 type of pubescence; pilose or tomentose; with the interface division terminating at base of with pubescence gray; with pubescence uniformly radicle; without margins recessed; white or yellow; distributed; with simple hairs; pliable; with hair bases inner face flat; glabrous around base of radicle. Embry- plain; eglandular; with or without spines; with spines onic axis deflexed; oblique to length of seed; without a persistent or broken off and their bases evident; with joint evident between the radicle and the cotyledons. spines same color as the rest of the fruit; not smooth; Radicle linear; lobe tip hooked or straight; deflexed and with elevated features; reticulately veined; tuberculate parallel to cotyledon length; centered between cotyle- (if apicies of spines removed); with solid tubercles on dons; less than 1/2 or 1/2 to nearly length of cotyledons. each valve; rugose, wrinkled, tuberculate, or muricate; Plumule rudimentary or moderately developed; not exfoliating; without cracks. Mesocarp thin; surface glabrous. not veined; 1-layered; without balsamic vesicles; without fibers; solid; ligneous (sub) or coriaceous. Distribution: Europe, Mediterranean region, Asia, and North Endocarp dull; monochrome; tan; smooth; septate; with America. septa thicker than paper, firm; with the septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds (1Ð)2Ð8; length parallel Hedysarum: H. coronarium C. Linnaeus (CÐE), H. spp. (AÐ with fruit length; neither overlapping nor touching; in 1 B). A, Fruits and articles (ϫ 1.4); B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

885 Genus: Taverniera A.-P. de Candolle adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome; brown (to greenish); Phylogenetic Number: 18.03. glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Tribe: Hedysareae. partially concealed; concealed by funicular remnant; without faboid split; punctiform; between cotyledon Species Studied—Species in Genus: 9 spp.—ca. 10 spp. and radicle lobe; recessed; within rim. Hilum rim color of testa or color darker than testa. Lens discernible; less Fruit a loment (with 2Ð4 joints); 1.25Ð1.5 ϫ 0.3Ð0.7 ϫ 0.5Ð than 0.5 mm in length or equal to or greater than 0.5 0.7 cm; with deciduous or persistent corolla; with mm in length; up to 0.5 mm long; with margins straight standard; with persistent or deciduous calyx; with calyx or curved; circular or oblong; not in groove of raphe; shorter than fruit; without orifice formed by curving of confluent with or adjacent to hilum; 0.1Ð0.5 mm from fruit or fruit segments; straight; not plicate; not twisted; hilum; mounded (light colored mound within darker asymmetrical or symmetrical; oblong; when asymmetri- area); similar color as or dissimilar color from testa; cal with 1 straight and 1 curved suture (or nearly darker than testa; black or brown (greenish); not within straight); narrowest near middle, B-shaped; not inflated; corona, halo, or rim. Endosperm thin; covering entire compressed; without beak; rounded or short tapered at embryo; adnate to testa. Cotyledons smooth; both outer apex; apex aligned or oblique with longitudinal axis of faces convex; both the same thickness; both more or fruit; rounded at base; base aligned with longitudinal less of equal length; not folded; margin entire 180 axis of fruit; with the apex and base uniform in texture; degrees from base of radicle; similar at apex; not coriaceous; seed chambers externally visible; with the concealing radicle; entire over radicle; without lobes; raised seed chambers torulose. Fruit margin constricted with the interface division terminating at base of along both margins; without sulcus; embellished or radicle; without margins recessed; green; inner face plain; with prickles or spines. Fruit wings absent. Fruit flat; glabrous around base of radicle. Embryonic axis substipitate. Fruit indehiscent. Replum invisible. deflexed; oblique to length of seed; without a joint Loment indehiscent. Loment segments (articles) evident between the radicle and the cotyledons. Radicle inconspicuous; widest across seed area; with all linear; deflexed and parallel to cotyledon length; essentially similar in shape; D-shaped or circular. centered between cotyledons; less than 1/2 length of Epicarp dull or glossy; monochrome; dark to reddish cotyledons. Plumule rudimentary; glabrous. brown or tan; pubescent and indurate; with hairs erect or appressed; with 1 type of pubescence; pilose; with Distribution: Horn of Africa and Socotra through Middle pubescence gray; with pubescence uniformly distrib- East to western India. uted; with simple hairs; pliable; with hair bases plain; eglandular; with spines (either glabrous or hairy and Notes: Polhill (1981j) noted that the genus has “about ten with straight or hooked apices) or without spines; with species (several undescribed).” spines persistent; with spines same color as the rest of the fruit; not smooth; with elevated features; veined or not veined; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth; septate; with septa thicker than paper, firm; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 2Ð3; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long or measured; 0.5 mm long; of 1 length only; filiform; curved. Aril absent.

Seed 1.5Ð2.5 ϫ 1.1Ð2 ϫ 0.5Ð1.1 mm; not overgrown; not Taverniera: T. lappacea (P. Forsskål) A.-P. de Candolle angular; asymmetrical; mitaform or reniform; com- (CÐE), T. spp. (AÐB). A, Fruits and articles (ϫ 3.6); B, pressed; with surface smooth; with visible radicle and seeds (ϫ 8); CÐD, testa (ϫ 50, ϫ 1000); E, embryos cotyledon lobes; without umbo on seed faces. Testa not (ϫ 8).

887 Genus: Stracheya G. Bentham not bifurcating; darker than testa; black; flush. Hilum fully concealed; concealed by funicular remnant; Phylogenetic Number: 18.04. without faboid split; punctiform; between cotyledon and radicle lobe; recessed; not within corona, halo, or Tribe: Hedysareae. rim. Lens discernible or not discernible; less than 0.5 mm in length; with margins curved; circular; not in Species Studied—Species in Genus: 1 sp.—1 sp. groove of raphe; adjacent to hilum; 0.8 mm from hilum; flush; dissimilar color from testa; darker than testa; Fruit a legume; unilocular; 2Ð3 ϫ 0.6Ð0.7 (excluding black; not within corona, halo, or rim. Endosperm thin; spines) ϫ 0.2 (excluding spines) cm; with deciduous covering entire embryo; adnate to testa. Cotyledons corolla; with persistent calyx; with calyx shorter than smooth; both outer faces convex; both the same fruit; without orifice formed by curving of fruit or fruit thickness; both more or less of equal length; not folded; segments; slightly curved or straight; not plicate; not margin entire 180 degrees from base of radicle; similar twisted; asymmetrical or symmetrical; linear or oblong; at apex; not concealing radicle; entire over radicle; when asymmetrical with both sutures nearly straight; without lobes; with the interface division terminating at not inflated; compressed (but spines along sutures and base of radicle; without margins recessed; tan; inner midvalve (4 rows of spines)); without beak; truncate at face flat; glabrous around base of radicle. Embryonic apex; apex aligned with longitudinal axis of fruit; short axis deflexed; oblique to length of seed; without a joint tapered at base; base aligned with longitudinal axis of evident between the radicle and the cotyledons. Radicle fruit; with the apex and base uniform in texture; linear; lobe tip straight; deflexed and parallel to coriaceous; seed chambers externally invisible. Fruit cotyledon length; centered between cotyledons; less margin not constricted; without sulcus; embellished; than 1/2 length of cotyledons. Plumule rudimentary; with spines. Fruit wings absent. Fruit nonstipitate. Fruit glabrous. indehiscent. Replum invisible. Epicarp dull; monochrome; reddish brown; pubescent and indurate; with Distribution: Kashmir, Tibet, Sikkim, and Nepal. hairs appressed; with 1 type of pubescence; with pubescence gray; with pubescence uniformly distrib- Notes: Little is known about this monotypic genus, and uted; with simple hairs; pliable; with hair bases plain; Viswanathan (1995) revised it. The fruits appear to be eglandular; with spines (along sutures and midvalves (4 indehiscent and are not loments. The fruits are defi- rows) on fruit); with spines persistent; with spines same nitely echinate-dentate and have spines in four rows. color as the rest of the fruit; not smooth; with elevated Only pressed fruits were seen. features; not veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface uniformly veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth; nonseptate (but seeds seated like a ball and socket in fruit); chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 5; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 2Ð2.7 ϫ 1.3Ð2 ϫ 0.7 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified or modified by a bloom; colored; monochrome; dark reddish brown; glabrous; not smooth; with elevated Stracheya: S. tibetica G. Bentham (AÐE). A, Fruits (ϫ 2.9); features; shagreen; coriaceous. Fracture lines absent. B, seeds (ϫ 7.4); CÐD, testa (ϫ 50, ϫ 1000); E, Rim absent. Wings absent. Raphe from hilum to lens; embryos (ϫ 10).

889 Genus: Onobrychis P. Miller Seed 2Ð5 ϫ 2.6Ð3 ϫ 2.1Ð2.2 mm; not overgrown; not angular; asymmetrical; reniform or oblong; com- Phylogenetic Number: 18.06. pressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle Tribe: Hedysareae. and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without Species Studied—Species in Genus: 66 spp.—ca. 130 spp. hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a Fruit a legume; unilocular; 0.4Ð1.5 ϫ 0.15Ð1 cm; with bloom; colored; monochrome or mottled; brown, deciduous corolla; with persistent or deciduous calyx; yellow, cream, or black; with black overlay; glabrous; with calyx shorter than, longer than, or equal in length smooth; coriaceous. Fracture lines absent. Rim absent. to fruit; without or with orifice formed by curving of Wings absent. Raphe not visible or visible; from hilum fruit or fruit segments; straight, curved, or 1-coiled to lens; not bifurcating; darker than testa or color of (nearly); not plicate; not twisted; asymmetrical; testa; reddish brown; recessed. Hilum partially con- circular, reniform, or samaroid; when asymmetrical cealed; concealed by funicular remnant; with faboid with 1 straight and 1 curved suture, both sutures split; with the lips of the faboid split the same color as parallelly curved or both unequally curved; widest near the rest of the hilum; punctiform; between cotyledon middle or D-shaped; not inflated; compressed or terete; and radicle lobe; flush; within halo. Hilum halo color without or with beak; hooked (at apex), straight, or darker than testa. Lens discernible; less than 0.5 mm in declined; with solid beak the same color and texture as length; with margins curved; circular or elliptic; not in fruit (3Ð10 mm long); short tapered at apex; apex or in groove of raphe; adjacent to or confluent with aligned with or almost reaching longitudinal axis of hilum; 1 mm from hilum; mounded; dissimilar color fruit; tapered or short tapered at base; base aligned with from or similar color as testa; darker than testa; reddish longitudinal axis of fruit; with the apex and base brown or black; not within corona, halo, or rim. uniform in texture; coriaceous; seed chambers exter- Endosperm thin; covering entire embryo; adnate to nally visible. Fruit margin constricted or not con- testa. Cotyledons smooth; both outer faces convex; both stricted; constricted along both margins; without sulcus; the same thickness; both more or less of equal length; embellished or plain; with spines, wing (entire or cut), not folded; margin entire 180 degrees from base of prickles, or fringe. Fruit wing absent or present; 1; up to radicle; similar at apex; not concealing radicle; entire 3 mm wide; samaroid, sutural, or continuous wing over radicle; without lobes; with the interface division around fruit; on both sutures. Fruit nonstipitate or terminating at base of radicle; without margins re- substipitate. Fruit indehiscent. Replum invisible. Fruit a cessed; tan; inner face flat; glabrous around base of nutlet; an intact article; entire. Epicarp dull; mono- radicle. Embryonic axis deflexed; oblique to length of chrome; brown or tan; glabrous or pubescent and seed; without a joint evident between the radicle and indurate; with hairs erect or appressed; with 1 type of the cotyledons. Radicle linear; lobe tip straight; pubescence; minutely puberulent or tomentose; with deflexed and parallel to cotyledon length; centered pubescence golden or gray; with pubescence uniformly between cotyledons; less to more than 1/2 length of distributed; with simple hairs; pliable; with hair bases cotyledons. Plumule rudimentary or moderately plain; eglandular; with or without spines; with spines developed; glabrous. persistent or broken off and their bases evident; with spines same color as the rest of the fruit; not smooth; Distribution: Europe, Mediterranean region, western and with elevated or recessed features; veined or not veined; central Asia, and Ethiopia. reticulately veined (and with or without spines and reticulum can be removed); not tuberculate or tubercu- Notes: Cooper and Carleton (1968) edited a symposium late; with solid tubercles on each valve; pitted (broad proceeding dealing with the agronomic aspects of pits); not exfoliating; without cracks. Mesocarp absent. sainfoin (O. viciifolia J.A. Scopali). Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seed 1; length parallel with fruit length. Funiculus less than 0.5 mm long; filiform; straight. Aril Onobrychis: O. melanotricha P.E. Boissier (CÐE), O. spp. absent. (AÐB). A, Fruits (ϫ 1.3); B, seeds (ϫ 3.7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

891 Genus: Ebenus C. Linnaeus with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; Phylogenetic Number: 18.07. marginal according to radicle tip or between cotyledon and radicle lobe; flush; not within corona, halo, or rim Tribe: Hedysareae. or within halo. Hilum halo color darker than testa. Lens not discernible or discernible; less than 0.5 mm in Species Studied—Species in Genus: 7 spp.—20 spp. length; with margins curved; circular; not in groove of raphe; adjacent to hilum; 0.2 mm from hilum; Fruit a legume; unilocular; 0.45Ð0.6 ϫ 0.3Ð0.4 ϫ 0.13Ð0.2 mounded; similar color as testa; darker than testa; cm; with deciduous corolla; with persistent calyx; with reddish brown or tan; not within corona, halo, or rim. calyx longer than fruit; without orifice formed by Endosperm thin; covering entire embryo; adnate to curving of fruit or fruit segments; straight; not plicate; testa. Cotyledons smooth; both outer faces convex; both not twisted; symmetrical; oblong or obovate; not the same thickness; both more or less of equal length; inflated; compressed; without beak; rounded at apex; not folded; margin entire 180 degrees from base of apex aligned with longitudinal axis of fruit; tapered at radicle; similar at apex; not concealing radicle; entire base; base aligned with longitudinal axis of fruit; with over radicle; without lobes; with the interface division the apex and base uniform in texture; membranous or terminating at base of radicle; without margins re- fragile, thinner than chartaceous like Trifolium; seed cessed; yellow or tan; inner face flat; glabrous around chambers externally invisible. Fruit margin not con- base of radicle. Embryonic axis deflexed; oblique to stricted; without sulcus; plain. Fruit nonstipitate. Fruit length of seed; without a joint evident between the indehiscent (rupturing at base). Replum invisible. radicle and the cotyledons. Radicle linear; lobe tip Epicarp dull; monochrome; tan; pubescent and indurate; straight; deflexed and parallel to cotyledon length; with 1 type of pubescence; tomentose; with pubescence centered between cotyledons; less than 1/2 length of golden; with apical pubescence different from basal cotyledons. Plumule moderately developed; glabrous. pubescence; with apical 3/4 tomentose and basal 1/4 glabrous or with apical 1/2 crinkly tomentose and basal Distribution: Mediterranean region. 1/2 densely villose with straight hairs; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1(Ð2); length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 2.5Ð3.2 ϫ 1.8Ð2 ϫ 1.6Ð2 mm; not overgrown; not angular; asymmetrical; mitaform or ovate; terete; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled; with frequent mottles; reddish brown or tan; with black overlay (minute mottles may be present on tan testae); glabrous; smooth or not smooth; with recessed features; faintly striate or Ebenus: E. cretica C. Linnaeus (DÐF), E. spp. (AÐC). A, concaved; coriaceous. Fracture lines absent. Rim Fruits in calyx (ϫ 2.1); B, fruits with calyx removed absent. Wings absent. Raphe not visible. Hilum (ϫ 7); C, seeds (ϫ 5.7); DÐE, testa (ϫ 50, ϫ 1000); F, partially concealed; concealed by funicular remnant; embryos (ϫ 8).

893 Fabeae (19.01Ð19.05) or thicker than paper, firm; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds Genus: Vicia C. Linnaeus 1Ð16; length parallel with fruit length (without regard to hilum position in V. faba); neither overlapping nor Phylogenetic Number: 19.01. touching or touching; in 1 series. Funiculus measured; up to 2 mm long; of 1 length only; thick or flattened; Tribe: Fabeae. straight or S-curved. Aril absent or present (not a true aril, but an expanded funiculus); fleshy; expanded Species Studied—Species in Genus: 85 spp.—166 spp. funiculus; covering less than 1/2 of seed; tan or white.

Fruit a legume; unilocular; 0.6Ð2.5 ϫ 0.2Ð3 ϫ 0.2Ð3 cm; Seed 1.5Ð30.5 (without regard to hilum position in V. faba) with persistent or deciduous calyx; with calyx shorter ϫ 1.4Ð17 ϫ 1.3Ð9 mm; not overgrown; not angular or than fruit; without orifice formed by curving of fruit or angular; asymmetrical or symmetrical; circular, oblong, fruit segments; straight to curved (to slightly curved); ovate, quadrangular, or triangular; terete or compressed; not plicate; not twisted; asymmetrical; linear, falcate, or without visible radicle and cotyledon lobes; with umbo rhombic; when asymmetrical with both sutures nearly on seed faces. Testa not adhering to endocarp; dull, straight; inflated (V. faba) or not inflated; compressed, glossy, or velvet; not modified or modified by a bloom; terete, or flattened; without or with beak; straight; with colored; monochrome or mottled and streaked; with solid beak the same color and texture as fruit; short frequent mottles; with frequent streaks; brown (includ- tapered to tapered to rounded at apex; apex aligned or ing most shades and combinations with other colors), oblique (slightly) with longitudinal axis of fruit; short tan (reddish, greenish), or black (purplish); with black, tapered to tapered to long tapered to rounded at base; brown (including most shades and combinations with base oblique or aligned with longitudinal axis of fruit; other colors), or red overlay; glabrous; smooth or not with the apex and base uniform in texture; coriaceous smooth; with elevated features; tuberculate; osseous or or leathery; seed chambers externally invisible or coriaceous. Fracture lines absent. Rim absent. Wings visible; with the raised seed chambers not torulose or absent. Raphe not visible. Hilum visible, partially torulose. Fruit margin not constricted or constricted; concealed, or fully concealed; concealed by funiculus, slightly constricted along both margins to constricted funicular remnant, or aril; with faboid split; with the along both margins or slightly constricted only on 1 lips of the faboid split lighter colored than the rest of margin; without sulcus; plain. Fruit wings absent. Fruit the hilum and therefore conspicuous or the same color nonstipitate or substipitate. Fruit with all layers as the rest of the hilum; larger than punctiform or dehiscing; splitting along sutures. Dehiscence of valves punctiform; 0.3Ð12 mm long; with curved, angular, or along both sutures; apical and down; active; with valves straight outline; circular, wedge-shaped, linear, or twisting (tightly or loosely). Replum invisible. Epicarp oblong; marginal according to radicle tip; flush; not dull; monochrome; brown (various shades and in within corona, halo, or rim or within halo. Hilum halo combination with other colors), tan, or black; glabrous color lighter than testa. Lens discernible or not discern- or pubescent and indurate to pubescent but soon ible; equal to or greater than 0.5 mm or less than 0.5 deciduous; with 1 type of pubescence; puberulent or mm in length; 0.7Ð4 mm long; with margins straight or pilose; with pubescence gray or golden; with simple or curved; irregular, linear, or circular; not in groove of glandular hairs; pliable; with hair bases swollen or raphe; adjacent to, confluent with, 180 degrees from, or plain; glandular; with glandular hairs; without spines; 270 degrees from hilum; up to 13 mm from hilum not smooth; with elevated or recessed features; reticu- (longest distances in V. faba); mounded; dissimilar lately veined; not tuberculate or tuberculate; with solid color from, similar color to, or same color as testa; tubercles (widely scattered) on each valve; pusticulate darker than testa; black, brown, or tan; not within (indurate bases of hairs) or tuberculate (scattered and corona, halo, or rim. Endosperm thin; covering entire rarely); occasionally punctate; not exfoliating; without embryo; adnate to testa (enclosing radicle in sheath). cracks. Mesocarp thin; surface not veined; 1-layered; Cotyledons smooth; both outer faces convex; both the without balsamic vesicles; without fibers; solid; same thickness; both more or less of equal length; not coriaceous. Endocarp dull; monochrome; tan; smooth; folded; margin entire 180 degrees from base of radicle; nonseptate (New World spp.) or septate (some Old similar at apex; not concealing radicle; split over World spp.); with septa thin (tissue paper-like), flexible radicle; with lobes; with lobes not touching; with the

894 interface division terminating at base of radicle; without margins recessed; white or yellow; inner face flat; glabrous around base of radicle. Embryonic axis deflexed; oblique to length of seed. Radicle triangular; deflexed and parallel to cotyledon length or width; centered between cotyledons; less than 1/2 length of cotyledons. Plumule well developed; glabrous.

Distribution: North and South America, Europe, Canary Islands, North and East Africa, Asia, and Hawaii.

Notes: This tribe has traditionally been called Vicieae. Article 19.4 of the International Code of Botanical Nomenclature (Greuter et al. 1994) states that, “The name of any subdivision of a family that includes the type of the adopted, legitimate name of the family to which it is assigned is to be based on the generic name equivalent to the type.” Faba P. Miller is the type of Fabaceae and is synonymous with Vicia. Therefore because Faba is included in this tribe, the tribe must be called Fabeae. After performing a cladistic analysis using morphological characters, including internal seed morphology, Endo and Ohashi (1997) proposed that Cicereae (20) and Fabeae (Vicieae) formed a monophyletic group whose sister group is Trifolieae (21). Kupicha (1976) treated, for the first time, Vicia on a worldwide basis. She recognized two subgenera and twenty-two sections, and some fruit and seed characters have value as sectional characters. Maxted (1993, 1995) revised Vicia subgen. Vicia and proposed nine sections and nine series. He distinguished Vicia sect. Hypechusa (F.G.C. Alefeld) P.F.A. Acherson & K.O.R.P.P. Graebner by its seed characteristics and revised the section with Colin Douglas (Maxted and Douglas 1997), recognizing two series, 14 species, and six subspecies in the section. Potokina (1997) revised the V. sativa aggregate for the former U.S.S.R. Like Lens (19.03) and Pisum (19.04), a few species of Vicia, such as V. faba and V. ervilia (C. Linnaeus) C.L. von Willdenow, appeared “in the Neolithic and Bronze Age cultures in the Near East and Europe” (Zohary and Hopf 1973). Hermann (1960) treated the genus in the United States; Gunn (1979) treated the genus in Mexico and Central America; Lassetter and Gunn (1979) monographed V. menziesii C.P.J. Sprengel, a native

species of the island of Hawaii; and Bastos and Miotto (1996) revised the native Brazilian^ species of Vicia. Gunn (1970a,b, 1971) and Z ertová (1962) studied the seed morphology of Vicia, and Gunn (1968) provided a key and diagrams for the seeds of 100 species of Vicia. Vicia sativa subsp. amphicarpa (C. Linnaeus) J.A. Battandier produces both aerial and subterranean fruits.

895 Vicia: V. faba C. Linnaeus (CÐE), V. sativa C. Linnaeus (F), V. spp. (AÐB). A, Fruits (intact and dehisced) (ϫ 1.1); B, seeds (ϫ 2.4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 1); F, embryos (ϫ 5).

897 Genus: Lathyrus C. Linnaeus Seed 1.5Ð10 ϫ 1.5Ð10 ϫ 1.3Ð7 mm; not overgrown; not angular or angular; symmetrical or asymmetrical (with Phylogenetic Number: 19.02. or without dents because of adjacent seed pressures); oblong, circular (to subcircular), rectangular, quadran- Tribe: Fabeae. gular, triangular (compressed), or irregular (angular); terete or compressed (rarely); without visible radicle Species Studied—Species in Genus: 75 spp.—161 spp. and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull or glossy; not modified Fruit a legume; unilocular; 2.5Ð10 ϫ 0.2Ð1 ϫ 0.15Ð0.4 cm; or modified by a bloom; colored; monochrome or with deciduous calyx; without orifice formed by mottled; with frequent mottles; brown (diverse shades curving of fruit or fruit segments; straight or curved and in combinations with other colors), black (with or (slightly); not plicate; not twisted; asymmetrical or without purplish tinge), or tan; with brown (blackish) symmetrical; linear to rhombic (linear); when asym- overlay; glabrous; smooth or not smooth; with elevated metrical with both sutures nearly straight; not inflated; features; tuberculate (either aligned or not in rows or flattened or compressed; without beak; short tapered at united into short ridges or not) or wrinkled (faintly); apex; apex aligned or oblique with longitudinal axis of coriaceous. Fracture lines absent. Rim absent. Wings fruit; long tapered or short tapered at base; base aligned absent. Raphe not visible or visible; from hilum through or oblique with longitudinal axis of fruit; with the apex lens and base of seed to point opposite hilum; not and base uniform in texture; coriaceous, membranous, bifurcating; darker than testa; black (color of mottles). chartaceous, or ligneous (L. lentiformis U. Plitman); Hilum visible, partially concealed, or fully concealed; seed chambers externally visible (faintly to visible) or concealed by funiculus or funicular remnant; with invisible; with the raised seed chambers not torulose or faboid split; with the lips of the faboid split lighter torulose. Fruit margin not constricted; without sulcus; colored than the rest of the hilum and therefore con- plain or embellished. Fruit wings absent or present; spicuous; larger than punctiform; up to 5 mm long; with sutural or valvular (occasionally); on both valves; on curved, angular, or straight outline; circular, elliptic, both sutures. Fruit nonstipitate or stipitate (infrequent, wedge-shaped, oblong, or linear; subapical or marginal see L. setifolius C. Linnaeus). Fruit with all layers according to radicle tip; flush or recessed; not within dehiscing or indehiscent (L. lentiformis); splitting along corona, halo, or rim (though some species with light sutures. Dehiscence of valves along both sutures; apical colored seeds with mottles concentrated around hilum and down; active; with valves twisting. Replum somewhat like a necklace). Lens discernible; less than invisible. Epicarp dull; monochrome; reddish brown or 0.5 mm or equal to or greater than 0.5 mm in length; up tan; glabrous or pubescent and indurate; with hairs erect to 1.5 mm long; with margins straight or curved; or appressed; with 1 type of pubescence; pilose; with oblong, linear, triangular, rhombic, or circular; not in pubescence gray; with simple hairs; pliable; with hair groove of raphe; adjacent to or confluent with hilum; bases swollen or plain; glandular; with glandular hairs touching to 1 mm from hilum; mounded or recessed; (L. cassius P.E. Boissier); without spines; not smooth; dissimilar color from testa; darker than testa; reddish with elevated features; reticulately veined; not tubercu- brown, black, or red; not within corona, halo, or rim. late; papillose (L. hirsutus C. Linnaeus) or glandular Endosperm absent or present; thin; covering entire dotted (section Orobus (C. Linnaeus) J.G. Baker); not embryo; adnate to testa (but enclosing radicle). Cotyle- exfoliating; without cracks. Mesocarp thin; surface not dons not smooth; both outer faces convex; both the veined; 1-layered; without balsamic vesicles; without same thickness or 1 thicker than the other; both more or fibers; solid; coriaceous or chartaceous. Endocarp dull; less of equal length; not folded; margin entire 180 monochrome; tan; smooth; nonseptate or septate; with degrees from base of radicle; similar at apex; not septa thin (tissue paper-like), flexible; with septa concealing radicle; entire over radicle; without lobes; eglandular; chartaceous; not exfoliating; remaining with the interface division terminating at base of fused to mesocarp and epicarp; entire. Seeds 1Ð25; radicle; without margins recessed; tan; inner face flat; length parallel with or transverse to fruit length; glabrous around base of radicle. Embryonic axis touching; in 1 series. Funiculus less than 0.5 mm long; deflexed; oblique to length of seed. Radicle linear or of 1 length only; thick; straight. Aril absent (funiculus bulbose; deflexed and parallel to cotyledon width; may be expanded and remain over hilum and is not an centered between cotyledons; less than 1/2 length of aril). cotyledons. Plumule well developed; glabrous.

898 Distribution: North and South America; Europe; North, East, and tropical Africa; and Asia.

Notes: Kupicha (1983) classified the Old World and New World species and recognized 13 sections. Asmussen and Liston (1998) conducted cladistic analyses of 42 Lathyrus species from 12 of Kupicha’s sections using chloroplast DNA. Their results indicated that Lathyrus should be organized in 6Ð8 sections, not 13. Hitchcock (1952) monographed the North American species. Bässler (1966, 1971, 1981) studied some of the Old World species, and Hung-Pin (1984) extended his studies to China. Butler (1986) investigated the testa of Lathyrus using the scanning electron microscope, and her results are published in the Kaul and Combes (1986) book “Lathyrus and Lathyrism.” Lathyrism is an animal expression of the toxic components of Lathyrus species, especially L. sativus. Several species produce aerial and subterranean fruits; for example, L. amphicarpos C. Linnaeus and L. ciliolatus K. H. Rechinger. Kupicha discussed the fruits of Lathyrus and illustrated their external variations in a full-page plate. The drawing of L. lentiformis fruit is especially interesting (fig. 5, k). Kupicha also recorded the relative hilum lengths for the 13 sections. The number of species is from Asmussen and Liston (1998).

899 Lathyrus: L. sativus C. Linnaeus (CÐE), L. spp. (AÐB). A, Fruits (intact and dehisced) (ϫ 1); B, seeds (ϫ 2.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

901 Genus: Lens P. Miller spicuous; larger than punctiform; 1Ð1.5 mm long; with straight outline; linear; marginal according to radicle Phylogenetic Number: 19.03. tip; flush; not within corona, halo, or rim. Lens discernible; equal to or greater than 0.5 mm to less than 0.5 Tribe: Fabeae. mm in length; 0.3Ð0.7 mm long; with margins straight or curved; linear or circular; not in groove of raphe; Species Studied—Species in Genus: 2 spp.—6 spp. adjacent to hilum; 0.5Ð1.2 mm from hilum; mounded; (Maxted, personal communication, 1998). dissimilar color from testa; darker than testa; black; not within corona, halo, or rim. Endosperm trace; restricted Fruit a legume; unilocular; 0.9Ð1.6 ϫ 0.4Ð1.2 ϫ 0.15Ð0.2 to region of embryo (around radicle area); adnate to cm; with persistent or deciduous calyx; with calyx testa. Cotyledons smooth; both outer faces convex; both shorter than fruit; without orifice formed by curving of the same thickness; both more or less of equal length; fruit or fruit segments; straight; not plicate; not twisted; not folded; margin entire 180 degrees from base of asymmetrical; oblong; when asymmetrical with both radicle; similar at apex; not concealing radicle; split sutures nearly straight; not inflated; compressed; over radicle; with lobes; with lobes not touching; with without beak; short tapered at apex; apex aligned with the interface division terminating at base of radicle; longitudinal axis of fruit; short tapered at base; base without margins recessed; white, yellow, or orange; oblique with longitudinal axis of fruit; with the apex inner face flat; glabrous around base of radicle. Embry- and base uniform in texture; chartaceous; seed cham- onic axis deflexed; oblique to length of seed. Radicle bers externally visible or invisible; with the raised seed linear; deflexed and parallel to cotyledon length (more chambers not torulose. Fruit margin not constricted; or less); centered between cotyledons; less than 1/2 without sulcus; plain. Fruit wings absent. Fruit length of cotyledons. Plumule well developed; gla- substipitate. Fruit with all layers dehiscing; splitting brous. along sutures. Dehiscence of valves along both sutures; apical and down; active or passive; with valves enroll- Distribution: Mediterranean region, West and Central Asia, ing or twisting. Replum invisible. Epicarp dull; mono- Ethiopia, and Zaire, and now widely cultivated. chrome; reddish brown or tan; glabrous or pubescent and indurate; with 1 type of pubescence; puberulent; Notes: The number of species in this genus has been open to with pubescence gray; pliable; with hair bases plain; question for many years. Most authors recognized the eglandular; without spines; not smooth; with elevated cultivated and commercially important lentil, L. features; reticulately veined; not tuberculate; not culinaris F.C. Medikus. In the past, at least three wild exfoliating; without cracks. Mesocarp thin; surface not species have been recognized. Ladizinsky et al. (1984), veined; 1-layered; without balsamic vesicles; without using extensive crossability and cytological studies, fibers; solid; chartaceous. Endocarp dull; monochrome; recognized one additional species, L. nigricans (F.A. tan; smooth; nonseptate; chartaceous; not exfoliating; Marchall von Bieberstein) D.A. Godman. The two other remaining fused to mesocarp and epicarp; entire. Seeds often recognized species of Lens, L. ervoides and L. (1Ð)2; length parallel with fruit length; neither overlap- orientalis (P.E. Boissier) H. Handel-Mazzetti, are now ping nor touching; in 1 series. Funiculus measured; 1 L. nigricans subsp. ervoides (G. de Brignoli di mm long; of 1 length only; filiform; straight. Aril Brunnhoff) G. Ladizinsky and L. culinaris subsp. absent (funiculus dilated). orientalis (P.E. Boissier) B.S. Williams, respectively. The fifth species, L. montbretii (F.E.L. von Fischer & Seed 3Ð8 ϫ 3Ð8 ϫ 1.5Ð3 mm; not overgrown; not angular; C.A.A. von Meyer) P.H. Davis & U. Plitmann, had symmetrical; circular; compressed; without visible been assigned to Lens (Barulina 1930, Davis and radicle and cotyledon lobes; without umbo on seed Plitmann 1970), but Ladizinsky and Sakar (1982) faces. Testa not adhering to endocarp; dull; colored; provided morphological and karyological data suggest- monochrome or mottled and streaked (mainly streaks); ing that this species be returned to Vicia (19.01) where with frequent mottles; with frequent streaks; reddish it was originally placed as V. montbretii (Hoffman et al. brown or tan (cream); with black overlay; glabrous; 1986). Ladizinsky (1997) described a new species of smooth; coriaceous. Fracture lines absent. Rim absent. Lens from southeastern Turkey, L. tomentosus G. Wings absent. Raphe not visible. Hilum visible; with Ladizinsky, using evidence from morphological, faboid split; with the lips of the faboid split lighter crossing, and cpDNA studies. Our species count of five colored than the rest of the hilum and therefore con- follows the count of Maxted (personal communication,

902 1998) for this genus. The history of L. culinaris extends as far back as agriculture itself. Like Pisum (19.04), Lens has been closely associated with grain cultivation in the Near East (Zohary and Hopf 1973).

903 Lens: L. ervoides (G. de Brignoli di Brunnhoff) L. Grande (CÐE), L. spp. (AÐB). A, Fruits (intact and dehisced) (ϫ 3.9); B, seeds (ϫ 4.9); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

905 Genus: Pisum C. Linnaeus split; with the lips of the faboid split the same color as the rest of the hilum or lighter colored than the rest of Phylogenetic Number: 19.04. the hilum and therefore conspicuous; larger than punctiform; 1.6Ð2.1 mm long; with curved outline; Tribe: Fabeae. elliptic; marginal according to radicle tip; flush; not within corona, halo, or rim. Lens discernible; equal to Species Studied—Species in Genus: 2 spp.—2 spp. or greater than 0.5 mm in length; 1Ð2 mm long; with margins curved; circular or elliptic; not in groove of Fruit a legume; unilocular; 3Ð12 ϫ 0.8Ð2.5 ϫ 0.25Ð0.5 cm; raphe; adjacent to hilum; 1.5Ð2 mm from hilum; with persistent calyx; with calyx shorter than fruit; mounded; dissimilar color from testa; darker than testa; without orifice formed by curving of fruit or fruit black, brown, or tan; not within corona, halo, or rim. segments; straight or curved (slightly); not plicate; not Endosperm thin; covering entire embryo (and forming twisted; asymmetrical; oblong or linear; when asym- sheath around radicle); adnate to testa. Cotyledons metrical with both sutures nearly straight or both smooth; both outer faces convex; both the same sutures parallelly curved; not inflated or inflated; terete thickness; both more or less of equal length; not folded; or compressed; without beak; short tapered at apex; margin entire 180 degrees from base of radicle; similar apex aligned with longitudinal axis of fruit; long at apex; not concealing radicle; split over radicle; with tapered at base; base aligned with longitudinal axis of lobes; with the interface division terminating at base of fruit; with the apex and base uniform in texture; radicle; without margins recessed; white or tan; inner coriaceous; seed chambers externally invisible or face flat; glabrous around base of radicle. Embryonic visible; with the raised seed chambers not torulose. axis deflexed; oblique to length of seed. Radicle Fruit margin not constricted; without sulcus; plain. Fruit bulbose; deflexed and parallel to cotyledon length; wings absent. Fruit nonstipitate. Fruit with all layers centered between cotyledons; 1/2 to nearly length of dehiscing; splitting along sutures. Dehiscence of valves cotyledons. Plumule well developed; glabrous. along both sutures; apical and down; active; with valves twisting. Replum invisible. Epicarp dull; monochrome; Distribution: Mediterranean region and now widely culti- brown, yellow, or green; glabrous; eglandular; without vated. spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Notes: The past literature on the genus Pisum is replete with Mesocarp thin; surface not veined; 1-layered; without numerous scientific names used at various taxonomic balsamic vesicles; without fibers; solid; coriaceous. levels because of the intense breeding of the pea, P. Endocarp dull; monochrome; brown; smooth; sativum, a major crop. Currently only two species of nonseptate; chartaceous; not exfoliating; remaining Pisum are recognized, and the other species is P. fulvum fused to mesocarp and epicarp; entire. Seeds up to 10; J. Sibthorp & J.E. Smith. Pisum fulvum can have both length parallel with fruit length; touching; in 1 series. aerial and subterranean fruits. Pisum sativum was Funiculus less than 0.5 mm long; of 1 length only; cultivated in “early Neolithic farming villages of the thick; straight. Aril absent (funiculus dilated at apex). Near East (7,000 to 6,000 B.C.)” (Zohary and Hopf 1973). Seed 3.7Ð9 ϫ 3.7Ð9 ϫ 3.7Ð7 mm; not overgrown; angular or not angular; symmetrical or asymmetrical; circular or irregular (angular); terete; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified or modified by a bloom; colored; monochrome or mottled and streaked; with frequent mottles; with frequent streaks; dark or dark reddish brown, tan (to greenish), red, or green; with brown (dark reddish) overlay; glabrous; smooth or not smooth; with elevated features; rugose, Pisum: P. sativum C. Linnaeus subsp. elatius (C. von Steven tuberculate (minutely), or wrinkled; chartaceous or ex F.A.M. von Bieberstein) P.F.A. Ascherson & coriaceous. Fracture lines absent. Rim absent. Wings K.O.R.P.P. Graebner (CÐE), P. spp. (AÐB). A, Fruits absent. Raphe not visible. Hilum visible or partially (ϫ 0.6); B, seeds (ϫ 3); CÐD, testa (ϫ 50, ϫ 1000); E, concealed; concealed by funicular remnant; with faboid embryos (ϫ 5).

907 Genus: Vavilovia A.A. Fedorov not in groove of raphe; adjacent to hilum; 0.7 mm from hilum; mounded; dissimilar color from testa; darker Phylogenetic Number: 19.05. than testa; black; not within corona, halo, or rim. Endosperm absent. Cotyledons smooth; both the same Tribe: Fabeae. thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; similar Species Studied—Species in Genus: 1 sp.—1 sp. at apex; partially concealing radicle; entire over radicle; without lobes; with the interface division terminating at Fruit a legume; unilocular; 1.5Ð3.5 ϫ 0.8Ð0.9 ϫ 0.8Ð0.9 base of radicle; without margins recessed; brown; inner (assumed) cm; with persistent calyx; with calyx shorter face flat; glabrous around base of radicle. Embryonic than fruit; without orifice formed by curving of fruit or axis straight; almost perpendicular to length of seed; fruit segments; straight; not plicate; not twisted; without a joint evident between the radicle and the asymmetrical; linear; when asymmetrical with both cotyledons. Radicle differentiated from cotyledon; sutures nearly straight; not inflated; terete; without triangular; lobe tip straight; straight with embryonic beak; rounded at apex; apex aligned with longitudinal axis; centered between cotyledons; less than 1/2 length axis of fruit; short tapered at base; base aligned with of cotyledons. Plumule rudimentary; glabrous. longitudinal axis of fruit; with the apex and base uniform in texture; coriaceous; seed chambers exter- Distribution: Turkey, Lebanon, Iraq, Iran, Russia, Georgia, nally invisible. Fruit margin not constricted; without and Armenia. sulcus; plain. Fruit wings absent. Fruit nonstipitate. Fruit with all layers dehiscing; splitting along sutures. Notes: Vavilovia formosa has been placed in Orobus C. Dehiscence of valves along both sutures; apical and Linnaeus (now Lathyrus, 19.02) and a segregate genus, down; active; with valves enrolling. Replum invisible. Alophotropis (H.F. Jaubert & É. Spach) A.A. Epicarp dull; monochrome; reddish brown or tan; Grossheim, now a synonym of Vavilovia. We were glabrous; eglandular; without spines; not smooth; with unable to obtain an intact fruit, and our studied seeds elevated features; reticulately veined; not tuberculate; were flattened during pressing. not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; reddish brown or tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 2Ð5; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 4Ð5 ϫ 3.3Ð4 ϫ 1.7Ð2.7 mm; not overgrown; not angular; symmetrical; oblong; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled; with frequent mottles; reddish brown; with black overlay; glabrous; smooth; chartaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible or partially concealed; concealed by funicular remnant; with faboid split; with the lips of the faboid split lighter colored than the rest of the hilum and therefore conspicuous; larger than punctiform; 0.7 mm long; with curved outline; elliptic; marginal according to radicle tip; flush; not within corona, halo, Vavilovia: V. formosa (C. von Steven) A.A. Federov (AÐE). or rim. Lens faintly discernible or not discernible; less A, Fruits (dehisced) (ϫ 3.1); B, seeds (ϫ 4); CÐD, testa than 0.5 mm in length; with margins curved; circular; (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

909 Cicereae (20.01) shagreen, tuberculate, warty, wrinkled, or echinate; coriaceous. Fracture lines absent. Rim absent. Raphe Genus: Cicer C. Linnaeus from lens to base of seed and bifurcating (C. arietinum); bifurcating at base of seed with each arm Phylogenetic Number: 20.01. going up antiraphe side, then turning (U-shaped) down and approaching bifurcation; color of or darker than Tribe: Cicereae. testa; tan, brown, or black; flush or recessed. Hilum visible; with faboid split; with the lips of the faboid Species Studied—Species in Genus: 28 spp.—43 spp. split the same color as the rest of the hilum; larger than punctiform; 0.5Ð1.7 mm long; with curved outline; Fruit a legume; unilocular; 1.8Ð3.6 ϫ 0.7Ð2 cm; with circular or elliptic; marginal according to radicle tip or deciduous calyx; without orifice formed by curving of between cotyledon and radicle lobe; recessed; within fruit or fruit segments; straight; not plicate; not twisted; rim or not within corona, halo, or rim. Hilum rim color symmetrical; elliptic to oblong to obovate to rhombic lighter than or color of testa. Lens discernible; equal to (elongate); not inflated; terete; without beak; short or greater than 0.5 mm in length; 2 mm long; with tapered at apex; apex aligned with longitudinal axis of margins straight or curved; triangular; elliptic; not in fruit; short tapered at base; base aligned with longitudi- groove of raphe; adjacent to hilum; 1.5Ð2 mm from nal axis of fruit; with the apex and base uniform in hilum; mounded or recessed; same color as or dissimi- texture; membranous; seed chambers externally lar color from testa; darker than testa; red; not within invisible. Fruit margin not constricted; without sulcus; corona, halo, or rim. Endosperm thin; covering entire plain. Fruit wings absent. Fruit nonstipitate. Fruit with embryo; adnate to testa. Cotyledons smooth or not all layers dehiscing; splitting along sutures. Dehiscence smooth; wrinkled (replicating testa surface); both outer of valves along both sutures; apical and down; active or faces convex; both the same thickness; both more or passive; with valves twisting. Replum invisible. Epicarp less of equal length; not folded; margin entire 180 dull; monochrome; brown or yellow (-brown); glabrous degrees from base of radicle; similar at apex; partially or pubescent and indurate; with 1 type of pubescence; concealing radicle; split over radicle; with lobes; with pilose; with pubescence gray; with pubescence uni- lobes touching (auriculate); with basal groin formed by formly distributed; with simple hairs; pliable; with hair lobes; with the interface division terminating at base of bases plain; glandular or eglandular; with glandular radicle; without margins recessed; white; inner face hairs; without spines; not smooth; with elevated flat; glabrous around base of radicle. Embryonic axis features; obliquely veined relative to fruit length; not straight, or deflexed; parallel or oblique to length of tuberculate; not exfoliating; without cracks. Mesocarp seed. Radicle linear; lobe tip straight or curved; straight thin; 1-layered; without balsamic vesicles; without with embryonic axis or oblique to cotyledons; centered fibers; solid; coriaceous. Endocarp glossy; mono- between cotyledons; less than 1/2 length of cotyledons. chrome; tan; smooth; nonseptate; chartaceous; not Plumule well developed; glabrous. exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð2(Ð7); touching, or neither overlapping Distribution: Turkey south to Israel and east to the nor touching; in 1 series. Funiculus measured; 1.5 mm Himalayas and Central Asia; also in Morocco, Egypt, long; of 1 length only; thick; straight. Aril absent. Ethiopia, Greece, and Crete. Cicer arietinum is widely cultivated. Seed 4Ð11 ϫ 2.5Ð9 ϫ 3Ð7 mm; overgrown, 1 seed filling entire fruit cavity; angular or not angular; asymmetri- Notes: After a cladistic analysis using morphological cal; ovate or bilobed, ramshead-shaped (typically characters, including internal seed morphology, Endo beaked); terete to quadrangular; with surface grooved; and Ohashi (1997) proposed that Cicereae and Fabeae with grooves longitudinal; with or without visible (19) form a monophyletic group whose sister group is radicle and cotyledon lobes; without umbo on seed Trifolieae (21). Van der Maesen (1972, 1987) faces. Testa not adhering to endocarp; dull; not modi- monographed Cicer. Lersten and Gunn (1981, 1982) fied by a bloom; colored or clear (C. arietinum); discussed the seed morphology and testa topography of monochrome or mottled; with frequent mottles; black, Cicer species, which supported the separation of brown, cream, gray, green, ivory, olive, orange (yellow- Cicereae from the Fabeae (19) (Werker 1997). Follow- ish), pink (gray-brown), red, tan, white, or yellow; ing traditions in the Indian subcontinent, Summerfield glabrous; smooth or not smooth; with elevated features; and Roberts (1985a) recognized two groups for the

910 guidance of plant breeders: Genotypes producing large, rounded, pale cream-colored seeds (greater than 25 g per 100 seeds) borne on tall plants (Kabuli), and genotypes producing smaller irregularly shaped, variously-colored seeds (less than 25 g per 100 seeds) borne on relatively short to prostrate plants (Desi).

911 Cicer: C. arietinum C. Linnaeus (CÐE), C. spp. (AÐB). A, Fruits (ϫ 1.6); B, seeds (ϫ 2.1); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

913 Trifolieae (21.01Ð21.06) visible radicle and cotyledon lobes; with deep hilar sinus or without hilar sinus; without umbo on seed Genus: Ononis C. Linnaeus faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome (occasionally Phylogenetic Number: 21.01. with white tubercles); dark to light reddish brown, tan, or yellow; glabrous; not smooth or smooth; with Tribe: Trifolieae. elevated or recessed features; tuberculate (some quite prominent in size and/or color) or rugose (rarely); Species Studied—Species in Genus: 27 spp.—ca. 75 spp. punctate; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe from hilum to lens; not bifurcat- Fruit a legume; unilocular; 0.8Ð1.6 ϫ 0.2Ð0.4 ϫ 0.02Ð0.04 ing; darker than testa; tan; flush. Hilum visible, fully cm; with persistent or deciduous calyx; with calyx concealed, or partially concealed; concealed by longer than, equal in length to, or shorter than fruit; funiculus or wing; with faboid split; with the lips of the without orifice formed by curving of fruit or fruit faboid split the same color as the rest of the hilum; segments; straight; not plicate; not twisted; asymmetri- punctiform; between cotyledon and radicle lobe; cal or symmetrical; linear, oblong, rhombic, or ovate recessed; within rim. Hilum rim color of testa. Lens (rarely); when asymmetrical with 1 straight and 1 discernible; less than 0.5 mm or equal to or greater than curved suture; widest near middle or D-shaped; not 0.5 mm in length; 0.5Ð1 mm long; with margins straight inflated; compressed; without beak; short tapered at or curved; linear, circular, elliptic, or oblong; not in apex; apex aligned with longitudinal axis of fruit; short groove of raphe; adjacent to hilum; 0.1Ð0.2 mm from tapered at base; base aligned with longitudinal axis of hilum; mounded; similar color as testa; darker than fruit; with the apex and base uniform in texture; testa; reddish brown, tan (reddish), or black (ish); not coriaceous; seed chambers externally invisible. Fruit within corona, halo, or rim. Endosperm thin; covering margin constricted or not constricted; slightly con- entire embryo; adnate to testa or embryo. Cotyledons stricted along both margins; without sulcus; plain. Fruit smooth; both outer faces convex; both the same wings absent. Fruit nonstipitate. Fruit with all layers thickness; both more or less of equal length; not folded; dehiscing; splitting along sutures. Dehiscence of valves margin entire 180 degrees from base of radicle; similar along both sutures; apical and down; active; with valves at apex; not concealing radicle; entire over radicle; twisting. Replum invisible. Epicarp dull; monochrome; without lobes; with the interface division terminating at tan; pubescent and indurate; with 1 type of pubescence; base of radicle; without margins recessed; yellow; inner pilose or puberulent; with pubescence golden or gray; face flat; glabrous around base of radicle. Embryonic with pubescence uniformly distributed; with glandular axis straight; oblique to length of seed. Radicle linear; and simple hairs; pliable; with hair bases plain or lobe tip straight, curved, or hooked; deflexed and swollen; glandular; with glandular hairs; without parallel to cotyledon length; centered between cotyle- spines; not smooth; with elevated features; not veined; dons; less than 1/2 length of cotyledons. Plumule not tuberculate; tuberculate (bases of hairs); not rudimentary or moderately developed; glabrous. exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without Distribution: Northern Europe to Canary Islands, Ethiopia, fibers; solid; coriaceous. Endocarp dull or glossy; and Iran. monochrome; tan; smooth; septate or nonseptate; with septa thin (tissue paper-like), flexible; with septa Notes: After a cladistic analysis using morphological eglandular; chartaceous; not exfoliating; remaining characters, including internal seed morphology, Endo fused to mesocarp and epicarp; entire. Seeds 5Ð10; and Ohashi (1997) have proposed that Cicereae (20) length parallel with fruit length; neither overlapping nor and Fabeae (19) formed a monophyetic group whose touching; in 1 series. Funiculus measured; up to 1 mm sister group is Trifolieae. Ononis and Parochetus long; of 1 length only; thick; straight. Aril absent. (21.02) “are not nearly as closely related to the remaining four genera as the latter are to each other, and Seed 0.5Ð5.5 ϫ 0.5Ð5 ϫ 1Ð4.5 mm; not overgrown; not indeed ... the two genera are not at all closely related to angular or angular; asymmetrical; mitaform, ovate, each other (or so far as I know to anything else)” (E. reniform, C-shaped (with cotyledon and radicles more Small, personal communication, 1997). Butler (1996) or less of equivalent size and on each side of a deep presented a table with eight seed characteristics of 14 hilar sinus), or triangular; compressed or terete; with Medicago (21.05) spp., 7 Melilotus (21.03) spp., 25

914 Trifolium (21.06) spp., 11 Trigonella (21.04) spp. and 2 Ononis spp. as an aid for their identification in archaeo- logical sites. Förther and Podlech (1991) revised the polymorphic O. natrix C. Linnaeus group of seven species. La Sota (1978) studied the testa morphology of 10 Ononis species.

915 Ononis: O. pubescens C. Linnaeus (CÐE), O. spp. (AÐB). A, Fruits with and without calyx (intact and dehisced) (ϫ 2.5); B, seeds (ϫ 6.4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

917 Genus: Parochetus F. Buchanan-Hamilton ex D. Don length; with margins curved; circular; not in groove of raphe; adjacent to hilum; 0.5 mm from hilum; flush; Phylogenetic Number: 21.02. similar color as or dissimilar color from testa; darker than testa; reddish brown. Endosperm quite thin; Tribe: Trifolieae. covering entire embryo; adnate to testa. Cotyledons smooth; both outer faces convex; both the same Species Studied—Species in Genus: 1 sp.—2 spp. thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; similar Fruit a legume; unilocular; 1.5Ð2.3 ϫ 0.4Ð0.5 cm; with at apex; not concealing radicle; entire over radicle; persistent calyx; with calyx shorter than fruit; without without lobes; with the interface division terminating at orifice formed by curving of fruit or fruit segments; base of radicle; without margins recessed; white; inner straight; not plicate; not twisted; symmetrical; linear or face flat; glabrous around base of radicle. Embryonic oblong (narrowly); inflated; without beak; short tapered axis deflexed; oblique to length of seed. Radicle linear; at apex; apex aligned with longitudinal axis of fruit; deflexed and parallel to cotyledon length; centered short tapered at base; base aligned with longitudinal between cotyledons; less than 1/2 length of cotyledons. axis of fruit; with the apex and base uniform in texture; Plumule moderately developed; glabrous. coriaceous; seed chambers externally invisible. Fruit margin not constricted; without sulcus; plain. Fruit Distribution: Tropical Africa, Asia (Himalaya Mountains to wings absent. Fruit substipitate. Fruit with all layers Sri Lanka and eastern China) to Indonesia (southeastern dehiscing (to tardily dehiscent); splitting along sutures. Java), and Europe (cultivated). Dehiscence of valves along both sutures; apical and down; active; with valves twisting. Replum invisible. Notes: For additional information, see the Notes for Ononis Epicarp dull; monochrome; dark brown; glabrous or (21.01). Polhill, in Beckett and Polhill (1991), named a pubescent but soon deciduous; with 1 type of pubes- second species, P. africanus R.M. Polhill. cence; rarely pilose; with pubescence gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; not veined; faintly tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; greenish tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 4Ð15; length transverse to fruit length; in 2 or more series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 1.5Ð2.5 ϫ 1.4Ð2 ϫ 1Ð1.5 mm; not overgrown; not angular; asymmetrical; oblong; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled and streaked; with frequent mottles; with frequent streaks; blackish brown; with brown overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; recessed; within rim. Hilum Parochetus: P. communis F. Buchanan-Hamilton ex D. Don rim color slightly darker than testa. Lens not discernible (AÐE). A, Fruits (dehisced) with calyx (ϫ 3.7); B, seeds or discernible (barely sometimes); less than 0.5 mm in (ϫ 11); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 15).

919 Genus: Melilotus P. Miller frequent streaks; brown (and yellowish, greenish), green (and yellowish), tan, or yellow; with purple Phylogenetic Number: 21.03. overlay; glabrous; smooth or not smooth; with elevated features; shagreen or warty; coriaceous. Fracture lines Tribe: Trifolieae. absent. Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by funicular Species Studied—Species in Genus: 16 spp.—19 spp. remnant; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform Fruit a legume; unilocular; 0.15Ð0.8 ϫ 0.15Ð0.45 ϫ 0.12Ð or larger than punctiform; with curved or angular 0.4 cm; with persistent or deciduous calyx; with calyx outline; circular or triangular; between cotyledon and shorter than fruit; without orifice formed by curving of radicle lobe; flush; not within corona, halo, or rim. Lens fruit or fruit segments; straight; not plicate; not twisted; discernible; less than 0.5 mm or equal to or greater than asymmetrical or symmetrical; circular, lanceolate, 0.5 mm in length; 0.6 mm long; with margins straight oblong, obovate, or ovate; when asymmetrical with or curved; oblong; not in groove of raphe; confluent both sutures parallelly curved; not inflated; compressed with hilum (at least discolor area); mounded; dissimilar or terete; without beak; short tapered or rounded at color from testa; darker than testa; brown; not within apex; apex aligned or oblique (slightly) with longitudi- corona, halo, or rim. Endosperm thick; covering entire nal axis of fruit; short tapered or rounded at base; base embryo; adnate to embryo. Cotyledons smooth; both aligned with longitudinal axis of fruit; with the apex outer faces convex; both the same thickness; not folded and base uniform in texture; membranous, chartaceous, or with both folded; not sufficiently folded for inner or fragile, thinner than chartaceous like Trifolium face to touch itself (essentially cotyledons longer than (21.06); seed chambers externally visible. Fruit margin testa); portions of inner folded face equal; margin entire not constricted; without sulcus; plain. Fruit wings 180 degrees from base of radicle; similar at apex; not absent. Fruit nonstipitate. Fruit indehiscent or with all concealing radicle; entire over or notched at radicle; layers dehiscing (M. altissimus J.L. Thuillier); splitting without lobes; with the interface division terminating at along suture. Dehiscence of valves along 1 suture; base of radicle; without margins recessed; tan; inner medial and up and down; passive. Replum invisible. face flat; glabrous around base of radicle. Embryonic Epicarp dull; monochrome; black, brown, or tan; axis deflexed; oblique to length of seed. Radicle glabrous or pubescent and indurate; with hairs ap- bulbose; deflexed and parallel to cotyledon length; pressed; with 1 type of pubescence; with pubescence centered between cotyledons; 1/2 to nearly length of gray; with pubescence uniformly distributed; with cotyledons. Plumule rudimentary or moderately simple hairs; pliable; with hair bases plain; eglandular; developed; glabrous. without spines; not smooth; with elevated features; veined or not veined; obliquely veined relative to fruit Distribution: Europe, Asia, and North Africa. length or reticulately veined; not tuberculate; concentric whorls like a fingerprint, ribbed, or wrinkled; not Notes: Stevenson (1969) reviewed Melilotus, recognized 18 exfoliating; without cracks. Mesocarp absent. Endocarp species, and illustrated its fruits. Jha and Pandey (1989) dull; monochrome; brown or tan; smooth; nonseptate; studied the seeds, especially the testae, of 11 species of chartaceous; not exfoliating; remaining fused to Melilotus. They concluded that the testa coat ornamen- epicarp; entire. Seeds 1 or 2; length parallel with fruit tation for a species is characteristic “and of great length; neither overlapping nor touching or touching; in importance in specific diagnosis of a seed.” Isely 1 series. Funiculus less than 0.5 mm long or measured; (1954) provided flower-fruit keys to the 20 species of up to 0.8 mm long; of 1 length only; filiform; straight or Melilotus. Voronchikhin and Bazilevskaya (1974) curved. Aril absent. described fruits and seeds of the nine species of Melilotus in the former U.S.S.R. Small (1989) noted Seed 1.5Ð4.5 ϫ 1Ð3 ϫ 1Ð2.2 mm; not overgrown; not that M. bicolor P.E. Boissier & B. Balansa may perhaps angular or angular (somewhat); asymmetrical; circular, be better placed in Trigonella (21.04). Our number of elliptic, mitaform, or reniform; compressed; with species includes M. bicolor and is the same as the count visible radicle and cotyledon lobes; without umbo on used by Small (1989); we do not agree with the species seed faces. Testa not adhering to endocarp; glossy or count of ca. 20 by Heyn (1981). We are not adopting dull; not modified by a bloom; colored; monochrome, Small’s selection of “a” to end species names currently mottled, or streaked; with frequent mottles; with ending in “us” in Wiersema et al. (1990). Schulz

920 (1901:667) noted that the two seeds in a pod may be differently colored: one light yellowish brown and the other dark reddish brown.

921 Melilotus: M. indicus (C. Linnaeus) C. Allioni (CÐE), M. spp. (AÐB). A, Fruits, most with calyx (ϫ 3); B, seeds (ϫ 6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

923 Genus: Trigonella C. Linnaeus touching; in 1 series. Funiculus less than 0.5 mm long or measured; up to 0.6 mm long; of 1 length only; Phylogenetic Number: 21.04. filiform; curved. Aril absent.

Tribe: Trifolieae. Seed 1.5Ð5 ϫ 0.5Ð3 ϫ 0.5Ð1.6 mm; not overgrown; angular or not angular; asymmetrical; mitaform, oblong, ovate, Species Studied—Species in Genus: 25 spp.—ca. 61 spp. quadrangular, rectangular, rhombic, elliptic, linear, or circular (sub); compressed; with visible radicle and Fruit a legume; unilocular; 1Ð11 ϫ 0.09Ð1.3 ϫ 0.07Ð0.08 cotyledon lobes; without umbo on seed faces. Testa not cm; with persistent or deciduous calyx; with calyx adhering to endocarp; dull; not modified by a bloom; shorter than fruit; without orifice formed by curving of colored; monochrome or mottled and streaked; with fruit or fruit segments; straight to curved (slightly), 0.5- frequent or infrequent mottles; with frequent or coiled, or contorted; loosely plicate; not twisted; infrequent streaks; yellowish, greenish, or dark brown, asymmetrical or symmetrical; linear, ovate, circular, yellow (or pale), orange, or green (grayish); with black moniliform, C-shaped, or falcate; when asymmetrical or brown overlay; glabrous; not smooth or smooth; with with 1 straight and 1 curved suture or both sutures elevated features; tuberculate, wrinkled, or verrucose; parallelly curved; widest near middle or D-shaped; coriaceous. Fracture lines absent. Rim absent. Wings inflated or not inflated; compressed, flattened, or terete; absent. Raphe not visible. Hilum visible or partially without or with beak; straight or declined; with solid concealed; concealed by funicular remnant or wing; beak the same color and texture as fruit; long tapered or with faboid split; with the lips of the faboid split the rounded at apex; apex aligned with longitudinal axis of same color as the rest of the hilum; punctiform; fruit; tapered or short tapered at base; base aligned with between cotyledon and radicle lobe; recessed; not longitudinal axis of fruit; with the apex and base within corona, halo, or rim. Lens not discernible or uniform in texture; chartaceous or coriaceous; seed discernible; less than 0.5 mm in length; with margins chambers externally visible; with the raised seed straight or curved; triangular or irregular; circular or chambers not torulose or torulose. Fruit margin not irregular; not in groove of raphe; adjacent to hilum; constricted or constricted; slightly constricted along 0.1Ð0.3 mm from hilum; mounded; similar color as both margins or constricted only on 1 margin; without testa; darker than testa; reddish brown; not within sulcus; plain or embellished; with prickles. Fruit wing corona, halo, or rim. Endosperm thin or thick; covering absent or present (rarely); 1; 1Ð2 mm wide; sutural; on entire embryo; adnate to testa. Cotyledons smooth; both both sutures or 1 suture. Fruit nonstipitate or outer faces convex; both the same thickness; both more substipitate. Fruit indehiscent (gaping or not along or less of equal length; not folded; margin entire 180 seed-bearing suture) or with all layers dehiscing degrees from base of radicle; similar at apex; not (rarely); splitting along sutures. Dehiscence of valves concealing radicle; entire over radicle; without lobes; along both sutures or 1 suture; apical and down; with the interface division terminating at base of passive. Replum invisible. Epicarp dull; monochrome; radicle; without margins recessed; yellow or tan; inner tan (to white); glabrous or pubescent and indurate; with face flat; glabrous around base of radicle. Embryonic hairs appressed or erect; with 1 type of pubescence; axis deflexed; oblique to length of seed. Radicle linear puberulent; with pubescence gray; with pubescence or bulbose; lobe tip straight; deflexed and parallel to uniformly distributed; with simple hairs; pliable; with cotyledon length; centered between cotyledons; less hair bases plain; eglandular; without spines; not than 1/2 or 1/2 to nearly length of cotyledons. Plumule smooth; with elevated features; reticulately veined or moderately developed; glabrous. longitudinally veined relative to fruit length (and veins twisted); not tuberculate; warty; not exfoliating; without Distribution: Central Europe to South Africa to Canary cracks. Mesocarp present or absent; thin; surface not Islands and Central Asia, and Australia (1 sp.). veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; Notes: There is a rich history of authors trying to separate tan; smooth; nonseptate; chartaceous; not exfoliating; Trigonella and Medicago (21.05), and most of this remaining fused to mesocarp and epicarp or to epicarp; history was summarized by Baum (1968). The recent entire. Seeds 1Ð20; length parallel with or transverse to publications of Small (1986, 1987a,b) have clarified fruit length; neither overlapping nor touching or this separation. Small and Jomphe (1989a) transferred

924 some species of Trigonella to Medicago, and our species count reflects these and other transfers. Our species count is based on Small (1989) and not on Heyn (1981), who had ca. 80. The number of perennial Asian species can only be roughly estimated (Small, personal communication, 1997).

925 Trigonella: T. foenum-graecum C. Linnaeus (CÐE), T. spp. (AÐB). A, Fruits (few with calyx) (ϫ 1.2); B, seeds (ϫ 6.7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

927 Genus: Medicago C. Linnaeus Seed 1.2Ð7 ϫ 1Ð4.5 ϫ 0.7Ð1.9 mm; not overgrown; not angular or angular; asymmetrical; mitaform, oblong, Phylogenetic Number: 21.05. rhombic, or triangular; compressed; with or without visible radicle and cotyledon lobes; without umbo on Tribe: Trifolieae. seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; brown (or Species Studied—Species in Genus: 83 spp.—86 spp. yellowish, reddish, or blackish) or yellow; glabrous; smooth or not smooth; with elevated features; rugose, Fruit a legume or nutlet; unilocular; 0.2Ð10 ϫ 0.1Ð1 ϫ 0.2Ð wrinkled, papillate, or transversely ridged; coriaceous. 1.5 cm; with persistent calyx; with calyx shorter than Fracture lines absent. Rim absent. Wings absent. Raphe fruit; with orifice formed by curving of fruit or fruit not visible. Hilum visible; with faboid split; with the segments (with 1 or more than 1 per fruit); straight, lips of the faboid split the same color as the rest of the curved (slightly), 0.5-coiled, 1-coiled, 1.5-coiled, 2- hilum; not within corona, halo, or rim. Cotyledons not coiled, 3-coiled, 4-coiled, or 5- to 10-coiled (rarely); concealing radicle; entire over radicle; brown to tan; not plicate or plicate (M. plicata (P.E. Boissier) G.I. with the interface division terminating at base of Sirjaev); not twisted or twisted; asymmetrical or radicle. Embryonic axis deflexed; oblique to length of symmetrical; circular, falcate, or coiled; when asym- seed. Radicle lobe tip straight; deflexed and parallel to metrical with 1 straight and 1 curved suture or both cotyledon length; 1/2 to nearly or equaling length of sutures parallelly curved; widest near middle or D- cotyledons. Plumule rudimentary; glabrous. shaped; not inflated; compressed, terete, or flattened; without beak; short tapered at base; base right angled Distribution: Europe to South Africa and western Asia; with longitudinal axis of fruit; with the apex and base widely cultivated. uniform in texture; coriaceous, membranous, or ligneous; seed chambers externally invisible. Fruit Notes: In an impressive number of papers over the past 20 margin not constricted or constricted; constricted along years dealing with Medicago and related genera, Small both margins; without sulcus; plain or embellished; has resolved many of the problems related to Medicago. with prickles or fringe. Fruit wing absent or present (M. Our species count follows that of Small et al. (1988), popovii (E.I. Korneva) G.I. Sirjaev); 1; sutural; on 1 Small and Jomphe (1989a,b), Small (1990a,b), and valve; on 1 suture. Fruit substipitate or nonstipitate. Small and Brookes (1991), who included in Medicago Fruit with all layers dehiscing or indehiscent; splitting species previously in Trigonella (21.04). We did not along sutures. Dehiscence of valves passive. Replum follow the species count of ca. 50 of Heyn (1981). invisible. Epicarp dull; monochrome; brown or black; Small and Jomphe illustrated seeds and fruits in line glabrous or pubescent and indurate; with 1 or 2 types of drawings, and Small et al. (1990) illustrated seeds in pubescence; tomentose (M. hypogaea E. Small); with photographs and micrographs. Factorovskya Eig, a pubescence gray or brown; with simple or glandular monotypic and geocarpic genus recognized by Heyn, hairs; pliable; with hair bases plain; glandular or was reduced to Medicago hypogaea by Small and eglandular; with glandular hairs; without or with spines Brookes (1984) and is included as such here. The (spines forked or more often not); smooth or not authors used the characteristic coiling of the fruit as one smooth; with elevated features; reticulately veined; not of the main traits supporting the transfer to M. tuberculate or tuberculate; with solid tubercles on each hypogaea. Small and Jomphe (1989b) presented a key valve; exfoliating in part; without cracks. Mesocarp to the 12 sections and 8 subsections of the genus and a absent. Endocarp dull; monochrome; nonseptate, comprehensive illustrated key to the 83 species. The subseptate, or septate; with septa thin (tissue paper- following year he circumscribed the genus based on like), flexible or thicker than paper, firm; with septa seed characters (Small et al. 1990). Some species of eglandular; chartaceous; not exfoliating; remaining Medicago are important crops throughout the temperate fused to epicarp; entire. Seeds 1Ð6(Ð20) (20 or more in world. The fruits of Medicago are as diverse as they are M. scutella (C. Linnaeus) P. Miller (Small, personal in any faboid genus: Straight, curved, or coiled; terete, communication, 1997)); length parallel with fruit compressed, or flat; and sutures spiny, tuberculate, length; neither overlapping nor touching or touching; in 1 series. Aril absent.

928 winged, frimbiate, or plain. Most Medicago species have numerous conspicuous veins in the fruit. These veins are located in the epicarp, unlike most other legumes, which have the veins in the mesocarp.

929 Medicago: M. ciliaris (C. Linnaeus) C. Allioni (CÐE), M. spp. (AÐB). A, Fruits with and without calyx (ϫ 1.2); B, seeds (ϫ 3.9); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

931 Genus: Trifolium C. Linnaeus glabrous; smooth or not smooth; with elevated or recessed features; shagreen, tuberculate (minutely), Phylogenetic Number: 21.06. warty (finely), or wrinkled; pitted with small separate pits; coriaceous. Fracture lines absent. Rim absent. Tribe: Trifolieae. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same Species Studied—Species in Genus: 121 spp.—ca. 250 spp. color as the rest of the hilum; punctiform; between cotyledon and radicle lobe. Endosperm thin; covering Fruit a legume or nutlet; unilocular; 0.14Ð1.5 ϫ 0.04Ð0.5 entire embryo. Cotyledons smooth; both outer faces cm; with persistent calyx; with calyx longer or shorter convex; both the same thickness; both more or less of than fruit; without orifice formed by curving of fruit or equal length; not folded; margin entire 180 degrees fruit segments; straight; not plicate; not twisted; from base of radicle; similar at apex; not concealing asymmetrical or symmetrical; oblong (to linear or radicle; entire over radicle; without lobes; with the ovate), ovate, elliptic, fusiform, lanceolate, linear, interface division terminating at base of radicle. dolabriform, or falcate; when asymmetrical with both Embryonic axis deflexed; oblique to length of seed. sutures parallelly curved; inflated or not inflated; Radicle bulbose or linear; lobe tip straight; deflexed compressed; without beak; short tapered or rounded at and parallel to cotyledon length; 1/2 to nearly or apex; with the apex and base uniform or differing in equaling length of cotyledons. Plumule rudimentary; texture; fragile, thinner than chartaceous, membranous, glabrous. chartaceous, coriaceous, or leathery (rarely); seed chambers externally visible or invisible. Fruit margin Distribution: Temperate and subtropical regions of the constricted or not constricted; slightly constricted along Northern and Southern Hemispheres. both margins; without sulcus; plain. Fruit wings absent. Fruit substipitate or nonstipitate. Fruit indehiscent or Notes: In discussing the evolution of the legume (their with all layers dehiscing (with 2 or 1 prominent legumen) in the genus Trifolium, Zohary and Heller sutures); splitting along sutures. Dehiscence of valves (1984) noted that the section Lotoidea has the most along 1 suture or both sutures; passive. Replum primitive legume in the genus. These legumes are often invisible. Epicarp dull; monochrome; brown (various 2Ð9-seeded and dehiscent by both sutures. From this shades, yellowish); glabrous or glabrate; with 1 type or primitive state, the advanced one-seeded “utricle or 2 types of pubescence; pilose or puberulent (slightly); nutlet” evolved with a membranous pericarp that may with pubescence gray; with apical pubescence different consist of only an epidermal layer. These legumes do from basal pubescence; with apical 1/4 tomentose and not dehisce but instead split transversely (circum- basal 3/4 glabrous or apical 3/4 tomentose and basal 1/4 scissly) or irregularly and are found in the section glabrous; with simple hairs; pliable; with hair bases Trifolium and even in advanced species in section plain; with elevated features; veined or not veined; Lotoidea H. J. N. von Crantz. Some fruits are opercu- longitudinally veined relative to fruit length (occasion- late. Unlike the fruiting heads of other legume genera, ally) or reticulately veined; not tuberculate; not exfoli- the fruiting heads of a few species of Trifolium separate ating; without cracks. Mesocarp absent. Endocarp from the plants and serve as the unit of dispersal. In remaining fused to epicarp. Seeds 1Ð10. Aril absent. Trifolium subterraneum C. Linnaeus, fertile flowers are pushed into the ground, where the fruits and seeds Seed 0.8Ð5 ϫ 0.8Ð3 mm; not overgrown; not angular; mature. See Arachis (14.26) and Medicago (21.05). symmetrical or asymmetrical; circular, mitaform, oblong, pyriform, quadrangular, reniform, rhombic, or triangular; terete, compressed, or flattened; with or without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome, mottled, or streaked; with frequent mottles; with frequent streaks; black (to reddish, Trifolium: T. pannonicum N. von Jacquin (CÐE), T. spp. greenish, purplish, yellowish, dark, or reddish), green (AÐB). A, Fruits (mostly within calyxes) (ϫ 2.1); B, (yellowish to pale), red, tan, or yellow (to lemon); with seeds (ϫ 6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos black, brown (various shades), or purple overlay; (ϫ 10).

933 Brongniartieae (22.01Ð22.02) or tan; with brown (dark) overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe Genus: Brongniartia K.S. Kunth visible or not visible; from lens to base of seed and terminating; color of to lighter than testa; tan; flush. Phylogenetic Number: 22.01. Hilum fully concealed; concealed by funiculus or funicular remnant; with faboid split; with the lips of the Tribe: Brongniartieae. faboid split the same color as the rest of the hilum; larger than punctiform; 0.8Ð1 mm long; with curved Species Studied—Species in Genus: 15 spp.—56 spp. outline; circular; subapical to radicle tip; recessed; within rim or not within corona, halo, or rim. Hilum rim Fruit a legume; unilocular; 1.3Ð6 ϫ 0.8Ð2.5 ϫ 0.4Ð0.5 cm; color of (but not mottled if testa mottled) or darker than with deciduous calyx; without orifice formed by testa (but not mottled if testa mottled). Lens discernible; curving of fruit or fruit segments; straight; not plicate; equal to or greater than 0.5 mm in length; 1Ð1.2 mm not twisted; asymmetrical or symmetrical; elliptic or long; with margins straight or curved; linear or elliptic; oblong; when asymmetrical with 1 straight and 1 not in groove of raphe; adjacent to hilum; 0.5 mm from curved suture, both sutures parallelly curved, or both hilum; mounded; same color as testa; brown or tan; not sutures unequally curved; not inflated; compressed; within corona, halo, or rim. Endosperm thin; covering without beak; short tapered at apex; apex aligned with entire embryo; adnate to testa. Cotyledons smooth; both longitudinal axis of fruit; tapered at base; base aligned outer faces convex; both the same thickness; both more with longitudinal axis of fruit; with the apex and base or less of equal length; not folded; margin entire 180 uniform in texture; coriaceous; seed chambers exter- degrees from base of radicle; similar at apex; partially nally invisible. Fruit margin not constricted; without concealing radicle; split over radicle; with lobes; with sulcus; plain. Fruit wings absent. Fruit nonstipitate. lobes overlapping; with basal groin formed by lobes; Fruit with all layers dehiscing; splitting along sutures. with the interface division terminating at base of Dehiscence of valves along both sutures; apical and radicle; without margins recessed; tan; inner face flat; down (assumed); active; with valves twisting. Replum glabrous around base of radicle. Embryonic axis invisible. Epicarp dull; monochrome; brown or tan; straight; parallel to length of seed. Radicle linear; pubescent and indurate; with 1 type of pubescence; with straight with embryonic axis; centered between pubescence gray; pliable; with hair bases plain; cotyledons; less than 1/2 length of cotyledons. Plumule eglandular; without spines; not smooth; obliquely well developed; glabrous. veined relative to fruit length; not tuberculate; not exfoliating; without cracks. Mesocarp thin; 1-layered; Distribution: Southern United States, Mexico, Central without balsamic vesicles; without fibers; solid; America, South America. coriaceous. Endocarp dull; monochrome; tan or yellow; fibrous (but fibrous around margins of seeds or not) or Notes: Arroyo (1981) noted that the “legumes and seeds in smooth; subseptate or nonseptate; with septa thin Harpalyce and Brongniartia are quite similar and on (tissue paper-like), flexible; with septa eglandular; this basis the genera are closely related.” She regarded coriaceous; not exfoliating; remaining fused to meso- this tribe as “a probably retictual American group, with carp and epicarp. Seeds 1Ð8; length transverse (to ancient austral roots.” Crisp and Weston (1987, pp. slightly oblique) or oblique to fruit length; neither 105Ð107) provided compelling evidence that the overlapping nor touching; in 1 series. Funiculus Templetonia group of the Bossiaeeae should be measured; 1Ð4 mm long; of 1 length only; flattened; transferred to the Brongniartieae, but they did not straight. Aril fleshy; hooked; covering less than 1/2 of resolve all aspects of the transfer. See the Notes for seed; brown to tan. Lamprolobium (23.02). Chappill’s (1995) cladistic analysis of the entire family indicated that the transfer Seed 6Ð20 ϫ 3.5Ð6.5 ϫ 2.5Ð4 mm; not overgrown; angular should not be made and that the two tribes should be or not angular; asymmetrical; irregular (because of circumscribed in the traditional manner (Crisp and pressure of adjacent seeds) or oblong to ovate; com- Weston 1995). pressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to Brongniartia: B. intermedia S. Moricand (CÐE), B. spp. (AÐ endocarp; glossy; not modified by a bloom; colored; B). A, Valves (ϫ 0.9); B, seeds (ϫ 1.5); CÐD, testa monochrome or mottled; with frequent mottles; brown (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

935 Genus: Harpalyce A.-P. de Candolle Arroyo fide Arroyo (1976)); smooth; coriaceous. Fracture lines absent. Rim absent. Raphe visible or not Phylogenetic Number: 22.02. visible; from lens to base of seed and terminating; color of testa; recessed. Hilum fully concealed; with faboid Tribe: Brongniartieae. split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.5Ð0.7 Species Studied—Species in Genus: 9 spp.—20 spp. mm long; with curved outline; elliptic; subapical to radicle tip; recessed; within rim or not within corona, Fruit a legume; unilocular; 1Ð12 ϫ 0.8Ð3.5 ϫ 0.3Ð0.6 halo, or rim. Hilum rim color of testa. Lens discernible (estimated) cm; with deciduous calyx; without orifice or not discernible; less than 0.5 mm in length; with formed by curving of fruit or fruit segments; straight to margins straight or curved; linear; punctiform; in curved (slightly); not plicate; not twisted; symmetrical; groove of raphe; adjacent to hilum; recessed; dissimilar oblong to obovate to ovate (when obovate, lower color from testa; darker than testa; black; not within portion of fruit sterile); not inflated; compressed; corona, halo, or rim. Endosperm thin; covering entire without beak; short tapered at apex; apex oblique with embryo; adnate to testa. Cotyledons smooth; both outer longitudinal axis of fruit; short tapered at base; base faces convex; both the same thickness; both more or aligned with longitudinal axis of fruit; with the apex less of equal length; not folded; margin entire 180 and base uniform in texture; fleshy (when fresh), degrees from base of radicle; similar at apex; partially coriaceous, or ligneous (H. mexicana J.N. Rose); seed concealing radicle; split over radicle; with lobes; with chambers externally visible; with the raised seed lobes touching (auriculate); with basal groin formed by chambers not torulose. Fruit margin not constricted; lobes; with the interface division terminating at base of without sulcus; plain. Fruit wings absent. Fruit radicle; without margins recessed; tan; inner face flat; nonstipitate. Fruit with all layers dehiscing. Dehiscence glabrous around base of radicle. Embryonic axis of valves along both sutures; apical and down; active or oblique; oblique to length of seed. Radicle linear; passive (H. mexicana); with valves revolute. Replum oblique to cotyledons; centered between cotyledons; invisible. Epicarp dull or glossy; monochrome; black, less than 1/2 length of cotyledons. Plumule rudimen- brown (reddish-brown to darker), purple, or tan; tary; glabrous. glabrous; eglandular; without spines; not smooth; with elevated features; obliquely veined relative to fruit Distribution: Mexico, Central America, Cuba, Brazil. length; not tuberculate; obscurely rugose; not exfoliating; without cracks. Mesocarp thin; 1-layered; without Notes: Arroyo (1976) monographed the genus and provided balsamic vesicles; without fibers; solid; subligneous. general fruit and seed descriptions for most recognized Endocarp dull or glossy (at least lustrous); mono- species. Our studied species were selected from all chrome; tan; cobwebby, smooth, or spongy; septate or three sections: Harpalyce, Brazilianae M.T.K. Arroyo, nonseptate; with septa thin (tissue paper-like), flexible and Cubenses P.A. Rydberg. The lens is a black line in or thicker than paper, firm; with septa eglandular; the raphe groove and is surrounded by a punctiform pit. coriaceous; not exfoliating. Seeds 1Ð15; length oblique (H. macrocarpa N.L. Britton & P. Wilson) or transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 2Ð4 mm long; of 1 length only; filiform or triangular; straight. Aril fleshy; topknotlike, cupshaped, or hooked; covering less than 1/2 of seed; brown or tan.

Seed 5Ð15 ϫ 2Ð12 ϫ 2.5Ð5 mm; not overgrown; not angular; asymmetrical; oblong to rectangular; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; black, brown, cream (H. parvifolia H.S. Harpalyce: H. brasiliana G. Bentham (CÐE), H. spp. (AÐB). Irwin & M.T.K. Arroyo), ivory, or olive; glabrous or A, Valves (ϫ 1); B, seeds (ϫ 4); CÐD, testa (ϫ 50, minutely pubescent (H. robusta H.S. Irwin & M.T.K. ϫ 1000); E, embryos (ϫ 5).

937 Bossiaeeae (23.01Ð23.10) adhering to endocarp; glossy; not modified by a bloom; colored; monochrome; yellowish to reddish or olive Genus: Templetonia R. Brown brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Phylogenetic Number: 23.01. Hilum fully concealed; concealed by aril; with faboid split; with the lips of the faboid split the same color as Tribe: Bossiaeeae. the rest of the hilum; larger than punctiform; 1 mm long; with curved outline; elliptic; subapical to radicle Group: Templetonia. tip; flush; within rim. Hilum rim color darker than testa. Lens discernible; equal to or greater than 0.5 mm in Species Studied—Species in Genus: 4 spp.—9 spp. length; 1 mm long; with margins straight or curved; rhombic; not in groove of raphe; adjacent to hilum; 1 Fruit a legume; unilocular; 1.5Ð8 ϫ 0.6Ð1.6 ϫ 0.25 cm; mm from hilum; barely mounded; dissimilar color from with deciduous corolla; with persistent or deciduous testa; darker than testa; brown; not within corona, halo, calyx; with calyx shorter than fruit; without orifice or rim. Endosperm thin; covering entire embryo; adnate formed by curving of fruit or fruit segments; straight; to testa. Cotyledons smooth; both outer faces convex; not plicate; not twisted; asymmetrical or symmetrical; both the same thickness; both more or less of equal elliptic or linear; when asymmetrical with both sutures length; not folded; margin entire 180 degrees from base parallelly curved to unequally curved; not inflated; of radicle; similar at apex; partially concealing radicle; flattened; without beak; short tapered or rounded at split over radicle; with lobes; with lobes touching apex; apex aligned to oblique with longitudinal axis of (auriculate); with the interface division terminating at fruit; short tapered or rounded at base; base aligned base of radicle; without margins recessed; yellow; inner with longitudinal axis of fruit; with the apex and base face flat; glabrous around base of radicle. Embryonic uniform in texture; coriaceous; seed chambers exter- axis almost straight; parallel to length of seed. Radicle nally visible or invisible; with the raised seed chambers nearly linear; straight with embryonic axis; centered not torulose. Fruit margin not constricted; without between cotyledons; less than 1/2 length of cotyledons. sulcus; plain. Fruit wings absent. Fruit stipitate or Plumule rudimentary; glabrous. nonstipitate; with the stipe up to 10 mm long. Fruit with all layers dehiscing; splitting along sutures. Dehiscence Distribution: Australia. of valves along both sutures; apical and down; active or passive; with valves twisting. Replum invisible. Epicarp Notes: The Bossiaeeae traditionally consisted of 10 genera dull or semiglossy; monochrome; dark reddish and dark and has been divided into two groups, in part on fruit greenish brown; glabrous; eglandular; without spines; and seed characters. The Templetonia group contains not smooth; with elevated features; reticulately veined; the first four genera (including Lamprolobium, 23.02) not tuberculate; not exfoliating; without cracks. and has nonwinged, coriaceous legumes with com- Mesocarp thin; surface not veined; 1-layered; without pressed seeds bearing a collarlike, often lipped aril balsamic vesicles; without fibers; solid; coriaceous. (except for Templetonia biloba (G. Bentham) R.M. Endocarp dull; monochrome; brown; smooth; septate or Polhill) and a short, straight radicle. The Bossiaea nonseptate; with septa thin (tissue paper-like), flexible; group contains the remaining genera and has legumes with septa eglandular; chartaceous; not exfoliating; that are keeled to winged or not so; the group has remaining fused to epicarp; entire. Seeds 1Ð12; length plump seeds that are often covered by a hooded caplike parallel or oblique to fruit length; neither overlapping aril (which is lacking in Muelleranthus (23.08) and nor touching; in 1 series. Funiculus less than 0.5 mm Ptychosema (23.09)) and a radicle that is slender, long; of 1 length only; thick; straight or hooked. Aril deflexed, and exserted from the cotyledons. Crisp and fleshy; caplike (T. biloba (G. Bentham) R.M. Polhill) or Weston (1987, pp. 105Ð107) in their cladistic analysis cupshaped (lipped or not); covering less than 1/2 of of the Bossiaeeae, Brongniartieae (22), and Mirbelieae seed; brown. (24), provided compelling evidence that the Bossiaeeae should be redefined to include only the Bossiaea group Seed 3.5Ð14.5 ϫ 2Ð8.5 ϫ 1.2Ð3.3 mm; not overgrown; not (genera 23.05Ð23.10). They proposed that the angular; symmetrical (except for hilum); elliptic to Templetonia group be moved to the Brongniartieae, ovate; compressed; without visible radicle and cotyle- becoming genera three through eight, after Brongniarta don lobes; without umbo on seed faces. Testa not (22.01) and Harpalyce (22.02). The proposed generic

938 sequence would therefore be: 3, Templetonia (23.01); 4, Lamprolobium; 5, Plagiocarpus (23.03); 6, ?Genus A (Templetonia incana J.H. Ross); 7, ?Genus B (Templetonia biloba (G. Bentham) R.M. Polhill); and 8, Hovea (23.04). They (Crisp and Weston 1995) retracted their proposal to transfer the Templetonia group because of Chappill’s (1995) cladistic analysis of the entire family, and we have used the traditional circumscriptions of the two tribes (Polhill 1994a,b). Polhill (1981n) reported 9 species in this genus, but Ross (1982) recognized 11 species, the number used in our study. Polhill (1976) has a full plate of the external and internal morphology of the seeds of Templetonia biloba (G. Bentham) R.M. Polhill.

939 Templetonia: T. retusa (E.P. Ventenat) R. Brown (CÐE), T. spp. (AÐB). A, Fruits (with and without calyx) and valve (ϫ 1.6); B, seeds (ϫ 4.1); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

941 Genus: Lamprolobium G. Bentham Hilum partially concealed; concealed by aril; with faboid split; with the lips of the faboid split the same Phylogenetic Number: 23.02. color as the rest of the hilum; larger than punctiform; 2 mm long; with curved outline; elliptic; apical according Tribe: Bossiaeeae. to radicle tip but marginal according to seed length; recessed; not within corona, halo, or rim. Lens discern- Group: Templetonia. ible; less than 0.5 mm in length; with margins curved; elliptic; in groove of raphe; adjacent to hilum; 1.5 mm Species Studied—Species in Genus: 1 sp.—2 spp. from hilum; mounded; similar color as testa; darker than testa; not within corona, halo, or rim. Endosperm Fruit a legume; unilocular; 4 ϫ 0.8Ð1 ϫ 0.5 (estimated) cm; thin; covering entire embryo; adnate to testa. Cotyle- with deciduous corolla; with deciduous calyx; without dons smooth; both outer faces convex; both the same orifice formed by curving of fruit or fruit segments; thickness; both more or less of equal length; margin straight; not plicate; not twisted; symmetrical; oblong; entire 180 degrees from base of radicle; similar at apex; not inflated; flattened; without beak; rounded at apex; partially concealing radicle; split over radicle; with apex aligned with longitudinal axis of fruit; short lobes; with lobes touching (auriculate); with the tapered at base; base aligned with longitudinal axis of interface division terminating at base of radicle; without fruit; with the apex and base uniform in texture; margins recessed; yellow; inner face flat; glabrous coriaceous; seed chambers externally visible; with the around base of radicle. Embryonic axis deflexed; raised seed chambers not torulose. Fruit margin not oblique to length of seed. Radicle bulbose; oblique to constricted; without sulcus; plain. Fruit wings absent. cotyledons; centered between cotyledons; less than 1/2 Fruit substipitate. Fruit with all layers dehiscing; length of cotyledons. Plumule rudimentary; glabrous. splitting along sutures. Dehiscence of valves along both sutures; apical and down (assumed); active; with valves Distribution: Australia (north Queensland). twisting. Replum invisible. Epicarp semiglossy; monochrome; brown; glabrous; eglandular; without Notes: The hilum length:seed length ratio is similar to the spines; not smooth; with elevated features; reticulately ratios seen in seeds of genera in tribe Phaseoleae (10) veined; not tuberculate; wrinkled (faintly); not exfoliat- and Hovea (23.04). ing; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; brown; smooth; subseptate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 5Ð7; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 3 mm long; of 1 length only; thick; straight. Aril fleshy; cupshaped (with 1 side longer (liplike) than other); covering less than 1/2 of seed; tan.

Seed 5.3Ð6 ϫ 3.5Ð4 ϫ 1.7Ð2.5 mm; not overgrown; not angular; asymmetrical; oblong; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome (when immature), mottled (when mature), or streaked (when mature); with frequent mottles; with frequent streaks; reddish brown; with brown (darker reddish) overlay; glabrous; smooth; coriaceous. Fracture lines absent. Lamprolobium: L. fruticosum G. Bentham (AÐE). A, Valve Rim absent. Wings absent. Raphe from hilum through (ϫ 2.3); B, seeds (ϫ 5.1); CÐD, testa (ϫ 50, ϫ 1000); lens to base of seed and terminating (beyond base of E, embryos (ϫ 5). seed); not bifurcating; darker than testa; recessed.

943 Genus: Plagiocarpus G. Bentham lens darker than between lens and base of seed); brown; flush. Hilum partially or fully concealed; concealed by Phylogenetic Number: 23.03. funiculus and aril; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; Tribe: Bossiaeeae. larger than punctiform; 0.9Ð1 mm long; with curved outline; elliptic; subapical to radicle tip; flush; not Group: Templetonia. within corona, halo, or rim. Lens discernible; less than 0.5 mm or equal to or greater than 0.5 mm in length; Species Studied—Species in Genus: 1 sp.—1 sp. 0.5 mm long; with margins curved; roughly circular; not in groove of raphe; adjacent to hilum; 0.6 mm from Fruit a legume; unilocular; 1 ϫ 0.4Ð0.5 cm; with deciduous hilum; slightly mounded; dissimilar color from testa; corolla; with deciduous calyx; without orifice formed darker than testa; black; not within corona, halo, or rim. by curving of fruit or fruit segments; straight; not Endosperm thin; covering entire embryo; adnate to plicate; not twisted; asymmetrical; oblong; when testa. Cotyledons smooth; both outer faces convex; both asymmetrical with both sutures nearly straight; not the same thickness; both more or less of equal length; inflated; compressed; without beak; rounded at apex; not folded; margin entire 180 degrees from base of apex right-angled with longitudinal axis of fruit; radicle; similar at apex; partially concealing radicle; rounded at base; base oblique with longitudinal axis of split over radicle; with lobes; with lobes touching fruit; with the apex and base uniform in texture; (auriculate); with the interface division terminating at ligneous; seed chambers externally invisible. Fruit base of radicle; without margins recessed; yellow or margin not constricted; without sulcus; plain. Fruit brown; inner face flat; glabrous around base of radicle. wings absent. Fruit nonstipitate. Fruit with all layers Embryonic nearly axis straight; parallel to length of dehiscing; splitting along sutures. Dehiscence of valves seed. Radicle bulbose; straight with embryonic axis; along both sutures; apical and down; passive. Replum centered between cotyledons; less than 1/2 length of invisible. Epicarp glossy; multicolored; bichrome (outer cotyledons. Plumule well developed; glabrous. margin of valve greenish-brown surrounding a tan inner area); glabrous; without spines; not smooth; with Distribution: Australia (northern Queensland). elevated features; reticulately veined (faintly); not tuberculate; faintly wrinkled; not exfoliating; without Notes: At a magnification of ϫ 10, the seed coat of cracks. Mesocarp thick; surface not veined; 1-layered; Plagiocarpus appears to be smooth. At ϫ 50 (fig. C) without balsamic vesicles; without fibers; solid; the surface of the testa appears to be minutely pitted; ligneous (sub). Endocarp dull; monochrome; tan; and at ϫ 1000 (fig. D) the surface cells of the testa have smooth; nonseptate; chartaceous; not exfoliating; thick lateral walls and the lumen of the cell is a pit. remaining fused to epicarp. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; curved. Aril fleshy; cupshaped (if funiculus absent; if funiculus is present, then aril center plugged by a curved, thick, stalklike funiculus); covering less than 1/2 of seed; tan.

Seed 4.5Ð6.5 ϫ 3Ð3.8 ϫ 1.5Ð3 mm; not overgrown; not angular; asymmetrical; elliptic; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; greenish to reddish or dark brown; glabrous; smooth (at ϫ 10, see Notes); coriaceous. Fracture lines absent. Rim absent. Raphe from hilum through lens to base of seed Plagiocarpus: P. axillaris G. Bentham (AÐE). A, Fruit with and terminating to from hilum to lens (and just beyond calyx and valves (ϫ 3.4); B, seeds (ϫ 4.3); CÐD, testa lens); not bifurcating; color of, lighter than, or darker (ϫ 50, ϫ 1000); E, embryos (ϫ 6). than testa (color of raphe may vary: Between hilum and

945 Genus: Hovea R. Brown ex W.T. Aiton mm long; with curved outline; elliptic; marginal according to radicle tip; flush; not within corona, halo, Phylogenetic Number: 23.04. or rim. Lens discernible; less than 0.5 mm in length; with margins curved; circular; not in groove of raphe; Tribe: Bossiaeeae. adjacent to hilum; 1 mm from hilum; mounded; similar color as testa; darker than testa; brown; not within Group: Templetonia. corona, halo, or rim. Endosperm thin; covering entire embryo; adnate to testa. Cotyledons smooth; both outer Species Studied—Species in Genus: 11 spp.—ca. 20 spp. faces convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 Fruit a legume; unilocular; 1.2 ϫ 2.1 ϫ 0.35Ð0.5 cm; with degrees from base of radicle; similar at apex; partially deciduous corolla; with deciduous calyx; without concealing radicle; split over radicle; with lobes; with orifice formed by curving of fruit or fruit segments; lobes touching (auriculate); with the interface division straight; not plicate; not twisted; asymmetrical; circular terminating at base of radicle; without margins re- (subcircular); when asymmetrical with both sutures cessed; yellow; inner face flat; glabrous around base of parallelly curved; not inflated; compressed; without radicle. Embryonic axis straight; oblique to length of beak; rounded at apex; apex aligned with longitudinal seed. Radicle bulbose; oblique to cotyledons; centered axis of fruit; rounded at base; base aligned with between cotyledons; less than 1/2 length of cotyledons. longitudinal axis of fruit; with the apex and base Plumule moderately developed; glabrous. uniform in texture; coriaceous; seed chambers externally invisible. Fruit margin not constricted; without Distribution: Eastern to southwestern Australia (Queensland sulcus; plain. Fruit wings absent. Fruit nonstipitate. to Tasmania). Fruit with all layers dehiscing; splitting along sutures. Dehiscence of valves along both sutures; apical and Notes: Polhill (1981n) listed ca. 12 species, Ross (1988) down; active; with valves enrolling. Replum invisible. listed ca. 20 species, and we are following Ross’s Epicarp dull; monochrome; brown; glabrous or pubes- count. Polhill (1976) provided a full-page plate of cent and indurate; with 1 type of pubescence; puberu- external and internal seed characters of Hovea lent or villous; with pubescence golden; with simple heterophylla A. Cunningham ex J.D. Hooker (now H. hairs; pliable; with hair bases plain; eglandular; without linearis (J. Smith) R. Brown). Valves of Hovea fruits spines; not veined; not tuberculate; not exfoliating; curve inward from the apex to the base, bringing both without cracks. Mesocarp thin; surface not veined; 1- closer together because they are wider than they are layered; without balsamic vesicles; without fibers; long. These fruit characters are unusual legume solid; coriaceous. Endocarp dull; monochrome; tan; characters because most faboid valves curve inward smooth; nonseptate; chartaceous; not exfoliating; from their sides, leaving the apex and base in the same remaining fused to epicarp; entire. Seed 1; length relative position, and are usually much longer than parallel with fruit length. Funiculus less than 0.5 mm wide. The seeds of Hovea species have a surprisingly long; thick; curved. Aril fleshy or dry (rarely); when large hilum. The hilum length:seed length ratio is fleshy cupshaped; entire; covering less than 1/2 of seed; similar to the ratios seen in seeds of some genera in when dry cupshaped; entire; covering less than 1/2 of tribe Phaseoleae (10) and Lamprolobium (23.02). seed; brown or tan.

Seed 4Ð6 ϫ 4Ð5 ϫ 2Ð2.5 mm; not overgrown; not angular; asymmetrical; elliptic to oblong; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe from hilum through lens to base of seed and terminating; not bifurcating; darker than testa. Hilum partially concealed; concealed by aril; with Hovea: H. elliptica A.-P. de Candolle (CÐE), H. spp. (AÐB). faboid split; with the lips of the faboid split the same A, Fruits and valves (ϫ 2.1); B, seeds (ϫ 5); CÐD, testa color as the rest of the hilum; larger than punctiform; 2 (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

947 Genus: Goodia R.A. Salisbury recessed; not within corona, halo, or rim. Lens not discernible. Endosperm thin; covering entire embryo; Phylogenetic Number: 23.05. adnate to testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of Tribe: Bossiaeeae. equal length; not folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing Group: Bossiaea. radicle; entire over radicle; without lobes; with the interface division terminating at base of radicle; with 1 Species Studied—Species in Genus: 2 spp.—2 spp. or both margins recessed; yellow; inner face flat; glabrous around base of radicle. Embryonic axis Fruit a legume; unilocular; 0.9Ð3.8 ϫ 0.55Ð0.8 ϫ 0.2Ð0.3 deflexed; oblique to length of seed. Radicle bulbose or (estimated) cm; with deciduous corolla; with deciduous linear; deflexed and parallel to cotyledon width; calyx; without orifice formed by curving of fruit or fruit centered between cotyledons; 1/2 to nearly length of segments; straight; not plicate; not twisted; asymmetri- cotyledons. Plumule rudimentary or moderately cal; oblong or rhombic; when asymmetrical with 1 developed; glabrous. straight and 1 curved suture or both sutures parallelly curved; widest near middle or D-shaped; not inflated; Distribution: Southern Australia including Tasmania. flattened; without beak; short tapered at apex; apex aligned with longitudinal axis of fruit; short tapered at Notes: Ross (1997) monographed the genus and recognized base; base aligned with longitudinal axis of fruit; with two species. The only reliable character separating the the apex and base uniform in texture; chartaceous; seed two species is the length of the aril foot at its point of chambers externally visible; with the raised seed attachment to the seed—up to 1.1 mm long in G. chambers not torulose. Fruit margin not constricted; medicagnea and 1.4Ð2 mm long in Goodia lotifolia G. without sulcus; plain or embellished; with thickened Salisbury. Polhill (1976) provided a full-page plate of sutural areas (margin thickened). Fruit wings absent. external and internal seed characters of G. lotifolia. Fruit stipitate or substipitate; with the stipe up to 5 mm long. Fruit with all layers dehiscing; splitting along sutures. Dehiscence of valves along both sutures; apical and down; passive. Replum invisible. Epicarp dull; monochrome; reddish brown; glabrous; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1- layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth; nonseptate; coriaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð4; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 1.5 mm long; of 1 length only; thick; hooked. Aril fleshy; hooked; covering less than 1/2 of seed; tan or brown.

Seed 2.7Ð4 ϫ 2.2Ð3 ϫ 2.2Ð3 mm; not overgrown; not angular; asymmetrical; oblong; terete; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; black; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum fully concealed; concealed by aril; with faboid split; with the Goodia: G. medicagnea F.H.J. von Mueller (CÐE), G. spp. lips of the faboid split the same color as the rest of the (AÐB). A, Fruits (closed and dehisced) (ϫ 1.7); B, seeds hilum; punctiform; marginal according to radicle tip; (ϫ 5.1); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

949 Genus: Bossiaea E.P. Ventenat Seed 3Ð3.5 ϫ 2.3Ð2.5 ϫ 1.8Ð2 mm; not overgrown; not angular; symmetrical or asymmetrical; elliptic to ovate; Phylogenetic Number: 23.06. compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering Tribe: Bossiaeeae. to endocarp; dull; not modified by a bloom; colored; monochrome or mottled; with frequent mottles; dark Group: Bossiaea. brown to brown; with black overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings Species Studied—Species in Genus: 15 spp.—ca. 40 spp. absent. Raphe not visible. Hilum fully concealed; concealed by aril; without faboid split; larger than Fruit a legume; unilocular; 1.2Ð1.7 ϫ 0.7Ð1 ϫ 0.15Ð0.25 punctiform; with curved outline; elliptic; marginal cm; with deciduous corolla; with deciduous calyx; according to radicle tip; not within corona, halo, or rim. without orifice formed by curving of fruit or fruit Lens not discernible (may be concealed by aril). segments; straight; not plicate; not twisted; asymmetri- Endosperm thin; covering entire embryo; adnate to cal or symmetrical; oblong, circular, or linear; when testa. Cotyledons smooth; both outer faces convex; both asymmetrical with 1 straight and 1 curved suture or the same thickness; both more or less of equal length; both sutures parallelly curved; widest near middle or D- not folded; margin entire 180 degrees from base of shaped; not inflated; flattened; without or with beak; radicle; similar at apex; partially concealing radicle; straight; with solid beak the same color and texture as split over radicle; with lobes; with lobes not touching; fruit; short tapered to rounded at apex; apex aligned with the interface division terminating at base of with longitudinal axis of fruit; short tapered at base; radicle; without margins recessed; yellow; inner face base aligned with longitudinal axis of fruit; with the flat; glabrous around base of radicle. Embryonic axis apex and base uniform in texture; coriaceous to deflexed; oblique to length of seed. Radicle linear; chartaceous; seed chambers externally visible or deflexed and parallel to cotyledon width; centered invisible; with the raised seed chambers not torulose. between cotyledons; less than 1/2 length of cotyledons. Fruit margin not constricted; without sulcus; plain or Plumule rudimentary; glabrous. embellished; with thickened sutural areas (thick or keeled especially upper suture). Fruit wings absent. Distribution: Temperate Australia, including Tasmania. Fruit stipitate to substipitate to nonstipitate; with the stipe to 5 mm long. Fruit with all layers dehiscing; splitting along sutures. Dehiscence of valves along both sutures; apical and down (assumed); active or passive; with valves revolute. Replum invisible. Epicarp dull; monochrome; black, brown, or gray; glabrous or pubescent and indurate; villous; with pubescence golden or gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; without spines; not smooth; with elevated features; veined or not veined; not tuberculate; striate (at right angles to fruit length); not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; brown; spongy (corklike); septate or nonseptate; with septa thicker than paper, firm; with septa eglandular; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð2(Ð3); length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; straight. Aril fleshy; flat from apex to near base; covering less than 1/2 of seed; tan. Bossiaea: B. cinerea R. Brown (CÐE), B. spp. (AÐB). A, Fruits (dehisced and closed) (ϫ 1.5); B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

951 Genus: Platylobium J.E. Smith coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum fully concealed; Phylogenetic Number: 23.07. concealed by aril; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; Tribe: Bossiaeeae. larger than punctiform; 1.3 mm long; with straight outline; oblong; marginal according to radicle tip; flush; Group: Bossiaea. not within corona, halo, or rim. Lens discernible; less than 0.5 mm in length; with margins curved; circular; Species Studied—Species in Genus: 4 spp.—4 spp. not in groove of raphe; confluent with hilum (at least hilar rim); flush; dissimilar color from testa; darker than Fruit a legume; unilocular; 1.3Ð5.5 ϫ 0.7Ð2 ϫ 0.2 cm; with testa; black; not within corona, halo, or rim. Endosperm deciduous corolla; with persistent calyx; with calyx thin; covering entire embryo; adnate to testa. Cotyle- shorter than fruit; without orifice formed by curving of dons smooth; both outer faces convex; both the same fruit or fruit segments; straight; not plicate; not twisted; thickness; both more or less of equal length; not folded; asymmetrical; when asymmetrical with 1 straight and 1 margin entire 180 degrees from base of radicle; similar curved suture; widest near middle or D-shaped; not at apex; not concealing radicle; entire over radicle; inflated; flattened; without beak; rounded or truncate at without lobes; with the interface division terminating at apex; apex oblique with longitudinal axis of fruit; base of radicle; without margins recessed; yellow; inner truncate at base; base oblique with longitudinal axis of face flat; glabrous around base of radicle. Embryonic fruit; with the apex and base uniform in texture; axis deflexed; oblique to length of seed. Radicle linear; chartaceous; seed chambers externally visible; with the deflexed and parallel to cotyledon length; centered raised seed chambers not torulose. Fruit margin not between cotyledons; less than 1/2 length of cotyledons. constricted; without sulcus; plain or embellished; with Plumule rudimentary; glabrous. wings. Fruit wings 2Ð5; 2Ð3 mm wide; sutural; on 1 suture (upper). Fruit stipitate to substipitate; with the Distribution: Eastern and southern Australia, including stipe 0.5Ð16 mm long. Fruit with all layers dehiscing. Tasmania. Dehiscence of valves along both sutures; active; with valves revolute. Replum invisible. Epicarp dull; Notes: Ross (1983) monographed the genus. monochrome; dark reddish brown; glabrous or pubescent and indurate; puberulent; with pubescence gray; with pubescence uniformly distributed; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; chartaceous. Endocarp dull; monochrome; brown and tan (especially dark- brown beneath seeds and tan along wing); smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds (1Ð)2Ð8; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 1 mm long; of 1 length only; thick; curved. Aril fleshy; hooked; covering less than 1/2 of seed; tan.

Seed 2.5Ð4 ϫ 1.6Ð2.5 ϫ 1.5Ð1.7 mm; not overgrown; not angular; asymmetrical; elliptic or ovate; compressed; without visible radicle and cotyledon lobes; with umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled; Platylobium: P. formosum J.E. Smith (CÐE), P. spp. (AÐB). with frequent mottles; reddish to dark (nearly black) A, Fruits (closed and dehisced) (ϫ 1.3); B, seeds reddish brown; with black overlay; glabrous; smooth; (ϫ 4.5); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

953 Genus: Muelleranthus J. Hutchinson lips of the faboid split the same color as the rest of the hilum; punctiform; marginal according to radicle tip; Phylogenetic Number: 23.08. recessed; within rim. Hilum rim color darker than testa. Lens discernible; less than 0.5 mm in length; not in Tribe: Bossiaeeae. groove of raphe; adjacent to hilum; 0.2 mm from hilum; mounded; dissimilar color from testa; darker than testa; Group: Bossiaea. black (ish); within rim. Lens rim color darker than testa. Endosperm thin; covering entire embryo; adnate to Species Studied—Species in Genus: 2 spp.—3 spp. testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; Fruit a legume; unilocular; (13Ð)17Ð25(Ð29) ϫ 0.4Ð0.5 not folded; margin entire 180 degrees from base of (Ð0.6) ϫ 0.12Ð0.2 cm; with deciduous corolla; with radicle; similar at apex; not concealing radicle; entire deciduous calyx; without orifice formed by curving of over radicle; without lobes; with the interface division fruit or fruit segments; straight; not plicate; not twisted; terminating at base of radicle; without margins re- asymmetrical or symmetrical; linear; when asymmetri- cessed; yellow; glabrous around base of radicle. cal with both sutures nearly straight; not inflated; Embryonic axis deflexed; oblique to length of seed. flattened; without beak; short tapered at apex; apex Radicle linear; deflexed and parallel to cotyledon aligned with longitudinal axis of fruit; short tapered at length; centered between cotyledons; less than 1/2 base; base aligned with longitudinal axis of fruit; with length of cotyledons. Plumule moderately developed; the apex and base uniform in texture; coriaceous; seed glabrous. chambers externally visible; with the raised seed chambers not torulose. Fruit margin not constricted; Distribution: Australia. without sulcus; plain. Fruit wings absent. Fruit substipitate. Fruit with all layers dehiscing; splitting Notes: Lee (1973) noted that M. trifoliatus (F.H. Mueller) J. along sutures. Dehiscence of valves along both sutures; Hutchinson ex A.T. Lee has at first a “reduced hooded apical and down; passive. Replum invisible. Epicarp aril as in Aenictophyton” (23.10) and later is “finally dull; monochrome; reddish brown; glabrous; without not arillate.” She also noted that M. stipularis seeds are spines; not smooth; with elevated features; reticulately not arillate. Based on our observations the seeds of both veined; not tuberculate; not exfoliating; without cracks. species of Muelleranthus and the one species of Mesocarp thin; surface not veined; 1-layered; without Aenictophyton have a dry rim aril. Perhaps Lee thought balsamic vesicles; without fibers; solid; coriaceous. the curved funiculus was a “hooded aril” that disap- Endocarp dull; monochrome; tan; smooth; nonseptate; peared on maturation. Fruits and seeds of M. crenulatus chartaceous; not exfoliating; remaining fused to A.T. Lee are unknown. The attractive seeds of M. epicarp; entire. Seeds 6Ð10; length parallel with fruit stipularis and M. trifoliatus resemble a stained glass length (because seeds are round, both length or width window. Similar colors may be found in seeds of some can be considered parallel to fruit length); neither species of Vicia (19.01). overlapping nor touching; in 1 series. Funiculus of 1 length only; thick; curved. Aril dry; rim-aril; white.

Seed 2Ð2.2 ϫ 2Ð2.2 ϫ 1.5 mm; not overgrown; not angular; symmetrical; circular; terete; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; mottled and streaked or monochrome (when immature); with frequent mottles; with frequent streaks; tan; with orange, yellow, brown, red (the orange overlay forming a reticulate pattern from hilum to base, the brown overlay reddish, and the red overlay with a brick tone), or black (when fully mature (Lee 1973)) overlay; glabrous; smooth; coriaceous. Fracture lines Muelleranthus: M. stipularis (J.M. Black) A.T. Lee (AÐE). absent. Rim absent. Raphe not visible. Hilum partially A, Fruits (dehisced and closed) (ϫ 2); B, seeds (ϫ 4.8); concealed; concealed by aril; with faboid split; with the CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

955 Genus: Ptychosema G. Bentham ex J. Lindley colored halo); within halo. Lens halo color lighter than testa. Endosperm thin; covering entire embryo; adnate Phylogenetic Number: 23.09. to testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal Tribe: Bossiaeeae. length; not folded; margin not entire 180 degrees from base of radicle; similar at apex; not concealing radicle; Group: Bossiaea. entire over radicle; without lobes; with the interface division terminating at base of radicle; without margins Species Studied—Species in Genus: 1 sp.—2 spp. recessed; yellow; inner face flat; glabrous around base of radicle. Embryonic axis deflexed; oblique to length Fruit a legume; unilocular; 1.4Ð2.2 ϫ 0.45Ð0.5 ϫ 0.08 cm; of seed. Radicle linear; deflexed and parallel to cotyle- with deciduous corolla; with persistent calyx; with don length; centered between cotyledons; less than 1/2 calyx shorter than fruit; straight; not plicate; not length of cotyledons. Plumule rudimentary; glabrous. twisted; symmetrical; oblong; not inflated; flattened; without beak; short tapered or emarginate at apex; apex Distribution: Central and Western Australia. aligned with longitudinal axis of fruit; rounded at base; base aligned or oblique with longitudinal axis of fruit; Notes: The fruits and seeds of the type species, Ptychosema with the apex and base uniform in texture; coriaceous; pusillum G. Bentham, are not known to science (Lee seed chambers externally visible (faintly) or invisible; 1973). with the raised seed chambers not torulose. Fruit margin not constricted; without sulcus; plain. Fruit wings absent. Fruit stipitate to substipitate; with the stipe 4Ð7 mm long. Replum invisible. Epicarp dull; multicolored; mottled and streaked; tan; with brown (reddish) overlay; glabrous; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 2Ð7; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril dry; tongue-aril; white.

Seed 2.3Ð2.5 ϫ 1.7Ð2 ϫ 0.5Ð0.7 mm; not overgrown; not angular; symmetrical (except for hilum); elliptic to oblong; compressed; with surface smooth; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; black (blackish-brown when mature) or green (greenish- brown when immature); glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum fully concealed; concealed by aril; without faboid split; punctiform; marginal according to radicle tip; flush; within halo. Hilum halo color lighter than testa. Lens faintly discernible; less than 0.5 mm in length; with margins curved; circular; not in groove of Ptychosema: P. anomalum G. Bentham (AÐE). A, Fruit raphe; adjacent to hilum; 0.1Ð0.2 mm from hilum; (ϫ 3.2); B, seed (ϫ 5); CÐD, testa (ϫ 50, ϫ 1000); barely mounded; similar color as testa (but in lighter E, embryos (ϫ 10).

957 Genus: Aenictophyton A. Lee embryo; adnate to testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or Phylogenetic Number: 23.10. less of equal length; not folded; margin entire 180 degrees from base of radicle; similar at apex; not Tribe: Bossiaeeae. concealing radicle; entire over radicle; without lobes; with the interface division terminating at base of Group: Bossiaea. radicle; without margins recessed; yellow; inner face flat; glabrous around base of radicle. Embryonic axis Species Studied—Species in Genus: 1 sp.—1 sp. deflexed; oblique to length of seed. Radicle linear; oblique to cotyledons; centered between cotyledons; Fruit a legume; unilocular; 1.5Ð1.7 ϫ 0.4Ð0.5 ϫ 0.1Ð0.2 less than 1/2 length of cotyledons. Plumule rudimen- cm; with deciduous corolla; with deciduous calyx; tary; glabrous. without orifice formed by curving of fruit or fruit segments; straight; not plicate; not twisted; asymmetri- Distribution: Australia (Northern Territory and Western cal; oblong; when asymmetrical with both sutures Australia). parallelly curved; not inflated; flattened; without beak; short tapered at apex; apex aligned with longitudinal Notes: Lee (1973) founded the genus and related it to other axis of fruit; tapered at base; base aligned with longitu- genera in the Bossiaeae. Only two seed-fruit samples dinal axis of fruit; with the apex and base uniform in were available for our study, and both were consistent texture; coriaceous; seed chambers externally invisible. with each other and with Lee’s limited seed-fruit data. Fruit margin not constricted; without sulcus; plain. Fruit wings absent. Fruit stipitate; with the stipe up to 10 mm long. Fruit with all layers dehiscing; splitting along sutures. Dehiscence of valves along both sutures; passive. Replum invisible. Epicarp dull; monochrome; reddish brown to brown; glabrous; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp present (and reddish-brown); thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; white (ish) or red (dish-brown because whitish thin endocarp does not conceal mesocarp color); smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1Ð4; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril dry; hooded; white.

Seed 4Ð5 ϫ 1.5Ð2.5 ϫ 1.5Ð2.5 mm; not overgrown; not angular; symmetrical; circular to oblong; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; black (ish-brown); glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum fully concealed; concealed by aril; without faboid split; punctiform; marginal according to radicle tip; flush; not within corona, halo, or rim. Lens Aenictophyton: A. reconditum A.T. Lee (AÐE). A, Fruits not discernible. Endosperm thin; covering entire (ϫ 3.7); B, seeds in situ (ϫ 4.7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

959 Mirbelieae (24.01Ð24.26) colored; monochrome or mottled; with frequent mottles; reddish brown, tan (reddish), orange, or black; Genus: Gompholobium J.E. Smith with black overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe Phylogenetic Number: 24.01. not visible. Hilum partially concealed; concealed by aril; with faboid split; with the lips of the faboid split Tribe: Mirbelieae. the same color as the rest of the hilum; larger than punctiform; 0.7 mm long; with curved outline; oval; Species Studied—Species in Genus: 16 spp.—ca. 35 spp. marginal according to radicle tip; recessed; within rim. Hilum rim color of testa. Lens discernible; less than 0.5 Fruit a legume; unilocular; 0.4Ð2 ϫ 0.4Ð1.3 ϫ 0.3Ð1 cm; mm in length; with margins curved; circular; not in with persistent or deciduous calyx; with calyx shorter groove of raphe; confluent with hilum; mounded; same than fruit; without orifice formed by curving of fruit or color as, similar color as, or dissimilar color from testa; fruit segments; straight; not plicate; not twisted; darker than testa; reddish brown or black; not within asymmetrical or symmetrical; circular or oblong; when corona, halo, or rim. Endosperm thin; covering entire asymmetrical with both sutures parallelly curved or embryo; adnate to testa or embryo. Cotyledons smooth; both sutures nearly straight; not inflated; terete; without both outer faces convex; both the same thickness; both beak; rounded at apex; apex oblique with longitudinal more or less of equal length; not folded; margin entire axis of fruit; rounded at base; base aligned with 180 degrees from base of radicle; similar at apex; not longitudinal axis of fruit; with the apex and base concealing radicle; split over radicle; with lobes; with uniform in texture; coriaceous; seed chambers exter- the interface division terminating at base of radicle; nally invisible. Fruit margin not constricted; without without margins recessed; pale tan or brown (reddish); sulcus; plain. Fruit wings absent. Fruit substipitate. inner face flat; glabrous around base of radicle. Embry- Fruit with all layers dehiscing; splitting along sutures. onic axis deflexed; oblique to length of seed; without a Dehiscence of valves along both sutures or 1 suture; joint evident between the radicle and the cotyledons. apical and down; passive. Replum invisible. Epicarp Radicle bulbose; lobe tip straight; deflexed and parallel dull; monochrome; black; glabrous or glabrate (with to cotyledon length; centered between cotyledons; less hairs along sutures); with 1 type of pubescence; than 1/2 length of cotyledons. Plumule moderately or puberulent; with pubescence gray; with pubescence well developed; glabrous. uniformly distributed; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated Distribution: Australia and New Guinea (G. papuanum E.D. features; not veined; not tuberculate; not exfoliating; Merrill & L.M. Perry). without cracks. Mesocarp thin; surface not veined; 1- layered; without balsamic vesicles; without fibers; Notes: Crisp and Weston (1987) analyzed the Mirbelieae, solid; coriaceous. Endocarp dull or glossy; mono- and we are following their species counts. We did not chrome or streaked; tan; with streaking (dark); with treat the following new taxa of Crisp and Weston: the brown overlay; smooth; nonseptate; chartaceous; not Pultenaea incurvata A. Cunningham group of five exfoliating; remaining fused to mesocarp and epicarp; species; Pultenaea neurocalyx P.K.N.S. Turczanivow, a entire. Seeds 6Ð12; length transverse to fruit length; single species; Aotus phyliocides G. Bentham, a single overlapping or touching; in 1 series. Funiculus mea- species; and Oxylobium microphyllum G. Bentham, a sured; up to 3 mm long; of 1 length only; thick; S- two-species genus. The genus Cupulanthus J. curved or hooked. Aril absent or present; dry; well Hutchinson is listed on page 85 of Crisp and Weston developed rim-aril; reddish brown. (1987) and not mentioned by them again. Crisp and Weston (1995) proposed the following two major Seed 1.6Ð3 ϫ 1.2Ð2.5 ϫ 1Ð2.1 mm; not overgrown; not generic changes: the resurrection of Podolobium R. angular; asymmetrical; mitaform, reniform, or circular; Brown with six species of Oxylobium (24.09, see Notes compressed or terete; with surface smooth; without for Oxylobium) and a new genus, tentatively to be visible radicle and cotyledon lobes; without hilar sinus; named Otion. Otion, as projected, will have six species, without umbo on seed faces. Cuticle inflated (like two new ones and four from four different genera: bubbles) or not inflated (around hilum). Testa not Aotus phylicoides G. Bentham, Burtonia simplicifolia adhering to endocarp; dull; not modified by a bloom; F. von Mueller & R. Tate, Oxylobium microphyllum G. Bentham, and Phyllota luehmannii F. von Mueller.

960 Crisp and Weston (1987), in Appendix II, united Burtonia R. Brown with Gompholobium under the latter name.

961 Gompholobium: G. latifolium J.E. Smith (E), G. scabra R. Brown (CÐD), G. spp. (AÐB). A, Fruits with and without calyx and valves (ϫ 2.3); B, seeds (ϫ 10); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 12).

963 Genus: Sphaerolobium J.E. Smith Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by aril; with faboid split; Phylogenetic Number: 24.02. with the lips of the faboid split the same color as the rest of the hilum; punctiform; marginal according to Tribe: Mirbelieae. radicle tip; flush; not within corona, halo, or rim or within rim. Hilum rim color of testa. Lens discernible; Species Studied—Species in Genus: 7 spp.—ca. 15 spp. less than 0.5 mm in length; with margins straight or curved; circular or oblong; not in groove of raphe; Fruit a legume; unilocular; 0.4Ð0.55 ϫ 0.3Ð0.4 ϫ 0.3Ð0.4 adjacent to or confluent with hilum; 0.2 mm from cm; with deciduous or persistent calyx; with calyx hilum; flush; dissimilar color from testa; darker than shorter than fruit; without orifice formed by curving of testa; black; not within corona, halo, or rim. Endosperm fruit or fruit segments; straight; not plicate; not twisted; thin; covering entire embryo; adnate to testa. Cotyle- asymmetrical or symmetrical; more or less circular or dons smooth; both outer faces convex; both the same oblong; when asymmetrical with 1 straight and 1 thickness; both more or less of equal length; not folded; curved suture or both sutures parallelly curved; widest margin entire 180 degrees from base of radicle; similar near middle or D-shaped; not inflated; terete; without at apex; not concealing radicle; split over radicle; with beak; rounded at apex; apex aligned with longitudinal lobes; with the interface division terminating at base of axis of fruit; rounded at base; base aligned with radicle; without margins recessed; yellow; inner face longitudinal axis of fruit; with the apex and base flat; glabrous around base of radicle. Embryonic axis uniform in texture; ligneous; seed chambers externally deflexed; oblique to length of seed; without a joint invisible. Fruit margin not constricted; without sulcus; evident between the radicle and the cotyledons. Radicle plain. Fruit wings absent. Fruit substipitate or stipitate; linear; deflexed and parallel to cotyledon length; with the stipe up to 5 mm long. Fruit with all layers centered between cotyledons; less than 1/2 length of dehiscing; splitting along sutures. Dehiscence of valves cotyledons. Plumule rudimentary; glabrous. along both sutures; apical and down; passive. Replum invisible. Epicarp dull; monochrome; reddish brown, Distribution: Southwestern and southeastern Australia. tan (to greenish or barely reddish tan), green, or black; glabrous; eglandular; without spines; not smooth; with Notes: The fruits of Sphaerolobium are slightly to notice- elevated features; veined or not veined; reticulately ably longitudinally compressed so that the long axis is veined; not tuberculate; wrinkled; not exfoliating; equivalent to the width of most other legume fruits. The without cracks. Mesocarp thick; surface not veined; 1- upper fruit in fig. A (of S. grandiflorum G. Bentham) is layered; without balsamic vesicles; without fibers; an example of this compression. solid; sub ligneous. Endocarp glossy; monochrome; reddish brown, tan, or white (nearly); smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with or transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened; straight (from a thickened suture wall). Aril dry; rim-aril; tan.

Seed 1.2Ð1.8 ϫ 1.1Ð1.3 ϫ 0.9 mm; not overgrown; not angular; asymmetrical; reniform, circular, or mitaform; compressed; with surface smooth; without or with visible radicle and cotyledon lobes; without or with external groove between radicle and cotyledon lobes; same color as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish Sphaerolobium: S. vimineum J.E. Smith (CÐE), S. spp. brown; glabrous; smooth or not smooth; with elevated (AÐB). A, Fruits with or without calyx (ϫ 6.9); B, seeds features; tuberculate; coriaceous. Fracture lines absent. (ϫ 10.5); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ13).

965 Genus: Viminaria J.E. Smith Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; not Phylogenetic Number: 24.03. folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing radicle; split over Tribe: Mirbelieae. radicle; with lobes; with the interface division terminating at base of radicle; without margins recessed; pale Species Studied—Species in Genus: 1 sp.—1 sp. tan; inner face flat; glabrous around base of radicle. Embryonic axis deflexed; oblique to length of seed; Fruit a legume; unilocular; 0.9Ð1 ϫ 0.5Ð0.6 ϫ 0.3 cm; with without a joint evident between the radicle and the persistent calyx; with calyx shorter than fruit (3/4 cotyledons. Radicle linear; deflexed and parallel to length of fruit); without orifice formed by curving of cotyledon width; centered between cotyledons; less fruit or fruit segments; straight; not plicate; not twisted; than 1/2 length of cotyledons. Plumule rudimentary; asymmetrical; oblong or ovate; when asymmetrical glabrous. with both sutures parallelly curved; not inflated; compressed; without beak; rounded at apex; apex Distribution: Southwestern and southeastern Australia. oblique with longitudinal axis of fruit; rounded at base; base slightly oblique with longitudinal axis of fruit; with the apex and base uniform in texture; membranous; seed chambers externally invisible. Fruit margin not constricted; without sulcus; plain. Fruit wings absent. Fruit nonstipitate. Fruit indehiscent. Replum invisible. Epicarp dull; monochrome; black; glabrous; eglandular; without spines; not smooth; with elevated features; not veined; not tuberculate; wrinkled (especially portion above calyx); not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome (but epicarp color is visible); tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp. Seed 1; length parallel with fruit length. Funiculus less than 0.5 mm long; flattened; straight. Aril dry; rim-aril; tan.

Seed 2Ð3 ϫ 1.4Ð2 ϫ 1Ð1.8 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim present. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.5 mm long; with curved outline; circular; between cotyledon and radicle lobe; recessed; not within corona, halo, or rim. Lens discernible; less than 0.5 mm in length (0.3Ð0.4 mm); with margins curved; circular; not in groove of raphe; adjacent to hilum; 0.1 mm from hilum; slightly mounded; dissimilar color from testa; darker than testa; Viminaria: V. juncea (H.A. Schader) J.C. von Hoffmann- black; not within corona, halo, or rim. Endosperm segg (AÐE). A, Fruits with calyx (ϫ 5.6); B, seeds thick; covering entire embryo; adnate to embryo. (ϫ 10); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 16).

967 Genus: Daviesia J.E. Smith oblong or elliptic; compressed; with surface smooth; without visible radicle and cotyledon; without hilar Phylogenetic Number: 24.04. sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; Tribe: Mirbelieae. monochrome or mottled and streaked; with frequent mottles; with frequent streaks; brown to greenish or Species Studied—Species in Genus: 22 spp.—120 spp. reddish brown, tan (to reddish tan), yellow, orange, or black; with black overlay; glabrous; smooth or not Fruit a legume; unilocular; 0.5Ð1.5 ϫ 0.4Ð1 ϫ 0.3Ð0.5 cm; smooth; with recessed features; punctate; coriaceous. with persistent calyx; with calyx shorter than fruit; Fracture lines absent. Rim absent. Wings absent. Raphe without orifice formed by curving of fruit or fruit not visible. Hilum partially concealed; concealed by segments; straight; not plicate; not twisted; more or less aril; with faboid split; with the lips of the faboid split asymmetrical; triangular or irregular; when asymmetri- the same color as the rest of the hilum; larger than cal with both sutures unequally curved, 1 straight and 1 punctiform; 1 mm long; with straight outline; oblong; curved suture, or both sutures parallelly curved; more marginal according to radicle tip; recessed; not within or less widest near middle or D-shaped; not inflated; corona, halo, or rim or within halo. Hilum halo color compressed; without or with beak; with solid beak the (black) darker than testa. Lens discernible or not same color and texture as fruit; tapered, short tapered, discernible; less than 0.5 mm in length; with margins or truncate at apex; apex aligned, oblique, or right- curved; elliptic; not in groove of raphe; confluent with angled with longitudinal axis of fruit; long tapered to hilum (at least with halo); mounded; dissimilar color tapered at base; base aligned or oblique with longitudi- from testa; darker than testa; black; not within corona, nal axis of fruit; with the apex and base uniform in halo, or rim. Endosperm thin; covering entire embryo; texture; membranous or chartaceous. Fruit margin not adnate to testa or embryo. Cotyledons smooth; both constricted; without sulcus; plain. Fruit wings absent. outer faces convex; both more or less of equal length; Fruit substipitate. Fruit with all layers dehiscing; not folded; margin entire 180 degrees from base of splitting along sutures. Dehiscence of valves along both radicle; similar at apex; not concealing radicle; split sutures; apical and down; active; with valves twisting over radicle; with lobes; with the interface division (each valve or both valves forming a cone). Replum terminating at base of radicle; without margins re- invisible. Epicarp dull or glossy; monochrome or cessed; yellow; inner face flat; glabrous around base of multicolored; mottled; light to dark or reddish brown, radicle. Embryonic axis deflexed; oblique to length of tan, green, purple, or gray; with tan overlay; glabrous or seed; without a joint evident between the radicle and pubescent and indurate; with 1 type of pubescence; the cotyledons. Radicle linear; lobe tip straight; villous; with pubescence golden; with pubescence deflexed and parallel to cotyledon length; centered uniformly distributed; with simple hairs; pliable; with between cotyledons; less than 1/2 length of cotyledons. hair bases plain; eglandular; without spines; smooth or Plumule rudimentary; glabrous. not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Distribution: Australia. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Notes: Crisp and Chandler (Crisp 1980, 1982, 1984, 1985, Endocarp dull; monochrome; brown; smooth, hairy 1991; Crisp and Chandler 1997; Chandler and Crisp (crinkled and golden), or cobwebby; nonseptate; 1998) have treated parts of Daviesia. Crisp (1985) dealt chartaceous; not exfoliating; remaining fused to with conservation of the genus. mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with fruit length; overlapping and touching or neither overlapping nor touching; in 1 series. Funiculus measured; 1 mm long; of 1 length only; thick; straight (may be bifurcate at base with seed on each stalk. Aril fleshy; cupshaped (with thicker apex); covering 1/2 to nearly all of seed; reddish brown or yellow. Daviesia: D. acicularis J.E. Smith (CÐE), D. spp. (AÐB). A, Seed 3Ð6 ϫ 1.5Ð3.5 ϫ 0.8Ð2.4 mm; overgrown, 1 seed Fruits and valves with calyx (ϫ 2.9); B, seeds (ϫ 5); filling entire fruit cavity; not angular; asymmetrical; CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

969 Genus: Isotropis G. Bentham not adhering to endocarp; dull; not modified by a bloom; clear; monochrome (though intertices may be Phylogenetic Number: 24.06. darker than ridges); light to dark reddish brown or tan; glabrous; not smooth; with elevated features; reticulate; Tribe: Mirbelieae. coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum partially or fully Species Studied—Species in Genus: 4 spp.—10 spp. concealed; concealed by radicle lobe or wing; with faboid split; with the lips of the faboid split the same Fruit a legume; unilocular; 1.5Ð3 ϫ 0.35Ð0.7 ϫ 0.37 cm; color as the rest of the hilum; punctiform; between with persistent calyx; with calyx shorter than fruit; cotyledon and radicle lobe; flush; within rim. Hilum without orifice formed by curving of fruit or fruit rim color of testa. Lens discernible or not discernible; segments; straight or curved (slightly); not plicate; not less than 0.5 mm in length; with margins curved; more twisted; asymmetrical; falcate, oblong (linear), linear, or less circular; not in groove of raphe; confluent with or lanceolate; when asymmetrical with both sutures hilum; mounded; dissimilar color from testa; darker nearly straight; not inflated; terete; without or with than testa; black; not within corona, halo, or rim. beak; straight or declined; with solid beak the same Endosperm thick; pluglike and resembling tip of color and texture as fruit; tapered or short tapered at radicle; covering entire embryo; adnate to testa or apex; apex aligned, oblique, or right-angled with embryo. Cotyledons smooth; both outer faces convex; longitudinal axis of fruit; short tapered at base; base both the same thickness; both more or less of equal aligned or oblique with longitudinal axis of fruit; with length; not folded; margin entire 180 degrees from base the apex and base uniform in texture; coriaceous; seed of radicle; similar at apex; not concealing radicle; split chambers externally invisible. Fruit margin not con- over radicle; with lobes; with the interface division stricted; without sulcus; plain. Fruit wings absent. Fruit terminating at base of radicle; without margins re- substipitate or nonstipitate. Fruit with all layers cessed; green, brown (greenish), or tan; inner face flat; dehiscing; splitting along sutures. Dehiscence of valves glabrous around base of radicle. Embryonic axis along both sutures; apical and down; active; with valves deflexed; oblique to length of seed; without a joint twisting. Replum invisible. Epicarp dull; monochrome; evident between the radicle and the cotyledons. Radicle tan; pubescent but soon deciduous; with 1 type of linear; lobe tip straight, curved, or hooked; deflexed and pubescence; tomentose or villous; with pubescence parallel to cotyledon length; centered between cotyle- golden or gray; with pubescence uniformly distributed; dons; less than 1/2 length of cotyledons. Plumule with simple hairs; pliable; with hair bases plain; rudimentary; glabrous. eglandular or glandular; with glandular dots (black); without spines; not smooth; with elevated features; not Distribution: Australia. veined; not tuberculate; shagreen; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1- layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds up to 11; length parallel with fruit length; overlapping and touching or neither overlapping nor touching; in 1 series. Funiculus measured; 1 mm long; of 1 length only; filiform; straight or curved. Aril dry; rim-aril; white.

Seed 2Ð3.75 ϫ 1.5Ð3 ϫ 0.8Ð1 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and Isotropis: I. wheeleri G. Bentham (CÐE), I. spp. (AÐB). A, cotyledon lobes; with external groove between radicle Fruits with and without calyx (closed and dehisced) and and cotyledon lobes same color as testa; with deep or valves (ϫ 2.8); B, seeds (ϫ 9.7); CÐD, testa (ϫ 50, shallow hilar sinus; without umbo on seed faces. Testa ϫ 1000); E, embryos (ϫ 10).

971 Genus: Jacksonia R. Brown ex J.E. Smith without umbo on seed faces. Cuticle wrinkled or not wrinkled. Testa not adhering to endocarp; dull; not Phylogenetic Number: 24.08. modified by a bloom; colored; monochrome or mottled and streaked; reddish brown; with black overlay; Tribe: Mirbelieae. glabrous; not smooth; with elevated features; wrinkled; coriaceous. Fracture lines absent. Rim absent. Wings Species Studied—Species in Genus: 10 spp.—ca. 50 spp. absent. Raphe not visible. Hilum partially concealed; concealed by radicle lobe or wing; with faboid split; Fruit a legume; unilocular; 0.5Ð1 ϫ 0.3Ð4.5 ϫ 0.25Ð0.35 with the lips of the faboid split the same color as the cm; with persistent or deciduous calyx; with calyx rest of the hilum; punctiform; between cotyledon and shorter than or equal in length to fruit; without orifice radicle lobe; recessed; not within corona, halo, or rim. formed by curving of fruit or fruit segments; straight or Lens discernible; less than 0.5 mm in length; with curved; not plicate; not twisted; symmetrical or margins curved; circular; not in groove of raphe; asymmetrical; oblong, obovate, or ovate; when asym- adjacent to hilum; 0.3 mm from hilum; mounded; same metrical with 1 straight and 1 curved suture or both color as testa; not within corona, halo, or rim. En- sutures nearly straight; widest near middle or D-shaped; dosperm thick; covering entire embryo; adnate to testa not inflated; terete or compressed; without or with beak; or embryo. Cotyledons smooth; both outer faces straight; with solid beak the same color and texture as convex; both the same thickness; both more or less of fruit; long tapered at apex; apex aligned or oblique with equal length; not folded; margin entire 180 degrees longitudinal axis of fruit; short tapered or long tapered from base of radicle; similar at apex; partially conceal- at base; base aligned or oblique with longitudinal axis ing or not concealing radicle; entire or split over of fruit; with the apex and base uniform in texture; radicle; without or with lobes; with lobes not touching; coriaceous; seed chambers externally invisible. Fruit with the interface division terminating at base of margin not constricted; without sulcus; plain. Fruit radicle; without margins recessed; greenish tan or wings absent. Fruit substipitate to nonstipitate. Fruit green; inner face flat; glabrous around base of radicle. with all layers dehiscing; splitting along sutures. Embryonic axis deflexed; oblique to length of seed; Dehiscence of valves along both sutures; apical and without a joint evident between the radicle and the down; passive. Replum invisible. Epicarp dull; mono- cotyledons. Radicle linear or bulbose; lobe tip hooked; chrome; reddish brown or tan (reddish); pubescent and deflexed and parallel to cotyledon length; centered indurate; with 1 type of pubescence; puberulent or between cotyledons; less than 1/2 length of cotyledons. tomentose; with pubescence gray or golden; with Plumule rudimentary; glabrous. pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without Distribution: Australia. spines; smooth or not smooth; with elevated features; veined or not veined; reticulately veined; not tubercu- Notes: Crisp (1984) presented notes on Jacksonia. late; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; reddish brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with fruit length; overlapping and touching or neither overlapping nor touching; in 1 series. Funiculus measured; 0.7 mm long; of 1 length only; filiform; curved. Aril present or absent; dry; tongue-aril; reddish brown or white.

Seed 2Ð3.5 ϫ 1Ð2.5 ϫ 0.5Ð1.5 mm; not overgrown; not angular or angular; asymmetrical; oblong or reniform; compressed; with surface smooth; with visible radicle Jacksonia: J. ramosissima G. Bentham (CÐE), J. spp. (AÐ and cotyledon lobes; without external groove between B). A, Fruits with calyx and valve (ϫ 3.5); B, seeds (ϫ radicle and cotyledon lobes; without hilar sinus; 10); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

973 Genus: Oxylobium H.C. Andrews radicle and cotyledon lobes; without or with external groove between radicle and cotyledon lobes; with Phylogenetic Number: 24.09. external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo Tribe: Mirbelieae. on seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome or Species Studied—Species in Genus: 13 spp.—ca. 15 spp. mottled and streaked; with frequent mottles; with frequent streaks; brown to dark to light reddish brown Fruit a legume; unilocular; 0.5Ð2 ϫ 0.3Ð0.8 ϫ 0.2Ð0.5 cm; or black; with brown (reddish), tan (reddish), or black with persistent calyx; with calyx shorter than fruit; overlay; glabrous; smooth; coriaceous. Fracture lines without orifice formed by curving of fruit or fruit absent. Rim absent. Wings absent. Raphe not visible. segments; straight or curved; not plicate; not twisted; Hilum visible; with faboid split; with the lips of the asymmetrical or symmetrical; ovate or C-shaped; when faboid split the same color as the rest of the hilum; asymmetrical with 1 straight and 1 curved suture or larger than punctiform; 0.5 mm long; with curved both sutures parallelly curved; widest near middle or D- outline; elliptic or oval; between cotyledon and radicle shaped; not inflated; compressed; without or with beak; lobe; recessed; not within corona, halo, or rim or within straight; with solid beak the same color and texture as halo. Hilum halo color darker or lighter than testa. Lens fruit; tapered or short tapered at apex; apex aligned or not discernible or discernible (faintly); less than 0.5 mm oblique with longitudinal axis of fruit; short tapered at or equal to or greater than 0.5 mm in length; up to 0.5 base; base aligned or oblique with longitudinal axis of mm long; with margins curved; circular or elliptic; not fruit; with the apex and base uniform in texture; in groove of raphe; adjacent to hilum; 0.3Ð0.5 mm from ligneous; seed chambers externally invisible or visible hilum; mounded; same or similar color as testa; not (faintly). Fruit margin not constricted; without sulcus; within corona, halo, or rim. Endosperm thick; covering plain. Fruit wings absent. Fruit substipitate. Fruit with entire embryo; adnate to embryo. Cotyledons smooth; all layers dehiscing; splitting along sutures. Dehiscence both outer faces convex; both the same thickness; both of valves along both sutures; apical and down (or more or less of equal length; not folded; margin entire apically only); passive. Replum invisible. Epicarp dull; 180 degrees from base of radicle; similar at apex; not monochrome; greenish brown, green, or black; pubes- concealing radicle; entire over radicle; without lobes; cent and indurate; with hairs erect or appressed; with 1 with the interface division terminating at base of type of pubescence; villous; with pubescence gray or radicle; without margins recessed; tan to reddish tan or golden; with pubescence uniformly distributed; with green; inner face flat; glabrous around base of radicle. simple hairs; stiff or pliable; with hair bases plain; Embryonic axis deflexed; oblique to length of seed; glandular; with glandular setae (short); without spines; without a joint evident between the radicle and the not smooth; with elevated features; reticulately veined; cotyledons. Radicle linear; lobe tip straight; deflexed not tuberculate; exfoliating in part, exfoliating, or not and parallel to cotyledon length; centered between exfoliating; without cracks. Mesocarp thin; surface not cotyledons; less than 1/2 length of cotyledons. Plumule veined; 1- or 2-layered; without balsamic vesicles; rudimentary; glabrous. without or with fibers; solid; with fibers over solid layer; subligneous. Endocarp glossy; monochrome; tan; Distribution: Australia. smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds Notes: Crisp and Weston (1987) noted that “in its strictest 2Ð10; length parallel with or oblique to fruit length; sense, Oxylobium may include only O. cordifolium neither overlapping nor touching or overlapping and [H.C. Andrews] and O. pultenaea” A.-P. de Candolle. touching; in 1 series. Funiculus measured; 0.7Ð0.8 mm Lebler (1977) described the fruits and seeds of five long; of 1 length only; filiform; curved, S-curved, or species of Oxylobium, also known as shaggy-peas. The straight. Aril dry; tongue-aril; reddish brown, tan (to plants are poisonous to livestock. reddish tan), or red.

Seed 1Ð2.7 ϫ 1.2Ð2 ϫ 1.2Ð1.5 mm; not overgrown; not Oxylobium: O. robustum J. Thompson (CÐE), O. spp. (AÐ angular or angular; asymmetrical; oblong, ovate, B). A, Fruits (dehisced or closed) with or without calyx reniform, mitaform, triangular, or irregular; com- (ϫ 2.5); B, seeds (ϫ 7); CÐD, testa (ϫ 50, ϫ 1000); E, pressed; with surface smooth; with or without visible embryos (ϫ 12).

975 Genus: Chorizema J.J.H. de Labillardière colored; monochrome or mottled and streaked; with frequent mottles; with frequent streaks; brown to Phylogenetic Number: 24.10. greenish to reddish brown, tan, or black; with black overlay; glabrous; smooth or not smooth; with elevated Tribe: Mirbelieae. features; reticulate; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Species Studied—Species in Genus: 21 spp.—25 spp. partially or fully (nearly) concealed; concealed by aril; with faboid split; with the lips of the faboid split the Fruit a legume; unilocular; 0.4Ð1.7 ϫ 0.3Ð0.8 cm; with same color as the rest of the hilum; larger than puncti- persistent or deciduous calyx; with calyx shorter than form; up to 0.5 mm long; with curved outline; oval; fruit; without orifice formed by curving of fruit or fruit marginal according to radicle tip or between cotyledon segments; straight; not plicate; not twisted; asymmetri- and radicle lobe; recessed; within rim. Hilum rim color cal or symmetrical; ovate, obovate, or elliptic; when of testa. Lens discernible; less than 0.5 mm or equal to asymmetrical with 1 straight and 1 curved suture; or greater than 0.5 mm in length; 0.5 mm long; with widest near middle or D-shaped; not inflated; terete or margins straight; linear; not in groove of raphe; compressed; with beak; straight, declined, or hooked; adjacent to hilum; 0.3 mm from hilum; flush; similar with solid beak the same color and texture as fruit; short color as testa; darker than testa; reddish brown; not tapered or tapered at apex; apex aligned or oblique with within corona, halo, or rim. Endosperm thick; covering longitudinal axis of fruit; long tapered, tapered, or short entire embryo; adnate to embryo or testa. Cotyledons tapered at base; base aligned with longitudinal axis of smooth; both outer faces convex; both the same fruit; with the apex and base uniform in texture; thickness; both more or less of equal length; not folded; coriaceous; seed chambers externally invisible or margin entire 180 degrees from base of radicle; similar visible. Fruit margin not constricted; without sulcus; at apex; not concealing radicle; split over radicle; with plain. Fruit wings absent. Fruit stipitate to substipitate lobes; with the interface division terminating at base of to nonstipitate. Fruit with all layers dehiscing; splitting radicle; without margins recessed; yellow or orange; along sutures. Dehiscence of valves along both sutures; inner face flat; glabrous around base of radicle. Embry- apical and down (to somewhat down); passive. Replum onic axis deflexed; oblique to length of seed; without a invisible. Epicarp dull; monochrome; brown; glabrous joint evident between the radicle and the cotyledons. or pubescent and indurate; with hairs appressed; with 1 Radicle linear; lobe tip straight; deflexed and parallel to type of pubescence; with pubescence golden; with cotyledon length; centered between cotyledons; less pubescence uniformly distributed; with simple hairs; than 1/2 length of cotyledons. Plumule rudimentary; pliable; with hair bases plain; eglandular; without glabrous. spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Distribution: Western Australia. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; subligneous. Notes: Taylor and Crisp (1992) revised Chorizema, and our Endocarp glossy; monochrome; reddish brown; smooth; species count agrees with theirs. All fruiting material nonseptate; chartaceous; not exfoliating; remaining available to us was dehisced, therefore we were unable fused to mesocarp and epicarp; entire. Seeds (number to determine the number of seeds per fruit. Taylor and unknown, see Notes) length parallel with fruit length; in Crisp did not report the number of seeds per fruit, but 1 series. Funiculus measured; 0.6 mm long; of 1 length they did report that the ovaries had 8Ð30 ovules and that only; filiform; curved. Aril dry; well developed rim- C. retrorsum J.M. Taylor & M.D. Crisp had as many as aril; reddish brown. 35 ovules per ovary.

Seed 1.5Ð3.5 ϫ 1Ð2 ϫ 1Ð1.3 mm; not overgrown; not angular; asymmetrical; D-shaped, oblong, or reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; Chorizema: C. ilicifolium J.J.H. de Labillardière (CÐE), C. without umbo on seed faces. Testa not adhering to spp. (AÐB). A, Fruits (dehisced) with and without calyx endocarp; glossy or dull; not modified by a bloom; (ϫ 5.3); B, seeds (ϫ 9.2); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

977 Genus: Mirbelia J.E. Smith smooth; with or without visible radicle and cotyledon lobes; without external groove between radicle and Phylogenetic Number: 24.11. cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull or Tribe: Mirbelieae. glossy; not modified by a bloom; colored; monochrome or mottled and streaked; with frequent mottles; with Species Studied—Species in Genus: 8 spp.—15–30 spp. frequent streaks; light reddish to dark reddish brown or black; with purple overlay; glabrous; smooth or not Fruit a legume; unilocular; 0.8Ð2 ϫ 0.3Ð1 ϫ 0.25Ð0.5 cm; smooth; with elevated features; wrinkled and shagreen; with persistent or deciduous calyx; with calyx shorter coriaceous. Fracture lines absent. Rim absent. Wings than fruit; without orifice formed by curving of fruit or absent. Raphe not visible. Hilum visible or partially fruit segments; straight; not plicate; not twisted; concealed; concealed by funicular remnant, radicle symmetrical; ovate or didymous; not inflated; com- lobe, or wing; with faboid split; with the lips of the pressed; with beak; straight; with solid beak the same faboid split the same color as the rest of the hilum; color and texture as fruit; short tapered or rounded at punctiform or larger than punctiform; up to 0.5 mm apex; apex aligned with longitudinal axis of fruit; short long; with curved or angular outline; circular or tapered or rounded at base; base aligned with longitudi- triangular; marginal according to radicle tip or between nal axis of fruit; with the apex and base uniform in cotyledon and radicle lobe; recessed; not within corona, texture; coriaceous; seed chambers externally invisible halo, or rim or within halo. Hilum halo color (reddish- or visible. Fruit margin not constricted or constricted; brown) lighter than testa or of testa. Lens not discern- constricted along both margins; without sulcus; plain. ible or discernible (on light-colored seeds); less than 0.5 Fruit wings absent. Fruit substipitate. Fruit with all mm or equal to or greater than 0.5 mm in length; up to layers dehiscing; splitting along sutures. Dehiscence of 0.5 mm long; with margins straight or curved; linear or valves along both sutures; apical and down or basal and irregular; not in groove of raphe; adjacent to or up; passive. Replum invisible. Epicarp dull; mono- confluent with hilum; 0.2 mm from hilum; flush; chrome; black, brown (reddish), or tan; glabrous or dissimilar color from testa; darker than testa; black; not pubescent and indurate; with hairs erect or appressed; within corona, halo, or rim. Endosperm thick; covering with 1 type of pubescence; puberulent; with pubescence entire embryo; adnate to testa or embryo. Cotyledons golden; with pubescence uniformly distributed; with smooth; both outer faces convex; both the same simple hairs; pliable; with hair bases plain; eglandular; thickness; both more or less of equal length; not folded; without spines; not smooth; with elevated features; not margin entire 180 degrees from base of radicle; similar veined or veined; reticulately veined and transversely at apex; partially concealing or not concealing radicle; veined relative to fruit length; not tuberculate; wrinkled entire or split over radicle; without or with lobes; with and shagreen; not exfoliating or exfoliating; without lobes not touching; with the interface division terminat- cracks. Mesocarp thin; surface not veined; 1- or 2- ing at base of radicle; without margins recessed; tan, layered; without balsamic vesicles; with fibers present brown (reddish), or yellow; inner face flat; glabrous or without fibers; solid; with fibers over solid layer; around base of radicle. Embryonic axis deflexed; coriaceous. Endocarp dull or glossy; monochrome; oblique to length of seed; without a joint evident reddish tan; smooth; with hairs restricted to sutures between the radicle and the cotyledons. Radicle linear; (some with hairs especially along funicular suture); lobe tip curved; deflexed and parallel to cotyledon nonseptate; chartaceous; not exfoliating; remaining width; centered between cotyledons; less than 1/2 fused to mesocarp and epicarp; entire. Seeds 2; length length of cotyledons. Plumule rudimentary; glabrous. parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; up to 2 mm Distribution: Australia. long; of 1 length only; flattened; straight or curved. Aril dry; rim-aril; reddish brown, red, or black. Notes: Crisp and Weston (1987) noted that its circumscription needs further study, especially in relation to Seed 1.5Ð3 ϫ 1Ð2.2 ϫ 0.5Ð1.2 mm; not overgrown; not Chorizema (24.10) and elements of Oxylobium (24.09). angular or angular; asymmetrical; mitaform, oblong, They (Crisp and Weston 1995) have carried out further reniform, rhombic, D-shaped, or irregular; compressed cladistic studies and resurrected Podolobium R. Brown or mounded on 1 side and straight on other side (shape (24.14) with six species that were accepted by Wiecek of valve with intruded lower suture); with surface (1991) as the first six species of Oxylobium. Pending

978 further evaluation by the botanical community, we have left those species in Oxylobium. The fruit has a spurious septum intruding from the lower suture.

979 Mirbelia: M. grandiflora W. Aiton (CÐE), M. spp. (AÐB). A, Fruits with calyx (closed and dehisced) (ϫ 3.2); B, seeds (ϫ 10.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

981 Genus: Callistachys É.P. Ventenat not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the Phylogenetic Number: 24.12. hilum; larger than punctiform; 0.4 mm long; with curved outline; circular; between cotyledon and radicle Tribe: Mirbelieae. lobe; recessed; within halo. Hilum halo color lighter than testa. Lens not discernible or discernible (faintly); Species Studied—Species in Genus: 1 sp.— 1 or 2 spp. less than 0.5 mm in length; with margins curved; circular; not in groove of raphe; adjacent to hilum; 0.3Ð Fruit a legume; unilocular; 1.5Ð2 ϫ 0.5Ð0.6 ϫ 0.4Ð0.45 cm; 0.4 mm from hilum; mounded or flush; similar color as with persistent or deciduous (rarely) calyx; with calyx testa; darker than testa; darker reddish brown; not shorter than fruit; without orifice formed by curving of within corona, halo, or rim. Endosperm thick; covering fruit or fruit segments; straight; not plicate; not twisted; entire embryo; adnate to embryo. Cotyledons smooth; asymmetrical; oblong; when asymmetrical with both both outer faces convex; both the same thickness; both sutures nearly straight; not inflated; terete; without more or less of equal length; not folded; margin entire beak; short tapered at apex; apex aligned with longitu- 180 degrees from base of radicle; similar at apex; not dinal axis of fruit; short tapered at base; base oblique concealing radicle; split over radicle; with lobes; with with longitudinal axis of fruit; with the apex and base the interface division terminating at base of radicle; uniform in texture; ligneous; seed chambers externally without margins recessed; tan; inner face flat; glabrous invisible. Fruit margin not constricted; without sulcus; around base of radicle. Embryonic axis deflexed; plain. Fruit wings absent. Fruit substipitate. Fruit with oblique to length of seed; without a joint evident all layers dehiscing; splitting along sutures. Dehiscence between the radicle and the cotyledons. Radicle linear; of valves along both sutures; apical and down; passive. lobe tip straight; deflexed and parallel to cotyledon Replum invisible. Epicarp dull; monochrome; dark length; centered between cotyledons; less than 1/2 reddish brown; pubescent and indurate; with 1 type of length of cotyledons. Plumule rudimentary; glabrous. pubescence; villous; with pubescence golden; with pubescence uniformly distributed; with simple hairs; Distribution: Southwestern Australia. pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; obliquely Notes: Callistachys, a segregate of Oxylobium (24.09), was veined relative to fruit length; not tuberculate; ribbed; recognized by Crisp and Weston (1987) but not by exfoliating (especially with age over and around ribs); Polhill (1981m). without cracks. Mesocarp thick; surface uniformly veined; 2-layered; without balsamic vesicles; without fibers; with veins over solid layer; ligneous. Endocarp dull; monochrome; tan; scurfy or smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 7; length transverse to fruit length; overlapping or touching; in 1 series. Funiculus measured; 7Ð15 mm long; of 1 length only; flattened; curved. Aril dry; rim-aril; tan.

Seed 2Ð2.5 ϫ 1.2Ð1.5 ϫ 1Ð1.2 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; mottled and streaked; with frequent mottles; with frequent streaks; dark reddish brown; with brown Callistachys: C. lanceolata É.P. Ventenat (AÐE). A, Fruits (darker reddish) overlay; glabrous; smooth; coriaceous. with calyx (ϫ 2.9); B, seeds (ϫ 9.7); CÐD, testa (ϫ 50, Fracture lines absent. Rim absent. Wings absent. Raphe ϫ 1000); E, embryos (ϫ 20).

983 Genus: Gastrolobium R. Brown with frequent mottles; reddish brown or black; with purple overlay; glabrous; smooth; coriaceous. Fracture Phylogenetic Number: 24.13. lines absent. Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by aril; Tribe: Mirbelieae. with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than puncti- Species Studied—Species in Genus: 10 spp.—ca. 50 spp. form; 1 mm long; with curved outline; oval; marginal according to radicle tip; recessed; within rim. Hilum Fruit a legume; unilocular; 0.7Ð2 ϫ 0.3Ð0.7 ϫ 0.2Ð0.65 cm; rim color darker than testa. Lens discernible; equal to or with persistent calyx; with calyx shorter than fruit; greater than 0.5 mm in length; up to 1 mm long; with without orifice formed by curving of fruit or fruit margins straight or curved; circular or oblong; not in segments; straight; not plicate; not twisted; symmetrical groove of raphe; confluent with or adjacent to hilum; up or asymmetrical; ovate or circular; when asymmetrical to 0.2 mm from hilum; mounded or flush; similar color with 1 straight and 1 curved suture; widest near middle as testa; darker than testa; brown; not within corona, or D-shaped; not inflated; terete; without beak; rounded halo, or rim. Endosperm thin; covering entire embryo; at apex; apex aligned with longitudinal axis of fruit; adnate to testa. Cotyledons smooth; both outer faces short tapered at base; base aligned or oblique with convex; both the same thickness; both more or less of longitudinal axis of fruit; with the apex and base equal length; not folded; margin entire 180 degrees uniform in texture; coriaceous or ligneous; seed from base of radicle; similar at apex; not concealing chambers externally invisible. Fruit margin not con- radicle; split over radicle; with lobes; with the interface stricted; without sulcus; plain. Fruit wings absent. Fruit division terminating at base of radicle; without margins stipitate to substipitate; with the stipe up to 9 mm long. recessed; tan or brown (reddish); inner face flat; Fruit with all layers dehiscing; splitting along sutures. glabrous around base of radicle. Embryonic axis Dehiscence of valves along both sutures; apical and deflexed; oblique to length of seed; without a joint down; passive. Replum invisible. Epicarp dull; mono- evident between the radicle and the cotyledons. Radicle chrome or multicolored; mottled; reddish brown; with linear; deflexed and parallel to cotyledon length; purple overlay; glabrous, pubescent and indurate, or centered between cotyledons; less than 1/2 length of glabrate; with hairs erect or appressed; with 1 type of cotyledons. Plumule rudimentary; glabrous. pubescence; villous; with pubescence gray; with pubescence uniformly distributed; with simple hairs; Distribution: Australia. pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately Notes: Crisp and Weston (1987) included the Oxylobium veined; not tuberculate; not exfoliating; without cracks. parviflorum J. Lindley group in Gastrolobium. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth or scurfy; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seed 1; length parallel with fruit length. Funiculus measured; up to 2 mm long; thick; straight or curved (near apex). Aril present or absent; fleshy (to somewhat dry, but better developed than rim-aril); topknotlike or marginal hilar; covering 1/2 to nearly all of seed; reddish tan or cream.

Seed 1.7Ð5 ϫ 1.3Ð3 ϫ 1Ð2.3 mm; not overgrown; angular or not angular; asymmetrical or symmetrical; mitaform, oblong, reniform, or circular; compressed; with surface smooth; with or without visible radicle and cotyledon lobes; without external groove between radicle and Gastrolobium: G. bilobum R. Brown (CÐE), G. spp. (AÐB). cotyledon lobes; without hilar sinus; without umbo on A, Fruits (dehisced and closed) with and without calyx seed faces. Testa not adhering to endocarp; glossy; not (ϫ 2.9); B, seeds (ϫ 6); CÐD, testa (ϫ 50, ϫ 1000); E, modified by a bloom; colored; monochrome or mottled; embryos (ϫ 10).

985 Genus: Nemcia K. Domin brown overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not Phylogenetic Number: 24.15. visible. Hilum partially concealed; concealed by aril; with faboid split; with the lips of the faboid split the Tribe: Mirbelieae. same color as the rest of the hilum; larger than punctiform; 0.7 mm long; with straight outline; oblong; Species Studied—Species in Genus: 9 spp.—28 spp. marginal according to radicle tip; recessed; not within corona, halo, or rim. Lens discernible; less than 0.5 mm Fruit a legume; unilocular; 0.8Ð1.5 ϫ 0.3Ð0.5 ϫ 0.2Ð0.4 or equal to or greater than 0.5 mm in length; 0.5 mm cm; with persistent calyx; with calyx shorter than fruit; long; with margins straight or curved; oblong; not in without orifice formed by curving of fruit or fruit groove of raphe; confluent with hilum; barely segments; straight; not plicate; not twisted; asymmetri- mounded; dissimilar color from testa; darker than testa; cal or symmetrical; ovate or oblong; when asymmetri- black; not within corona, halo, or rim. Endosperm thin; cal with 1 straight and 1 curved suture, both sutures covering entire embryo; adnate to testa or embryo. unequally curved, or both sutures parallelly curved; Cotyledons smooth; both outer faces convex; both the widest near middle or D-shaped; not inflated; terete; same thickness; both more or less of equal length; not with or without beak; straight or declined; with solid folded; margin entire 180 degrees from base of radicle; beak the same color and texture as fruit; short tapered at similar at apex; not concealing radicle; entire over apex; apex aligned with longitudinal axis of fruit; radicle; without lobes; with the interface division tapered at base; base aligned with longitudinal axis of terminating at base of radicle; without margins re- fruit; with the apex and base uniform in texture; cessed; reddish tan; inner face flat; glabrous around coriaceous; seed chambers externally invisible. Fruit base of radicle. Embryonic axis deflexed; oblique to margin not constricted; without sulcus; plain. Fruit length of seed; without a joint evident between the wings absent. Fruit substipitate or nonstipitate. Fruit radicle and the cotyledons. Radicle linear or bulbose; with all layers dehiscing; splitting along sutures. deflexed and parallel to cotyledon length; centered Dehiscence of valves along both sutures; apical and between cotyledons; less than 1/2 length of cotyledons. down (at least 1/2 way); passive. Replum invisible. Plumule rudimentary; glabrous. Epicarp dull; monochrome; dark to light reddish brown; pubescent and indurate; with 1 type of pubescence; Distribution: Southwestern Australia. villous; with pubescence golden or gray; with pubescence uniformly distributed; with simple hairs; pliable; Notes: Crisp and Weston (1987) emended the generic with hair bases plain; eglandular; without spines; not circumscription to include series Axillares of smooth; with elevated features; reticulately veined; not Gastrolobium (24.13). tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; pale tan; smooth or reticulate; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seed 1; length parallel with fruit length. Funiculus measured; 1 mm long; thick; curved. Aril fleshy or dry; when fleshy topknotlike; covering less than 1/2 of seed; when dry rim-aril; reddish brown.

Seed 2.3Ð3 ϫ 1.5Ð1.8 ϫ 1Ð1.5 mm; not overgrown; not angular; asymmetrical; oblong, reniform, or D-shaped; compressed; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to Nemcia: N. dilatata (G. Bentham) M.D. Crisp (CÐE), N. endocarp; glossy; not modified by a bloom; colored; spp. (AÐB). A, Fruits (dehisced and closed) with and monochrome or mottled and streaked; with frequent without calyx (ϫ 4.8); B, seeds (ϫ 10.1); CÐD, testa mottles; with frequent streaks; reddish brown; with (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

987 Genus: Brachysema R. Brown absent. Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by aril; with Phylogenetic Number: 24.16. faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 1 Tribe: Mirbelieae. mm long; with curved outline; oval; between cotyledon and radicle lobe; recessed; not within corona, halo, or Species Studied—Species in Genus: 7 spp.—8 spp. rim or within halo. Hilum halo nearly color of testa. Lens discernible or not discernible; less than 0.5 mm to Fruit a legume; unilocular; 1Ð2.5 ϫ 0.5Ð1 ϫ 0.5Ð0.6 cm; equal to or greater than 0.5 mm in length; up to 0.7 mm with persistent calyx; with calyx shorter than fruit; long; with margins curved; key-hole shaped; not in without orifice formed by curving of fruit or fruit groove of raphe; confluent with hilum; flush; dissimilar segments; straight; not plicate; not twisted; symmetri- color from testa; darker than testa; dark reddish brown; cal; oblong; not inflated; terete; without beak; tapered not within corona, halo, or rim. Endosperm thick; to short tapered at apex; apex aligned with longitudinal covering entire embryo; adnate to embryo. Cotyledons axis of fruit; rounded at base; base aligned with smooth; both outer faces convex; both the same longitudinal axis of fruit; with the apex and base thickness; both more or less of equal length; not folded; uniform in texture; ligneous; seed chambers externally margin entire 180 degrees from base of radicle; similar invisible. Fruit margin not constricted; without sulcus; at apex; not concealing radicle; split over radicle; with plain. Fruit wings absent. Fruit substipitate to lobes; with the interface division terminating at base of nonstipitate. Fruit with all layers dehiscing; splitting radicle; without margins recessed; yellow or tan; inner along sutures. Dehiscence of valves along both sutures; face flat; glabrous around base of radicle. Embryonic apical and down; passive. Replum invisible. Epicarp axis deflexed; oblique to length of seed; without a joint dull; monochrome or multicolored; mottled (faintly in evident between the radicle and the cotyledons. Radicle some species); reddish brown or tan; with gray overlay; linear or bulbose; lobe tip straight; deflexed and parallel pubescent and indurate; with 1 type of pubescence; to cotyledon length; centered between cotyledons; less tomentose; with pubescence gray; with pubescence than 1/2 length of cotyledons. Plumule rudimentary or uniformly distributed (or restricted to sutures); with moderately developed; glabrous. simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; Distribution: Australia. reticulately veined; not tuberculate; shagreen or wrinkled; exfoliating in part; with or without cracks; Notes: Brachysema is “under revision by M.D. Crisp” cracking transverse to fruit length. Mesocarp thick; (Crisp and Weston 1987), but Crisp and Weston (1987) surface not veined; 1-layered; without balsamic did not recognize Cupulanthus J. Hutchinson (old vesicles; without fibers; solid; ligneous. Endocarp dull; phylogenetic number 25.03). Cupulanthus bracteolosus monochrome; reddish tan; transversely wrinkled; (F.H.J. von Mueller) J. Hutchinson now is B. nonseptate; ligneous; not exfoliating; remaining fused bracteolosus F.H.J. von Mueller. to mesocarp and epicarp; entire. Seeds 7; length parallel with fruit length; overlapping and touching; in 1 series. Funiculus measured; 1 mm long; of 1 length only; thick; straight or curved (slightly). Aril fleshy; cupshaped; covering less than 1/2 of seed; reddish brown or tan.

Seed 2Ð3.1 ϫ 1.5Ð2 ϫ 1Ð1.5 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not Brachysema: B. lanceolata C.D.F. Meissner (CÐE), B. spp. modified by a bloom; colored; monochrome; black or (AÐB). A, Fruits (dehisced and closed) with and without brown (reddish); glabrous; smooth or not smooth; with calyx (ϫ 2.1); B, seeds (ϫ 9.6); CÐD, testa (ϫ 50, recessed features; punctate; coriaceous. Fracture lines ϫ 1000); E, embryos (ϫ 13.3).

989 Genus: Latrobea C.D.F. Meissner a bloom; colored; monochrome; dark reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Phylogenetic Number: 24.18. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid Tribe: Mirbelieae. split the same color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; somewhat re- Species Studied—Species in Genus: 2 spp.—7 spp. cessed; within rim. Hilum rim color of testa. Lens discernible; less than 0.5 mm in length; with margins Fruit a legume; unilocular; 0.7Ð1 ϫ 0.2Ð0.55 ϫ 0.15Ð0.2 curved; circular; not in groove of raphe; adjacent to cm; with persistent calyx; with calyx shorter than fruit; hilum; 0.5 mm from hilum; mounded; same color as without orifice formed by curving of fruit or fruit testa; not within corona, halo, or rim. Endosperm thin; segments; straight; not plicate; not twisted; asymmetri- covering entire embryo; adnate to testa. Cotyledons cal; ovate or falcate; when asymmetrical with both smooth; both outer faces convex; both the same sutures parallelly curved; not inflated; compressed; thickness; both more or less of equal length; not folded; without or with beak; straight; with solid beak the same margin entire 180 degrees from base of radicle; similar color and texture as fruit; tapered or short tapered at at apex; not concealing radicle; entire over radicle; apex; apex aligned or oblique with longitudinal axis of without lobes; with the interface division terminating at fruit; tapered or rounded at base; base aligned with base of radicle; without margins recessed; reddish longitudinal axis of fruit; with the apex and base brown; inner face flat; glabrous around base of radicle. uniform in texture; coriaceous; seed chambers exter- Embryonic axis deflexed; oblique to length of seed; nally invisible. Fruit margin not constricted; without without a joint evident between the radicle and the sulcus; plain. Fruit wings absent. Fruit nonstipitate. cotyledons. Radicle linear; deflexed and parallel to Fruit with all layers dehiscing; splitting along sutures. cotyledon length; centered between cotyledons; less Dehiscence of valves along both sutures; apical and than 1/2 length of cotyledons. Plumule rudimentary; down; passive. Replum invisible. Epicarp dull; mono- glabrous. chrome or multicolored; mottled; reddish brown, tan (to greenish tan), purple, or green; with purple overlay; Distribution: Southwestern Australia. pubescent and indurate; with hairs erect or appressed; with 1 type of pubescence; villous; with pubescence golden or gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; vitreous (fractured); coriaceous. Endocarp dull; monochrome or mottled (somewhat because of darker mesocarp layer); grayish brown or gray; with brown (reddish) overlay; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 0.5 mm long; of 1 length only; filiform; straight. Aril fleshy; cupshaped; covering less than 1/2 of seed; tan.

Seed 2 ϫ 1.2 ϫ 0.8 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon Latrobea: L. tenella G. Bentham (CÐE), L. spp. (AÐB). A, lobes; without hilar sinus; without umbo on seed faces. Fruits with calyx (ϫ 2.8); B, seeds (ϫ 10); CÐD, testa Testa not adhering to endocarp; glossy; not modified by (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

991 Genus: Phyllota (A.-P. de Candolle) G. Bentham ceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with Phylogenetic Number: 24.19. the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.5 mm long; with Tribe: Mirbelieae. curved outline; elliptic; between cotyledon and radicle lobe; recessed; within halo. Hilum halo color darker Species Studied—Species in Genus: 4 spp.—10 spp. than testa. Lens discernible; equal to or greater than 0.5 mm in length; 0.8Ð1 mm long; with margins straight or Fruit a legume; unilocular; 0.45Ð0.5 ϫ 0.25Ð0.3 ϫ 0.15Ð0.2 curved; oblong or linear; not in groove of raphe; cm; with persistent calyx; with calyx shorter than fruit; confluent with hilum; mounded; dissimilar color from without orifice formed by curving of fruit or fruit testa; darker than testa; reddish brown; not within segments; straight; not plicate; not twisted; asymmetri- corona, halo, or rim. Endosperm thin; covering entire cal or symmetrical; obovate or ovate; when asymmetri- embryo; adnate to testa. Cotyledons smooth; both outer cal with both sutures parallelly curved; not inflated; faces convex; both the same thickness; both more or compressed; without beak; short tapered at apex; apex less of equal length; not folded; margin entire 180 aligned or oblique with longitudinal axis of fruit degrees from base of radicle; similar at apex; not (slightly); rounded at base; base aligned with longitudi- concealing radicle; split over radicle; with lobes; with nal axis of fruit; with the apex and base uniform in the interface division terminating at base of radicle; texture; ligneous; seed chambers externally invisible. without margins recessed; green; inner face flat; Fruit margin not constricted; without sulcus; plain. Fruit glabrous around base of radicle. Embryonic axis wings absent. Fruit nonstipitate. Fruit with all layers deflexed; oblique to length of seed; without a joint dehiscing; splitting along sutures. Dehiscence of valves evident between the radicle and the cotyledons. Radicle along both sutures; apical and down; passive or active; bulbose; deflexed and parallel to cotyledon length; with valves enrolling. Replum invisible. Epicarp dull; centered between cotyledons; less than 1/2 length of monochrome; reddish brown or tan; pubescent and cotyledons. Plumule rudimentary; glabrous. indurate; with hairs appressed or erect; with 1 type of pubescence; tomentose; with pubescence golden or Distribution: Australia. gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; not veined or veined; reticulately veined; not tuberculate; shagreen; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy; monochrome; reddish brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened; straight. Aril absent.

Seed 2.3Ð4 ϫ 1.5Ð2 ϫ 1.2Ð1.5 mm; not overgrown; not angular or angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Cuticle inflated; inflated around hilum. Testa not adhering to endocarp; dull; not modified by a bloom; colored; mottled and streaked; with frequent mottles; with frequent streaks; reddish Phyllota: P. pleurandroides F.H.J. von Mueller (AÐE). A, brown; with black overlay; glabrous; smooth; coria- Fruits with calyx (ϫ 8.7); B, seeds (ϫ 9.5); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 20).

993 Genus: Euchilopsis F.H.J. von Mueller Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; Phylogenetic Number: 24.20. larger than punctiform; 0.5 mm long; with straight outline; oblong; between cotyledon and radicle lobe; Tribe: Mirbelieae. flush; within rim, corona, and halo. Hilum corona color (brown) darker than testa. Hilum halo color (reddish- Species Studied—Species in Genus: 1 sp.—1 sp. brown) lighter than testa. Hilum rim color of testa. Lens discernible; equal to or greater than 0.5 mm in length; 1 Fruit a legume; unilocular; 0.9Ð1 ϫ 0.5Ð0.6 ϫ 0.35Ð0.45 mm long; with margins straight; linear; not in groove of cm; with persistent calyx; with calyx shorter than fruit; raphe; confluent with hilum; flush; dissimilar color without orifice formed by curving of fruit or fruit from testa; darker than testa; darker reddish brown; not segments; straight; not plicate; not twisted; asymmetri- within corona, halo, or rim. Endosperm thin; restricted cal or symmetrical; ovate; when asymmetrical with to region of embryo; adnate to testa. Cotyledons both sutures parallelly curved; not inflated; terete; smooth; both outer faces convex; both the same without beak; rounded at apex; apex aligned or oblique thickness; both more or less of equal length; not folded; (slightly) with longitudinal axis of fruit; short tapered or margin entire 180 degrees from base of radicle; similar rounded at base; base aligned with longitudinal axis of at apex; not concealing radicle; split over radicle; with fruit; with the apex and base uniform in texture; lobes; with the interface division terminating at base of ligneous; seed chambers externally invisible. Fruit radicle; without margins recessed; green; inner face margin not constricted; without sulcus; plain. Fruit flat; glabrous around base of radicle. Embryonic axis wings absent. Fruit substipitate; with the stipe 2.5Ð3 deflexed; oblique to length of seed; without a joint mm long. Fruit with all layers dehiscing; splitting along evident between the radicle and the cotyledons. Radicle sutures. Dehiscence of valves along both sutures; apical linear; deflexed and parallel to cotyledon length; and down; passive. Replum invisible. Epicarp dull; centered between cotyledons; less than 1/2 length of monochrome; reddish brown; pubescent but soon cotyledons. Plumule rudimentary; glabrous. deciduous; with 1 type of pubescence; villous; with pubescence pale golden; with pubescence uniformly Distribution: Southwestern Australia. distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with Notes: Unfortunately, we were able to study only three fruit recessed features; not veined; not tuberculate; pitted; samples, each with minimal material, and only one not exfoliating; without cracks. Mesocarp thin; surface seed. not veined; 2-layered; without balsamic vesicles; without fibers; with vitreous layer over solid layer (vitreous layer beadlike); ligneous (or subligneous). Endocarp glossy; monochrome; reddish brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching or overlapping and touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick or triangular; straight. Aril absent.

Seed 4 ϫ 2.3 ϫ 1 mm; not overgrown; not angular; asymmetrical; D-shaped or rectangular; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; coriaceous. Fracture lines Euchilopsis: E. linearis (G. Bentham) F.H.J. von Mueller absent. Rim absent. Wings absent. Raphe not visible. (AÐE). A, Fruits and valve (ϫ 5.7); B, seed (ϫ 10); CÐD, testa (ϫ 50, ϫ 1000).

995 Genus: Aotus J.E. Smith endocarp; dull; not modified by a bloom; colored; monochrome (cuticle may be raised, causing silver Phylogenetic Number: 24.21. patches); dark to light reddish brown or black; glabrous; not smooth; with elevated features; reticulate or Tribe: Mirbelieae. reticulate and shagreen; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Species Studied—Species in Genus: 8 spp.—16–18 spp. Hilum visible or partially concealed; concealed by funicular remnant, radicle lobe, or wing; with faboid Fruit a legume; unilocular; 0.5Ð0.6 ϫ 0.4Ð0.45 ϫ 0.3Ð0.45 split; with the lips of the faboid split the same color as cm; with persistent calyx; with calyx shorter than fruit; the rest of the hilum; punctiform to larger than puncti- without orifice formed by curving of fruit or fruit form; up to 0.5 mm long; with curved outline; elliptic; segments; straight; not plicate; not twisted; symmetrical marginal according to radicle tip or between cotyledon or asymmetrical; more or less circular; when asym- and radicle lobe; recessed; not within corona, halo, or metrical with both sutures parallelly curved; not rim or within halo. Hilum halo color lighter than testa. inflated; terete; without beak; rounded at apex; apex Lens discernible or not discernible; less than 0.5 mm oblique or aligned with longitudinal axis of fruit; short long or equal to or greater than 0.5 mm; up to 0.5 mm tapered at base; base aligned with longitudinal axis of long; with margins straight or curved; triangular or fruit; with the apex and base uniform in texture; more or less circular; not in groove of raphe; confluent ligneous; seed chambers externally invisible. Fruit with hilum; flush; dissimilar color from testa; darker margin not constricted; without sulcus; embellished. than testa; black (ish); not within corona, halo, or rim. Fruit wings absent. Fruit stipitate or substipitate; with Endosperm thick (reflecting testa reticulation); covering the stipe up to 15 mm long. Fruit with all layers entire embryo; adnate to testa or embryo. Cotyledons dehiscing; splitting along sutures. Dehiscence of valves smooth; both outer faces convex; both the same along both sutures; apical and down; passive. Replum thickness; both more or less of equal length; not folded; invisible. Epicarp dull; monochrome; dark reddish margin entire 180 degrees from base of radicle; similar brown; pubescent and indurate; with 1 type of pubes- at apex; not concealing radicle; split over radicle; with cence; villous; with pubescence golden or gray; with lobes; with the interface division terminating at base of pubescence uniformly distributed; with simple hairs; radicle; without margins recessed; green or tan; inner pliable; with hair bases plain; glandular; with glandular face flat; glabrous around base of radicle. Embryonic dots; without spines; not smooth; with elevated fea- axis deflexed; oblique to length of seed; without a joint tures; not veined; not tuberculate; shagreen and evident between the radicle and the cotyledons. Radicle subvesicular; not exfoliating; without cracks. Mesocarp bulbose; lobe tip straight or hooked; deflexed and thick; surface not veined; 1-layered; without balsamic parallel to cotyledon length; centered between cotyle- vesicles; without fibers; solid; ligneous. Endocarp dons; less than 1/2 length of cotyledons. Plumule glossy; monochrome or mottled; dark reddish brown; rudimentary; glabrous. with mottling (dark); with brown (reddish) overlay; smooth; nonseptate; chartaceous; not exfoliating; Distribution: Australia. remaining fused to mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlap- Notes: The thick cuticle on the testa exfoliates during ping nor touching or overlapping and touching; in 1 soaking of the seed. series. Funiculus measured or less than 0.5 mm long; up to 0.5 mm long; of 1 length only; thick; straight. Aril present or absent; dry; rim-aril; white.

Seed 2.2Ð3 ϫ 1.7Ð3 ϫ 1Ð2 mm; not overgrown; not angular or angular (somewhat); asymmetrical; D-shaped or oblong (more or less); compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes; Aotus: A. ericoides (É.P. Ventenat) G. Don (CÐE), A. spp. with external groove between radicle and cotyledon (AÐB). A, Fruits (closed and dehisced) with calyx lobes the same color as testa; without hilar sinus; (ϫ 3.5); B, seeds (ϫ 9.8); CÐD, testa (ϫ 50, ϫ 1000); without umbo on seed faces. Testa not adhering to E, embryos (ϫ 10).

997 Genus: Urodon P.K.N.S. Turczaninow rest of the hilum; larger than punctiform; 0.5 mm long; with curved outline; elliptic; subapical to radicle tip; Phylogenetic Number: 24.22. recessed; not within corona, halo, or rim or within rim (slight). Hilum rim color of testa. Lens discernible; less Tribe: Mirbelieae. than 0.5 mm in length; with margins curved; circular; not in groove of raphe; adjacent to hilum; 0.3 mm from Species Studied—Species in Genus: 1 sp.—1–3 spp. hilum; mounded; dissimilar color from testa; darker than testa; black; not within corona, halo, or rim. Fruit a legume; unilocular; 0.6Ð0.7 ϫ 0.3Ð0.35 ϫ 0.14Ð0.16 Endosperm thin; covering entire embryo; adnate to cm; with persistent calyx; with calyx shorter than fruit; testa. Cotyledons smooth; both outer faces convex; both without orifice formed by curving of fruit or fruit the same thickness; both more or less of equal length; segments; straight; not plicate; not twisted; asymmetri- not folded; margin entire 180 degrees from base of cal; when asymmetrical with 1 straight and 1 curved radicle; similar at apex; not concealing radicle; split suture; widest near middle or D-shaped; not inflated; over radicle; with lobes; with the interface division compressed; without beak; short tapered at apex; apex terminating at base of radicle; without margins realigned with longitudinal axis of fruit; short tapered at cessed; tan; inner face flat; glabrous around base of base; base oblique with longitudinal axis of fruit; with radicle. Embryonic axis deflexed; oblique to length of the apex and base uniform in texture; membranous; seed; without a joint evident between the radicle and seed chambers externally invisible. Fruit margin not the cotyledons. Radicle linear; deflexed and parallel to constricted; without sulcus; plain. Fruit wings absent. cotyledon length; centered between cotyledons; 1/2 to Fruit nonstipitate. Fruit with all layers dehiscing; nearly length of cotyledons. Plumule rudimentary; splitting along suture. Dehiscence of valves along 1 glabrous. suture; apical and down; active; with valves revolute. Replum invisible. Epicarp dull; monochrome; dark Distribution: Southwestern Australia. reddish brown; pubescent and indurate; with hairs appressed; with 1 type of pubescence; with pubescence Notes: Crisp and Weston (1987) resurrected Urodon. This gray; with pubescence uniformly distributed; with genus has three described species, but they were simple hairs; stiff; with hair bases plain; eglandular; uncertain whether these species were truly separate or without spines; with elevated features; faintly, trans- were instead one variable species. versely veined relative to fruit length; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp dull; monochrome; grayish brown or gray; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching or overlapping and touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened; straight. Aril fleshy; topknotlike; fimbriate; covering less than 1/2 of seed; reddish brown.

Seed 2.7 ϫ 1.6 ϫ 0.8 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; monochrome; dark reddish brown; glabrous; smooth or not smooth (occasionally); with recessed features; occasionally pitted with small separate pits; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Urodon: U. capitatus P.K.N.S. Turczaninow (AÐE). A, partially concealed; concealed by aril; with faboid split; Fruits with calyx (closed and dehisced) (ϫ 6); B, seeds with the lips of the faboid split the same color as the (ϫ 13); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 20).

999 Genus: Pultenaea J.E. Smith endocarp; glossy or dull; not modified by a bloom; colored; mottled and streaked or monochrome; with Phylogenetic Number: 24.23. frequent mottles; with frequent streaks; reddish brown; with black overlay; glabrous; smooth; coriaceous. Tribe: Mirbelieae. Fracture lines absent. Rim present. Wings absent. Raphe not visible. Hilum partially concealed; concealed Species Studied—Species in Genus: 14 spp.—ca. 150 spp. by aril; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than Fruit a legume; unilocular; 0.4Ð0.7 ϫ 0.3Ð0.35 ϫ 0.2 cm; punctiform; 0.5Ð0.6 mm long; with curved outline; with persistent calyx; with calyx shorter or longer than elliptic; between cotyledon and radicle lobe; recessed; fruit; without orifice formed by curving of fruit or fruit within rim. Hilum rim color lighter than testa. Lens not segments; straight; not plicate; not twisted; asymmetri- discernible or discernible; less than 0.5 mm or equal to cal or symmetrical; ovate; when asymmetrical with 1 or greater than 0.5 mm in length; 0.3Ð0.5 mm long; straight and 1 curved suture or both sutures parallelly with margins curved; circular; not in groove of raphe; curved; widest near middle or D-shaped; not inflated; confluent with hilum; flush; dissimilar color from testa; compressed or terete; without or with beak; straight or darker than testa; reddish brown; not within corona, hooked; with solid beak the same color and texture as halo, or rim. Endosperm thin; covering entire embryo; fruit; short tapered or rounded at apex; apex aligned or adnate to testa. Cotyledons smooth; both outer faces oblique with longitudinal axis of fruit; rounded or convex; both the same thickness; both more or less of truncate at base; base aligned with longitudinal axis of equal length; not folded; margin entire 180 degrees fruit; with the apex and base uniform in texture; from base of radicle; similar at apex; not concealing coriaceous; seed chambers externally invisible. Fruit radicle; split over radicle; with lobes; with the interface margin not constricted; without sulcus; plain. Fruit division terminating at base of radicle; without margins wings absent. Fruit substipitate or nonstipitate. Fruit recessed; reddish brown, tan, or yellow; inner face flat; with all layers dehiscing; splitting along suture. glabrous around base of radicle. Embryonic axis Dehiscence of valves along 1 suture; apical and down; deflexed; oblique to length of seed; without a joint passive. Replum invisible. Epicarp dull; monochrome; evident between the radicle and the cotyledons. Radicle reddish brown or black; glabrous or pubescent but soon linear or bulbose; deflexed and parallel to cotyledon deciduous; with hairs appressed or erect; with 1 type of length; centered between cotyledons; less than 1/2 pubescence; villous; with pubescence golden or gray; length of cotyledons. Plumule rudimentary; glabrous. with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without Distribution: Australia. spines; not smooth; with elevated features; reticulately veined; not tuberculate; minutely warty; exfoliating in part or exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; reddish brown or black; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with fruit length; overlapping or overlapping and touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened; straight. Aril fleshy; topknotlike or cupshaped; entire or fimbriate; covering less than 1/2 of seed; tan to reddish tan.

Seed 1.5Ð4 ϫ 0.6Ð2 ϫ 0.7Ð1.3 mm; not overgrown; not angular; asymmetrical; oblong, mitaform, or reniform; compressed; with surface smooth; with visible radicle Pultenaea: P. obcordata (R. Brown) G. Bentham (CÐE), P. and cotyledon lobes; without external groove between spp. (AÐB). A, Fruits within calyx (closed and dehisced) radicle and cotyledon lobes; without hilar sinus; (ϫ 3.3); B, seeds (ϫ 6.7); CÐD, testa (ϫ 50, ϫ 1000); without umbo on seed faces. Testa not adhering to E, embryos (ϫ 10).

1001 Genus: Almaleea M.D. Crisp & P.H. Weston colored; mottled or monochrome; with frequent mottles; reddish brown; with brown (reddish) overlay; Phylogenetic Number: 24.24. glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Tribe: Mirbelieae. partially concealed; concealed by aril; with faboid split; with the lips of the faboid split the same color as the Species Studied—Species in Genus: 5 spp.—5 spp. rest of the hilum; larger than punctiform; 0.5 mm long; with straight outline; oblong; between cotyledon and Fruit a legume; unilocular; 0.3Ð0.6 ϫ 0.2Ð0.4 ϫ 0.2Ð0.3 radicle lobe; recessed; not within corona, halo, or rim or cm; with persistent calyx; with calyx shorter than or within halo. Hilum halo color lighter than testa. Lens equal in length to fruit; without orifice formed by discernible; less than 0.5 mm or equal to or greater than curving of fruit or fruit segments; straight; not plicate; 0.5 mm in length; 0.5 mm long; with margins curved; not twisted; asymmetrical; ovate; when asymmetrical more or less circular; not in groove of raphe; confluent with both sutures parallelly curved; not inflated; terete; with hilum; flush; similar color as testa; darker than without beak; rounded or short tapered at apex; apex testa; reddish brown; not within corona, halo, or rim. oblique or aligned with longitudinal axis of fruit; short Endosperm thin; covering entire embryo; adnate to tapered or rounded at base; base aligned with longitudi- testa. Cotyledons smooth; both outer faces convex; both nal axis of fruit; with the apex and base uniform in the same thickness; both more or less of equal length; texture; coriaceous; seed chambers externally invisible. not folded; margin entire 180 degrees from base of Fruit margin not constricted; without sulcus; plain. Fruit radicle; similar at apex; not concealing radicle; split wings absent. Fruit nonstipitate. Fruit with all layers over radicle; with lobes; with the interface division dehiscing; splitting along sutures. Dehiscence of valves terminating at base of radicle; without margins re- along both sutures or 1 suture; apical and down (about cessed; yellow or tan; inner face flat; glabrous around 1/2 way down); passive. Replum invisible. Epicarp base of radicle. Embryonic axis deflexed; oblique to dull; monochrome; dark reddish brown; pubescent and length of seed; without a joint evident between the indurate; with 1 type of pubescence; villous; with radicle and the cotyledons. Radicle bulbose; lobe tip pubescence gray; with pubescence uniformly distrib- straight; deflexed and parallel to cotyledon length; uted; with simple hairs; pliable; with hair bases plain or centered between cotyledons; less than 1/2 length of swollen; eglandular; without spines; not smooth; with cotyledons. Plumule moderately developed; glabrous. elevated or recessed features; reticulately veined; not tuberculate; tuberculate (base of hairs); punctate (when Distribution: Southeastern Australia. hair bases absent); not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without Notes: Almaleea includes species formerly placed in balsamic vesicles; without fibers; solid; coriaceous. Pultenaea (24.23) and Dillwynia (24.25) (Crisp and Endocarp glossy; monochrome; grayish or reddish tan Weston 1991). Crisp and Weston noted that Almaleea or gray; smooth; nonseptate; chartaceous; exfoliating in was closely related to Eutaxia (24.26) and Dillwynia part; remaining fused to mesocarp and epicarp; entire. (24.25). Seeds 1 or 2; length parallel with fruit length; overlapping, touching, or neither overlapping nor touching; in 1 or 2 or more series. Funiculus measured; 0.3Ð0.8 mm long; of 1 length only; thick; straight. Aril fleshy; cupshaped; covering less than 1/2 of seed; tan.

Seed 1Ð2.7 ϫ 1Ð2 ϫ 0.6Ð1.2 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with surface smooth; with visible radicle and cotyledon lobes; with or without external groove between radicle and cotyledon lobes; with external groove between Almaleea: A. subumbellata (W.D. Hooker) M.D. Crisp & radicle and cotyledon lobes same color as testa; without P.E. Weston (CÐE), A. spp. (AÐB). A, Fruits with calyx hilar sinus; without umbo on seed faces. Testa not (ϫ 7.1); B, seeds (ϫ 9.5); CÐD, testa (ϫ 50, ϫ 1000); adhering to endocarp; dull; not modified by a bloom; E, embryos (ϫ 15).

1003 Genus: Dillwynia J.E. Smith colored; streaked; with frequent streaks; dark brown or black; with brown (lighter) or purple overlay; glabrous; Phylogenetic Number: 24.25. smooth or not smooth; with recessed features; punctate; coriaceous. Fracture lines absent. Rim absent. Wings Tribe: Mirbelieae. absent. Raphe not visible. Hilum fully concealed; concealed by aril; with faboid split; with the lips of the Species Studied—Species in Genus: 5 spp.—ca. 20 spp. faboid split the same color as the rest of the hilum; larger than punctiform; 0.5Ð0.6 mm long; with curved Fruit a legume; unilocular; 0.4Ð0.7 ϫ 0.25Ð0.45 ϫ 0.3Ð0.4 outline; circular; marginal according to radicle tip or cm; with persistent or deciduous calyx; with calyx between cotyledon and radicle lobe; recessed; within slightly shorter than fruit; without orifice formed by halo. Hilum halo color lighter than testa. Lens discern- curving of fruit or fruit segments; straight; not plicate; ible; equal to or greater than 0.5 mm in length; 0.6 mm not twisted; asymmetrical or symmetrical; circular, long; with margins straight or curved; oblong; not in elliptic, or ovate; when asymmetrical with both sutures groove of raphe; confluent with hilum (at least halo); parallelly curved; not inflated; terete; without or with barely mounded; similar color as testa; darker than beak; declined; with solid beak the same color and testa; reddish brown; not within corona, halo, or rim. texture as fruit; rounded at apex; apex oblique with Endosperm thick; covering entire embryo; adnate to longitudinal axis of fruit; short tapered at base; base testa or embryo. Cotyledons smooth; both outer faces aligned with longitudinal axis of fruit; with the apex convex; both the same thickness; both more or less of and base uniform in texture; ligneous; seed chambers equal length; not folded; margin entire 180 degrees externally invisible. Fruit margin not constricted; from base of radicle; similar at apex; not concealing without sulcus; plain. Fruit wings absent. Fruit radicle; split over radicle; with lobes; with the interface substipitate. Fruit with all layers dehiscing; splitting division terminating at base of radicle; without margins along sutures. Dehiscence of valves along both sutures; recessed; yellow or brown (reddish); inner face flat; apical and down; passive. Replum invisible. Epicarp glabrous around base of radicle. Embryonic axis dull; monochrome; reddish brown; pubescent but soon deflexed; oblique to length of seed; without a joint deciduous; with hairs erect or appressed; with 1 type of evident between the radicle and the cotyledons. Radicle pubescence; villous; with pubescence gray; with linear; deflexed and parallel to cotyledon length; pubescence uniformly distributed; with simple hairs; centered between cotyledons; less than 1/2 to 1/2 to pliable; with hair bases plain; eglandular; without nearly length of cotyledons. Plumule rudimentary; spines; not smooth; with elevated features; not veined glabrous. or veined (rarely); reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface Distribution: Australia. not veined; 1-layered; without balsamic vesicles; without fibers; solid; subligneous. Endocarp glossy; Notes: Two species of Dillwynia, known as parrot-pea, are monochrome; reddish brown or purple; smooth; Australian wildflowers and were discussed by Lebler nonseptate; chartaceous; not exfoliating; remaining (1976). fused to mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching or overlapping and touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril fleshy; topknotlike; covering less than 1/2 of seed; reddish tan or yellow.

Seed 2.5Ð3.5 ϫ 2Ð2.5 ϫ 1.3Ð1.7 mm; not overgrown; not angular or angular; asymmetrical; reniform, mitaform, or triangular; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external Dillwynia: D. floribunda J. Smith (CÐE), D. spp. (AÐB). groove between radicle and cotyledon lobes; without A, Fruits with or without calyx (closed and dehisced) hilar sinus; without umbo on seed faces. Testa not (ϫ 4.7); B, seeds (ϫ 9.4); CÐD, testa (ϫ 50, ϫ 1000); adhering to endocarp; dull; not modified by a bloom; E, embryos (ϫ 10).

1005 Genus: Eutaxia R. Brown reddish brown; with brown (dark reddish) or tan overlay; glabrous; smooth or not smooth; with elevated Phylogenetic Number: 24.26. features; reticulate; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum Tribe: Mirbelieae. partially concealed; concealed by aril and radicle lobe or wing; with faboid split; with the lips of the faboid Species Studied—Species in Genus: 4 spp.—10 spp. split the same color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; flush; within rim. Fruit a legume; unilocular; 0.4Ð0.6 ϫ 0.25Ð0.3 ϫ 0.1Ð0.25 Hilum rim color lighter (reddish) than testa. Lens cm; with persistent or deciduous calyx; with calyx discernible; less than 0.5 mm in length; with margins shorter than fruit; without orifice formed by curving of straight or curved; somewhat wedge-shaped; elliptic; fruit or fruit segments; straight; not plicate; not twisted; not in groove of raphe; confluent with hilum; mounded; symmetrical or asymmetrical; ovate; when asymmetri- similar color as testa; darker than testa; brown; not cal with 1 straight and 1 curved suture; widest near within corona, halo, or rim. Endosperm thin; covering middle or D-shaped; not inflated; compressed or entire embryo; adnate to embryo. Cotyledons smooth; flattened; without or with beak; hooked; with solid both outer faces convex; both the same thickness; both beak the same color and texture as fruit; rounded at more or less of equal length; not folded; margin entire apex; apex aligned or oblique with longitudinal axis of 180 degrees from base of radicle; similar at apex; not fruit; rounded at base; base aligned with longitudinal concealing radicle; split over radicle; with lobes; with axis of fruit; with the apex and base uniform in texture; the interface division terminating at base of radicle; coriaceous; seed chambers externally invisible. Fruit without margins recessed; yellow; inner face flat; margin not constricted; without sulcus; plain. Fruit glabrous around base of radicle. Embryonic axis wings absent. Fruit substipitate. Fruit with all layers deflexed; oblique to length of seed; without a joint dehiscing; splitting along sutures. Dehiscence of valves evident between the radicle and the cotyledons. Radicle along both sutures; apical and down; passive. Replum bulbose; lobe tip straight; deflexed and parallel to invisible. Epicarp dull; monochrome; reddish to dark cotyledon length; centered between cotyledons; 1/2 to reddish brown; pubescent and indurate; with 1 type of nearly length of cotyledons. Plumule rudimentary; pubescence; puberulent; with pubescence gray; with glabrous. pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without Distribution: Southern Australia. spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; reddish tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1 or 2; length parallel with fruit length; neither overlapping nor touching or overlapping and touching; in 1 series. Funiculus measured; 0.5Ð1 mm long; of 1 length only; filiform or thick; straight. Aril fleshy; topknotlike; covering less than 1/2 of seed; yellow.

Seed 2.2Ð2.7 ϫ 1.5Ð2 ϫ 1Ð1.5 mm; not overgrown; not angular or angular (somewhat); asymmetrical; mitaform; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not Eutaxia: E. microphylla (R. Brown) J.M. Black (CÐE), E. adhering to endocarp; dull; not modified by a bloom; spp. (AÐB). A, Fruits with or without calyx (closed and colored; monochrome or mottled and streaked; with dehisced) (ϫ 6.5); B, seeds (ϫ 9.2); CÐD, testa (ϫ 50, frequent mottles; with frequent streaks; brown to ϫ 1000); E, embryos (ϫ 10).

1007 Podalyrieae (25.01Ð25.09) (with a narrow opening on the micropylar side) or horseshoe shaped (with the opening towards the Genus: Xiphotheca C.F. Ecklon & J.M. Zeyher micropyle); entire; covering less than 1/2 of seed; orangish tan or white. Phylogenetic Number: 25.01. Seed 3.5Ð4.5 2.5Ð3.5 ca. 2 mm; not overgrown; not Tribe: Podalyrieae. angular; asymmetrical; reniform (oblong) or oblong; compressed; with surface smooth; without visible Subtribe: Xiphothecinae. radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces; without medial ridge on Species Studied—Species in Genus: 1 sp.—9 spp. each face. Testa without pieces of adhering epicarp; not adhering to endocarp; free from endocarp; glossy; not Fruit a legume; unilocular; 1.5Ð3.8 0.6Ð0.9 0.3Ð0.4 modified by a bloom; colored; mottled or monochrome cm; with persistent calyx or deciduous calyx; with (rarely); with frequent mottles; greenish tan to tan or calyx shorter than fruit; without orifice formed by brown; with brown (darker) overlay; glabrous; smooth; curving of fruit or fruit segments; straight or curved coriaceous. Fracture lines absent. Rim absent. Wings (rarely); not plicate; not twisted; asymmetrical or absent. Raphe from hilum through lens and terminating symmetrical (rarely); obliquely obovate, obliquely before base of seed; not bifurcating; color of or darker ovate, or oblong (rarely); when asymmetrical with both than testa; brown; flush. Hilum partially concealed; sutures unequally or parallelly (rarely) curved; not concealed by aril; with faboid split; with the lips of the inflated or inflated (rarely); compressed; with beak (1Ð5 faboid split the same color as the rest of the hilum; mm long); straight; with solid beak the same color and larger than punctiform; 0.5Ð1 mm long; with curved texture as fruit; tapered to long tapered at apex; apex outline; elliptic; apical according to radicle tip but aligned or oblique (slightly) with longitudinal axis of marginal according to seed length; recessed; within rim fruit; tapered or long tapered at base; base oblique or or not within corona, halo, or rim. Hilum rim color aligned with longitudinal axis of fruit; with the apex darker than testa. Lens discernible; equal to or greater and base uniform in texture; chartaceous; seed cham- than 0.5 mm in length; 1.2Ð1.5 mm long; with margins bers externally invisible or visible; with the raised seed straight; oblong; not in groove of raphe; confluent with chambers not torulose. Fruit margin not constricted or hilum; mounded; similar color as or dissimilar color constricted; slightly constricted along both margins; from testa; darker than testa; brown; not within corona, without sulcus; plain. Fruit wings absent. Fruit halo, or rim. Endosperm thick; not pluglike and not nonstipitate. Fruit with all layers dehiscing; splitting resembling tip of radicle; covering entire embryo; along sutures. Dehiscence of valves along both sutures adnate to embryo. Cotyledons smooth; both outer faces (assumed); active (assumed); with valves twisting convex; both the same thickness; both more or less of (slightly). Replum invisible. Epicarp dull; mono- equal length; not folded; margin entire 180 degrees chrome; brown to tan; with surface texture uniform; from base of radicle; similar at apex; not concealing pubescent and indurate or glabrous (rarely); with hairs radicle; entire over radicle; without lobes; with the erect; with 1 type of pubescence; tomentose, villous interface division terminating at base of radicle; without (densely), velutinous, or hirsute; with pubescence margins recessed; tan; inner face flat; glabrous around brown; with pubescence uniformly distributed; with base of radicle. Embryonic axis oblique; perpendicular simple hairs; pliable; with hair bases plain; straight; to length of seed; without a joint evident between the straight at apex; eglandular; without spines; smooth; not radicle and the cotyledons. Radicle bulbose; lobe tip veined; not tuberculate; not exfoliating; without cracks. straight; deflexed and parallel to cotyledon width; Mesocarp trace; surface not veined; 1-layered; without centered between cotyledons; less than 1/2 length of balsamic vesicles; without fibers; without reniform cotyledons. Plumule rudimentary; glabrous. canals; solid. Endocarp glossy; opaque; monochrome; tan; smooth; without adhering pieces of testa; Distribution: South Africa (Cape Province). nonseptate; taceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð6; length parallel Notes: Wyk and Schutte (1995) considered Liparieae and with fruit length; neither overlapping nor touching; in 1 Podalyrieae to each be monophyletic and Sophoreae (2) series. Funiculus measured; 0.4Ð0.8 mm long; of 1 to be their sister group. Using Crotalarieae (27) as the length only; flattened; straight. Aril fleshy; cupshaped outgroup, Schutte and Wyk (1998a,b) found that the

1008 genera of Liparieae and Podalyrieae coalesced into two closely related clades. Liparia (25.04) was in the Podalyria (25.06) clade. This supported earlier sugges- tions (Polhill 1976, 1981o; Wyk and Schutte 1995) that Liparieae and Podalyrieae should be merged. Schutte and Wyk (1998a,b) merged the two tribes as Podalyrieae, recognized the two clades as subtribes, Xiphothecinae and Podalyriinae, and erected a monotypic tribe for Hypocalyptus (26.01), Hypocalypteae (26). The generic enumeration and number of species in each genus follows Schutte and Wyk (1998a). Schutte and Wyk (1993) recently reinstated Xiphotheca, a segregate genus of Priestleya A.-P. de Candolle, and Schutte (1997) revised the genus.

1009 Xiphotheca: X. fruticosa (C. Linnaeus) A.L. Schutte & B.-E. van Wyk (AÐE). A, Fruits ( 4.2); B, seeds ( 3.3); CÐ D, testa ( 50, 1000); E, embryos ( 5).

1011 Genus: Amphithalea C.F. Ecklon & C.L.P. Zeyher radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to Phylogenetic Number: 25.02 endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth; Tribe: Podalyrieae. coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible (perhaps concealed by aril). Subtribe: Xiphothecinae. Hilum visible; with faboid split; with the lips of the faboid split the same color (nearly so) as the rest of the Species Studied—Species in Genus: 4 spp.—21 spp. hilum; larger than punctiform; 0.5 mm long; with curved outline; circular; between cotyledon and radicle Fruit a legume; unilocular; 0.7Ð0.8 0.3Ð0.4 0.2 cm; lobe; recessed; within rim or halo. Hilum halo color with persistent calyx; with calyx shorter than fruit; lighter than testa. Hilum rim color darker than testa. without orifice formed by curving of fruit or fruit Lens not discernible (perhaps concealed by aril). segments; straight; not plicate; not twisted; asymmetri- Endosperm absent. Cotyledons smooth; both outer cal; oblong to ovate; when asymmetrical with 1 straight faces convex; both the same thickness; both more or and 1 curved suture, both sutures unequally curved, or less of equal length; not folded; entire 180 degress from both sutures parallelly curved; widest near middle or D- base of radicle; similar at apex; not concealing radicle; shaped; not inflated; compressed; without beak; short entire over radicle; without lobes; with the interface tapered at apex; apex aligned with longitudinal axis of division terminating at base of radicle; without margins fruit; short tapered or rounded at base; base aligned recessed; tan; inner face flat; glabrous around base of with longitudinal axis of fruit; with the apex and base radicle. Embryonic axis deflexed; oblique to length of uniform in texture; coriaceous; seed chambers exter- seed; without a joint evident between the radicle and nally invisible. Fruit margin not constricted; without the cotyledons. Radicle linear; lobe tip straight; sulcus; plain. Fruit wings absent. Fruit substipitate or deflexed and parallel to cotyledon width; centered nonstipitate. Fruit with all layers dehiscing; splitting between cotyledons; 1/2 to nearly length of cotyledons. along sutures. Dehiscence of valves along both sutures; Plumule rudimentary; glabrous. apical and down; passive or active (barely); with valves enrolling (somewhat). Replum invisible. Epicarp dull; Distribution: South Africa (southern Cape Province). monochrome; greenish to dirty or reddish brown (concealed by long straight dense hairs) or gray Notes: Polhill (1981p) noted that Amphithalea may be (because of hairs); pubescent and indurate; with 1 type combined with Coelidium (26.04). Schutte (1998) of pubescence; villous (A. ericifolia (C. Linnaeus) C.F. carried out a cladistic analysis using morphological and Ecklon & J.M. Zeyher); with pubescence gray or alkaloid characters of Amphithalea and Coelidium with golden; with pubescence uniformly distributed; with Liparia (26.01) and Xiphotheca (26.03) as outgroups. simple hairs; pliable; with hair bases plain; eglandular; She concluded that the two genera are synonymous, and without spines; smooth; not veined; not tuberculate; not transferred the Coelidium species to Amphithalea. exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; spongy; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð4 (several); length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick or triangular; straight. Aril fleshy; cupshaped (with tonguelike extension); covering less than 1/2 of seed; reddish brown.

Seed 2.7Ð3 1.5Ð1.7 0.7Ð1.1 mm; not overgrown; not angular; asymmetrical; reniform (oblong) or oblong; Amphithalea: A. cuneifolia C.F. Ecklon & C.L.P. Zeyher compressed; with surface smooth; with visible radicle (BÐE), A. spp. (A). A, Fruits ( 3.3); B, seed ( 10); CÐ and cotyledon lobes; without external groove between D, testa ( 50, 1000); E, embryos ( 20).

1013 Genus: Coelidium (B.C. Vogel) G.W. Walpers frequent streaks; brown; with black overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Phylogenetic Number: 25.03. Wings absent. Raphe not visible (perhaps concealed by aril). Hilum visible; with faboid split; with the lips of Tribe: Podalyrieae. the faboid split lighter colored than the rest of the hilum and therefore conspicuous; larger than punctiform; 0.5 Subtribe: Xiphothecinae. mm long; with curved outline; elliptic; between cotyledon and radicle lobe; recessed; within rim or Species Studied—Species in Genus: 5 spp.—21 spp. halo. Hilum halo color lighter than testa. Hilum rim color darker than testa. Lens discernible (but concealed Fruit a legume; unilocular; 0.3Ð1.5 0.2Ð0.7 0.2 cm; under aril); not within corona, halo, or rim. Endosperm with persistent calyx; with calyx shorter than fruit; thin; covering entire embryo; adnate to embryo. without orifice formed by curving of fruit or fruit Cotyledons smooth; both outer faces convex; both the segments; straight; not plicate; not twisted; asymmetri- same thickness; both more or less of equal length; not cal; ovate or dolabriform; when asymmetrical with both folded; margin entire 180 degrees from base of radicle; sutures nearly straight; not inflated; compressed; with similar at apex; not concealing radicle; entire over beak; straight; with solid beak the same color and radicle; without lobes; with the interface division texture as fruit; rounded at apex; apex aligned with terminating at base of radicle; without margins re- longitudinal axis of fruit; rounded at base; base oblique cessed; green; inner face flat; glabrous around base of with longitudinal axis of fruit; with the apex and base radicle. Embryonic axis deflexed; oblique to length of uniform in texture; coriaceous; seed chambers exter- seed; without a joint evident between the radicle and nally invisible. Fruit margin not constricted; without the cotyledons. Radicle bulbose; lobe tip straight; sulcus; plain. Fruit wings absent. Fruit substipitate or oblique to cotyledons; centered between cotyledons; nonstipitate. Fruit with all layers dehiscing; splitting less than 1/2 length of cotyledons. Plumule rudimen- along sutures. Dehiscence of valves along both sutures; tary; glabrous. apical and down; passive. Replum invisible. Epicarp dull; monochrome; brown; pubescent and indurate; with Distribution: South Africa (southern Cape Province). 1 type of pubescence; tomentose, villous, or sericeous; with pubescence gray; with pubescence uniformly Notes: Polhill (1981p) noted that this genus is “doubtfully distributed; with simple hairs; pliable; with hair bases distinct from Amphithalea” (26.05). Granby (1980) plain; eglandular; without spines; not smooth; with monographed the genus, recognizing 19 species (7 elevated features; not veined; not tuberculate; faintly new), but Polhill recognized only ca. 15 species. wrinkled; not exfoliating; without cracks. Mesocarp Schutte (1998) carried out a cladistic analysis using thin; surface uniformly veined; 1-layered; without morphological and alkaloid characters of Amphithalea balsamic vesicles; without fibers; solid; coriaceous. and Coelidium with Liparia (26.01) and Xiphotheca Endocarp dull; monochrome; tan; smooth; nonseptate; (26.03) as outgroups. She concluded that the two genera chartaceous; not exfoliating; remaining fused to are synonymous, and transferred the Coelidium species mesocarp and epicarp; entire. Seeds 1 or 2; length to Amphithalea. According to Granby, fruits of several parallel with fruit length; neither overlapping nor species were unknown. touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick or triangular; straight. Aril fleshy; cupshaped (with tonguelike extension); covering less than 1/2 of seed; reddish brown.

Seed 1.5Ð5.5 0.9Ð3 0.5Ð1.2 mm; not overgrown; not angular; asymmetrical; oblong; compressed; with surface smooth; with (sligthly) or without visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar Coelidium: C. muraltioides G. Bentham (C), C. pageae sinus; without umbo on seed faces. Testa not adhering H.M.L. Bolus (B, DÐF), C. spp. (A). A, Fruits ( 5); BÐ to endocarp; dull; not modified by a bloom; colored; C, seed ( 7, 12.5); DÐE, testa ( 50, 1000); F, streaked and mottled; with frequent mottles; with embryos ( 7).

1015 Genus: Liparia C. Linnaeus radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to Phylogenetic Number: 25.04. endocarp; glossy or dull; not modified by a bloom; colored; monochrome, streaked (mainly), or mottled; Tribe: Podalyrieae. with frequent mottles; with frequent streaks; dark or reddish brown or tan; with black or purple overlay; Subtribe: Podalyriinae. glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe from hilum through Species Studied—Species in Genus: 8 spp.—20 spp. lens to base of seed and terminating or from hilum to lens (at base of seed); not bifurcating; darker than testa; Fruit a legume; unilocular; 1.5Ð3.5 0.5Ð1 cm; with reddish to blackish brown; flush. Hilum partially deciduous or persistent calyx; with calyx shorter than concealed; concealed by aril; with faboid split; with the fruit; without orifice formed by curving of fruit or fruit lips of the faboid split the same color as the rest of the segments; straight; not plicate; not twisted; asymmetri- hilum; larger than punctiform; 1 mm long; with curved cal or symmetrical; oblong (obliquely ovate); when outline; circular or elliptic; between cotyledon and asymmetrical with both sutures parallelly curved or 1 radicle lobe; recessed; not within corona, halo, or rim or straight and 1 curved suture; widest near middle or D- within halo. Hilum halo color darker than testa. Lens shaped; not inflated; compressed; with or without beak; discernible; equal to or greater than or less than 0.5 mm declined or straight; with solid beak the same color and in length; 0.5Ð1 mm long; with margins curved; texture as fruit; short tapered at apex; apex slightly roughly elliptic; not in groove of raphe; adjacent to oblique or aligned with longitudinal axis of fruit; hilum; 1.2Ð2 mm from hilum; mounded; similar color rounded or short tapered at base; base aligned with as or dissimilar color from testa; darker than testa; longitudinal axis of fruit; with the apex and base reddish to blackish brown; not within corona, halo, or uniform in texture; coriaceous; seed chambers exter- rim. Endosperm present (with or without 3 layers: nally invisible. Fruit margin not constricted; without white, cloudy, translucent (thin)); thick; covering entire sulcus; plain. Fruit wings absent. Fruit nonstipitate. embryo; adnate to embryo. Cotyledons smooth; both Fruit with all layers dehiscing; splitting along sutures. outer faces convex; both the same thickness; both more Dehiscence of valves along both sutures; apical and or less of equal length; not folded; margin entire 180 down; active or passive; with valves twisting. Replum degrees from base of radicle; similar at apex; not invisible. Epicarp dull; black or brown (dark); pubes- concealing radicle; entire over radicle; without lobes; cent and indurate or pubescent but soon deciduous; with with the interface division terminating at base of 1 type of pubescence; villous; with pubescence golden radicle; without margins recessed; tan or green; inner or gray; with pubescence uniformly distributed; with face flat; glabrous around base of radicle. Embryonic simple hairs; pliable; with hair bases plain; eglandular; axis deflexed; oblique to length of seed; with a joint without spines; with elevated features; veined or not evident between the radicle and the cotyledons. Radicle veined; reticulately veined; not tuberculate; not bulbose; lobe tip straight; deflexed and parallel to exfoliating; without cracks. Mesocarp thin; surface not cotyledon width; centered between cotyledons; less veined; 1-layered; without balsamic vesicles; without than 1/2 length of cotyledons. Plumule moderately fibers; solid; coriaceous. Endocarp dull; monochrome; developed or well developed; glabrous. brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Distribution: South Africa (Cape Province). Seeds 1Ð5; length parallel with or transverse to fruit length; neither overlapping nor touching; in 1 series. Notes: Bos (1967) monographed Liparia, s.s. Our treatment Funiculus measured; 1Ð2 mm long; of 1 length only; follows the reappraisal of Liparia and Priestleya A.-P. triangular or thick; straight. Aril fleshy; cupshaped (but de Candolle (26.02) by Schutte and Wyk (1994), who not a complete circle); covering less than 1/2 of seed; placed Priestleya into synonymy under Liparia. Wyk reddish brown to brown or yellow (pale). and Schutte (1995) carried out cladistic analyses of tribes Crotalarieae (27), Liparieae, and Podalyrieae (25) Seed 3.5Ð5.6 1.5Ð4 1.5Ð1.8 mm; not overgrown; not and their genera, and presented Priestleya as a good angular; asymmetrical; reniform (oblong) or oblong; genus distinct from Liparia. Wyk (personal communi- compressed; with surface smooth; with visible radicle cation, 1997; see also Schutte and Wyk 1998a) in- and cotyledon lobes; without external groove between

1017 Liparia: L. splendens (N.L. Burman) J.J. Bos & H.C.D. de Wit (AÐE). A, Valves ( 3.4); B, seeds ( 4.8); CÐD, testa ( 50, 1000); E, embryos ( 5).

1019 Genus: Cyclopia E.P. Ventenat ceous. Fracture lines absent. Rim absent. Raphe from hilum through lens to base of seed and terminating; not Phylogenetic Number: 25.05. bifurcating; lighter than testa; flush. Hilum partially concealed; concealed by aril; with faboid split; with the Tribe: Podalyrieae. lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 1Ð1.2 mm long; with Subtribe: Podalyriinae. curved outline; circular or elliptic; between cotyledon and radicle lobe; recessed; not within corona, halo, or Species Studied—Species in Genus: 6 spp.—23 spp. rim. Lens discernible or not discernible; less than 0.5 mm in length; with margins curved; circular; not in Fruit a legume; unilocular; 2Ð3.5 0.7Ð1.5 0.3Ð0.4 groove of raphe; adjacent to hilum; mounded or flush; (estimated) cm; with deciduous calyx; without similar color as or dissimilar color from testa; darker orifice formed by curving of fruit or fruit seg- than testa; reddish brown; not within corona, halo, or ments; straight; not plicate; not twisted; asym- rim. Endosperm thick; covering entire embryo; adnate metrical; oblong; when asymmetrical with 1 to embryo. Cotyledons smooth; both outer faces straight and 1 curved suture or both sutures convex; both the same thickness; both more or less of parallelly curved; widest near middle or D- equal length; not folded; margin entire 180 degrees shaped; not inflated; compressed; without beak; from base of radicle; similar at apex; not concealing short tapered at apex; apex aligned with longitudi- radicle; entire over radicle; without lobes; with the nal axis of fruit; rounded at base; base aligned to interface division terminating at base of radicle; green; oblique with longitudinal axis of fruit; with the inner face flat; glabrous around base of radicle. Embry- apex and base uniform in texture; coriaceous; seed onic axis oblique; perpendicular to length of seed. chambers externally invisible. Fruit margin not Radicle bulbose; lobe tip curved; deflexed and parallel constricted; without sulcus; plain. Fruit wings to cotyledon width; centered between cotyledons; less absent. Fruit substipitate. Fruit with all layers than 1/2 length of cotyledons. Plumule moderately dehiscing; splitting along sutures. Dehiscence of developed; glabrous. valves along both sutures; assumed apical and down; passive. Replum invisible. Epicarp dull; Distribution: South Africa (Cape). monochrome; brownish black or brown (reddish); glabrous; eglandular; without spines; not smooth; Notes: Kies (1951) monographed Cyclopia and noted that with elevated features; faintly reticulately veined; most species are used as a type of tea and “lately ‘bush- not tuberculate; not exfoliating; without cracks. tea’ has become a marketable commodity.” Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; reddish brown; cobwebby (around funicular area) or smooth; nonseptate; coriaceous; not exfoliating; remaining fused to mesocarp and epicarp. Seeds 2Ð11; length parallel with or transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; up to 1 mm long; of 1 length only; thick; straight. Aril fleshy; annular; covering less than 1/2 of seed; tan.

Seed 3.5Ð4 2Ð2.5 1.5Ð1.7 mm; not overgrown; not angular; asymmetrical; nearly oblong; compressed; without or with (barely) visible radicle and cotyledon lobes; with umbo on seed faces. Testa not adhering to endocarp; dull; not Cyclopia: C. galioides (B. Bergius) A.-P. de Candolle (A, modified by a bloom; colored; monochrome; CÐE), C. spp. (B). A, Valves ( 3); B, seeds ( 6); CÐ greenish brown or tan; glabrous; smooth; coria- D, testa ( 50, 1000); E, embryos ( 5).

1021 Genus: Podalyria C.L. von Willdenow absent. Raphe from hilum through lens to base of seed and terminating; lighter than testa; flush. Hilum visible; Phylogenetic Number: 25.06. with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than puncti- Tribe: Podalyrieae. form; 2 mm long; with straight outline; oblong; marginal according to radicle tip or between cotyledon Subtribe: Podalyriinae. and radicle lobe; recessed; within corona (when aril removed) or not within corona, halo, or rim. Hilum Species Studied—Species in Genus: 7 spp.—19 spp. corona color darker than testa. Lens not discernible or discernible; less than 0.5 mm or equal to or greater than Fruit a legume; unilocular; 2.5Ð3 1.2Ð1.3 0.3Ð0.4 0.5 mm in length; up to 0.5 mm long; with margins (estimated) cm; with deciduous calyx; without orifice straight or curved; irregular; not in groove of raphe; formed by curving of fruit or fruit segments; straight; adjacent to hilum; 1 mm from hilum; mounded; similar not plicate; not twisted; asymmetrical; when asym- color as testa; lighter or darker than testa; brownish red; metrical with 1 straight and 1 curved suture; widest not within corona, halo, or rim. Endosperm thick; near middle or D-shaped; not inflated; compressed; covering entire embryo; adnate to embryo. Cotyledons without beak; rounded at apex; apex aligned with smooth; both outer faces convex; both the same longitudinal axis of fruit; tapered at base; base aligned thickness; both more or less of equal length; not folded; with longitudinal axis of fruit; with the apex and base margin entire 180 degrees from base of radicle; similar uniform in texture; coriaceous; seed chambers exter- at apex; not concealing radicle; entire over radicle; nally invisible. Fruit margin not constricted; without without lobes; with the interface division terminating at sulcus; plain. Fruit wings absent. Fruit nonstipitate. base of radicle; without margins recessed; green; inner Fruit with all layers dehiscing; splitting along sutures. face flat; glabrous around base of radicle. Embryonic Dehiscence of valves along both sutures; assumed axis oblique; oblique to length of seed. Radicle bulbose; apical and down; passive. Replum invisible. Epicarp oblique to cotyledons; centered between cotyledons; dull; monochrome; green or brown (greenish); pubes- less than 1/2 length of cotyledons. Plumule moderately cent and indurate; with 1 type of pubescence; tomen- developed; glabrous. tose; with pubescence golden; with pubescence uniformly distributed; with simple hairs; pliable; with Distribution: South Africa (Cape region into Natal). hair bases plain; eglandular; without spines; not smooth; with recessed features; not veined; not tuberculate; minutely pitted; not exfoliating; without cracks. Mesocarp thin; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth; nonseptate; coriaceous; not exfoliating; remaining fused to mesocarp and epicarp. Seeds 8; length oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 0.5 mm long; of 1 length only; thick; straight. Aril fleshy; annular; covering less than 1/2 of seed; cream, tan, or white.

Seed 4.5Ð5 3Ð4 2.5Ð3 mm; not overgrown; not angular; asymmetrical; oblong; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; monochrome, mottled, or streaked; with frequent mottles; with frequent streaks; reddish, blackish, greenish, or dark brown, Podalyria: P. sericea (H.C. Andrews) R. Brown (A, CÐE), green, or tan; with black overlay; glabrous; smooth; P. spp. (B). A, Valves ( 3); B, seeds ( 7); CÐD, testa coriaceous. Fracture lines absent. Rim absent. Wings ( 50, 1000); E, embryos ( 5).

1023 Genus: Virgilia J.L.M. Poiret testa; flush. Hilum partially or fully concealed; concealed by aril or aril remnant; with faboid split; with the Phylogenetic Number: 25.08. lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 1.3Ð1.5 mm long; with Tribe: Podalyrieae. curved outline; elliptic; marginal according to radicle tip; recessed; not within corona, halo, or rim. Lens Subtribe: Podalyriinae. discernible or not discernible (at most a discolored darker and flush area); equal to or greater than 0.5 mm Species Studied—Species in Genus: 2 spp.—2 spp. in length; 0.8 mm long; with margins curved; elliptic; not in groove of raphe; adjacent to hilum; 1.5 mm from Fruit a legume; unilocular; 1.5Ð7 0.7Ð1.5 0.1Ð0.2 cm; hilum; flush; similar color as testa; darker than testa; with deciduous calyx; without orifice formed by gray (ish); not within corona, halo, or rim. Endosperm curving of fruit or fruit segments; slightly curved; not thick; covering entire embryo; adnate to embryo. plicate; not twisted; asymmetrical or symmetrical; Cotyledons smooth; both outer faces convex; both the linear to oblong; when asymmetrical with both sutures same thickness; both more or less of equal length; not nearly straight; not inflated; flattened; without beak; folded; margin entire 180 degrees from base of radicle; rounded at apex; apex oblique with longitudinal axis of similar at apex; not concealing radicle; entire over fruit; tapered at base; base oblique (slightly) with radicle; without lobes; with the interface division longitudinal axis of fruit; with the apex and base terminating at base of radicle; without margins re- uniform in texture; coriaceous or ligneous; seed cessed; green (ish); inner face flat; glabrous around chambers externally visible; with the raised seed base of radicle. Embryonic axis oblique; oblique to chambers not torulose. Fruit margin slightly constricted length of seed. Radicle linear; oblique to cotyledons; along both margins; without sulcus; plain. Fruit wings centered between cotyledons; less than 1/2 length of absent. Fruit substipitate to nonstipitate. Fruit with all cotyledons. Plumule moderately developed; glabrous. layers dehiscing; splitting along sutures. Dehiscence of valves along 1 suture or both sutures; assumed apical Distribution: South Africa (southern Cape Province). and down; passive. Replum invisible. Epicarp dull; monochrome; brown; pubescent and indurate (widely Notes: Wyk (1986) monographed Virgilia. scattered); with 1 type of pubescence; pilose; with pubescence golden; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; transversely veined relative to fruit length; not tuberculate; not exfoliating; with cracks; cracking transverse to fruit length. Mesocarp apparently absent. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp. Seeds 4Ð6; length oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only. Aril fleshy; annular; covering less than 1/2 of seed; tan or white.

Seed 5Ð9 3Ð6 2.5Ð3.5 mm; not overgrown; angular or not angular; symmetrical or asymmetrical; sub circular to oblong to reniform; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; glossy to dull; not modified by a bloom; colored; monochrome; black, brown, or yellow (-brown); glabrous; smooth; coria- Virgilia: V. oroboides (B. Bergius) T.M. Salter (A, CÐE), V. ceous. Fracture lines absent. Rim absent. Wings absent. spp. (B). A, Valves ( 2); B, seeds ( 4.5); CÐD, testa Raphe from hilum to lens; not bifurcating; color of ( 50, 1000); E, embryos ( 5).

1025 Genus: Calpurnia E.H.F. Meyer visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to Phylogenetic Number: 25.09. endocarp; glossy to dull; not modified by a bloom; colored or clear; monochrome or mottled; with infre- Tribe: Podalyrieae. quent mottles; brown or tan; with brown (darker) overlay; glabrous; smooth; chartaceous. Fracture lines Subtribe: Podalyriinae. absent. Rim absent. Wings absent. Raphe visible or not visible; from hilum through lens to base of seed and Species Studied—Species in Genus: 4–5 spp.—7 spp. terminating; not bifurcating; darker than testa; brown; flush. Hilum partially concealed; concealed by aril; Fruit a legume; unilocular; 2.2Ð13 0.5Ð1.9 1Ð3 cm; with faboid split; with the lips of the faboid split the with persistent or deciduous calyx; with calyx shorter same color as the rest of the hilum; larger than puncti- than fruit; without orifice formed by curving of fruit or form; 0.4Ð0.6 mm long; with curved outline; circular to fruit segments; straight or curved (slightly); not plicate; elliptic; subapical to radicle tip or apical according to not twisted; asymmetrical or symmetrical; linear or radicle tip but marginal according to seed length; falcate (slightly); when asymmetrical with both sutures recessed; within rim. Hilum rim color of testa. Lens parallelly curved; not inflated; flattened; with beak; discernible; equal to or greater than 0.5 mm in length; straight or hooked; with solid beak the same color and 0.5Ð0.8 mm long; with margins straight; diamond- texture as fruit; tapered or rounded at apex; apex shaped; not in groove of raphe; adjacent to or confluent aligned or oblique with longitudinal axis of fruit; long with hilum; 1 mm from hilum; mounded; similar color tapered or tapered at base; base aligned or oblique with as testa; lighter or darker than testa; brown; not within longitudinal axis of fruit; with the apex and base corona, halo, or rim. Endosperm thick; covering entire uniform in texture; chartaceous to coriaceous (sub); embryo; adnate to testa or embryo. Cotyledons not seed chambers externally visible; with the raised seed smooth; apically sulcate; both outer faces convex; both chambers not torulose. Fruit margin not constricted; the same thickness; both more or less of equal length; without sulcus; embellished; with wing. Fruit wing 1; not folded; margin entire 180 degrees from base of 0.2Ð2.5 mm wide; sutural; on 1 suture. Fruit stipitate or radicle; similar at apex; not concealing radicle; notched substipitate; with the stipe 4Ð15 mm long. Fruit at radicle; without lobes; with the interface division indehiscent. Replum invisible. Epicarp dull; mono- terminating at base of radicle; without margins re- chrome or multicolored; mottled; greenish tan; with cessed; yellow; inner face flat; glabrous around base of brown overlay; glabrous or pubescent and indurate; radicle. Embryonic axis right angled; parallel to length with 1 type of pubescence; sparsely strigose; with of seed; without a joint evident between the radicle and pubescence golden; with pubescence uniformly the cotyledons. Radicle differentiated from cotyledon; distributed; with simple hairs; stiff; with hair bases linear; lobe tip curved; oblique to cotyledons or with plain; eglandular; without spines; not smooth; with 90-degree turn; centered between cotyledons; less than elevated features; reticulately veined; not tuberculate; 1/2 length of cotyledons. Plumule moderately devel- sometimes dotted; not exfoliating; without cracks. oped; glabrous. Mesocarp present or absent; thin; surface not veined; 1- layered; without balsamic vesicles; solid; chartaceous. Distribution: South Africa (eastern Cape) and southern Endocarp glossy; monochrome; golden tan; smooth; India. nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; remaining fused to Notes: Brummitt (1967) treated Calpurnia aurea and epicarp; entire. Seeds 2Ð10; length oblique or transverse suggested that the genus has six or seven species. to fruit length; neither overlapping nor touching; in 1 Yakovlev (1971) recognized nine species in the genus. series. Funiculus measured; 1.5Ð2.5 mm long; of 1 length only; filiform; curved. Aril dry; rim-aril; entire; cream or tan.

Seed 4Ð10.5 2.5Ð6.5 1.5Ð3.5 mm; not overgrown; not angular; asymmetrical; elliptic to ovate to irregular; Calpurnia: C. aurea (W. Aiton) G. Bentham (CÐE), C. spp. terete or compressed; with surface smooth; without (AÐB). A, Fruits ( 1.1); B, seeds ( 6.3); CÐD, testa ( 50, 1000); E, embryos ( 5).

1027 Hypocalypteae (26.01) overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Raphe visible or not visible; from Genus: Hypocalyptus C.P. Thunberg hilum through lens to base of seed and terminating; not bifurcating; darker than testa; reddish brown or black; Phylogenetic Number: 26.01. flush. Hilum partially concealed; concealed by aril; with faboid split; with the lips of the faboid split the Tribe: Hypocalypteae. same color as the rest of the hilum; larger than punctiform; 1 mm long; with straight outline; oblong; Species Studied—Species in Genus: 3 spp.—3 spp. between cotyledon and radicle lobe or marginal according to radicle tip; recessed; not within corona, Fruit a legume; unilocular; 0.2Ð6.5 ϫ 0.4Ð1.4 ϫ 0.25 cm; halo, or rim. Lens discernible or not discernible; less with persistent calyx; with calyx shorter than fruit; than 0.5 mm in length; with margins curved; circular without orifice formed by curving of fruit or fruit (outline difficult to see); not in groove of raphe; segments; straight to curved (slightly); not plicate; not adjacent to hilum; 1 mm from hilum; mounded; twisted; symmetrical or asymmetrical; linear, oblong, or dissimilar color from testa; darker than testa; reddish obovate; when asymmetrical with both sutures nearly brown or black; not within corona, halo, or rim. straight; not inflated or inflated; compressed or terete; Endosperm thick; covering entire embryo; adnate to with beak; short tapered at apex; apex aligned with embryo. Cotyledons smooth; both outer faces convex; longitudinal axis of fruit; tapered at base; base oblique both the same thickness; both more or less of equal with longitudinal axis of fruit; with the apex and base length; not folded; margin entire 180 degrees from base uniform in texture; coriaceous or chartaceous; seed of radicle; similar at apex; not concealing radicle; entire chambers externally visible; with the raised seed over radicle; without lobes; with the interface division chambers torulose. Fruit margin constricted or not terminating at base of radicle; without margins re- constricted; slightly constricted along both margins; cessed; yellow; inner face flat; glabrous around base of without sulcus; plain. Fruit wings absent. Fruit radicle. Embryonic axis deflexed; oblique to length of substipitate. Fruit with all layers dehiscing; splitting seed; without a joint evident between the radicle and along sutures. Dehiscence of valves along both sutures; the cotyledons. Radicle linear; deflexed and parallel to medial and up and down; passive. Replum invisible. cotyledon width; centered between cotyledons; less Epicarp dull; reddish or blackish brown or brown, than 1/2 length of cotyledons. Plumule rudimentary black, or yellow (pale); glabrous; eglandular; without glabrous. spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Distribution: South Africa (southern Cape Province). Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Notes: Schutte and Wyk (1998a,b) carried out extensive Endocarp dull; monochrome; tan; smooth; nonseptate; cladistic analyses of the tribes Liparieae and chartaceous; not exfoliating; remaining fused to Podalyrieae and related genera. They concluded that all mesocarp and epicarp; entire. Seeds 1Ð6; length parallel genera of Liparieae, except Hypocalyptus, should be with fruit length; neither overlapping nor touching; in 1 transferred to Podalyrieae and that a monotypic tribe series. Funiculus less than 0.5 mm long; of 1 length should be created for Hypocalyptus. Dahlgren (1972) only; triangular; straight. Aril fleshy; cupshaped; monographed Hypocalyptus and correctly noted the covering less than 1/2 of seed; white. fruit variation. Hypocalyptus coluteoides (J.B.A.P. de M. de Lamarck) R.M.T. Dahlgren has an inflated Seed 3.4Ð8 ϫ 2Ð3 ϫ 1.3Ð3 mm; not overgrown; not legume like Colutea (16.05), while the other two angular; asymmetrical; reniform (oblong) or ovate; species have linear to obovate legumes that are not compressed; with surface smooth; with (faintly) or inflated without visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull and glossy or dull; not modified by a bloom; colored; monochrome or mottled to streaked; with frequent mottles; with frequent Hypocalyptus: H. sophoroides (P.J. Bergius) H.E. Baillon streaks; black to brown (dark); with black (brownish) (AÐE). A, Fruits and valve (ϫ 1.7); B, seeds (ϫ 5.5); CÐ D, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5). 1028 A

1029 Crotalarieae (27.01Ð27.11) concealed; concealed by funicular remnant; with faboid split; with the lips of the faboid split the same color as Genus: Pearsonia R.A. Dummer the rest of the hilum; punctiform; between cotyledon and radicle lobe; flush; within rim. Hilum rim color of Phylogenetic Number: 27.01. or lighter than testa. Lens discernible; equal to or greater than 0.5 mm in length; 0.5Ð0.7 mm long; with Tribe: Crotalarieae. margins straight or curved; linear or oblong; not in groove of raphe; adjacent to hilum; 0.1 mm from hilum; Species Studied—Species in Genus: 7 spp.—12 spp. mounded; dissimilar color from testa; darker than testa; reddish brown; not within corona, halo, or rim. En- Fruit a legume; unilocular; 0.5Ð3.8 ϫ 0.35Ð0.8 ϫ 0.7Ð0.8 dosperm thin; covering entire embryo; adnate to cm; with persistent calyx; with calyx equal in length to embryo. Cotyledons smooth; both outer faces convex; or shorter than fruit; without orifice formed by curving both the same thickness; both more or less of equal of fruit or fruit segments; straight or curved (slightly); length; not folded; similar at apex; not concealing not plicate; not twisted; asymmetrical or symmetrical; radicle; entire over radicle; without lobes; with the elliptic or oblong (lanceolate); when asymmetrical with interface division terminating at base of radicle; without both sutures nearly straight; not inflated; compressed; margins recessed; yellow; inner face flat; glabrous without beak; tapered at apex; apex aligned with around base of radicle. Embryonic axis deflexed; longitudinal axis of fruit; tapered at base; base aligned oblique to length of seed. Radicle bulbose; deflexed and with longitudinal axis of fruit; with the apex and base parallel to cotyledon length; centered between cotyle- uniform in texture; coriaceous; seed chambers exter- dons; less than 1/2 length of cotyledons. Plumule nally invisible. Fruit margin not constricted; without rudimentary; glabrous. sulcus; plain. Fruit wings absent. Fruit nonstipitate. Fruit with all layers dehiscing (assumed); splitting Distribution: Africa (south of the equator) and Madagascar along sutures. Dehiscence of valves along both sutures; (1 sp.). apical and down; assumed active; with valves twisting (assumed). Replum invisible. Epicarp dull; mono- Notes: Polhill (1981q) broadly defined the tribe Crotalarieae chrome; brown (because of numerous golden hairs); with two generic groups. The first group, which does pubescent and indurate; with 1 type of pubescence; not have a two-lipped calyx, formed a tight cluster pilose, tomentose, villous, or sericeous; with pubes- around Lebeckia (27.10) in southern Africa. The second cence golden; with simple hairs; pliable; with hair bases group, which has a two-lipped calyx, had more scat- plain; eglandular; without spines; not smooth; with tered distributions and uncertain affinities. Wyk (1991) elevated features; veined or not veined; reticulately followed Polhill (1981q) and transferred Argyrolobium veined; not tuberculate; shagreen (may not be best (30.03) from Genisteae (30) to the second group. answer); not exfoliating; without cracks. Mesocarp thin; Crotalarieae and related tribes are rich in alkaloids, surface not veined; 1-layered; without balsamic which have been extensively studied in the last decade vesicles; without fibers; solid; coriaceous. Endocarp (Hussain et al. 1988; Wyk and Verdoorn 1989a,b,c, dull; monochrome; brown; smooth; nonseptate; 1990, 1991a,b; Wyk et al. 1989, 1993; Verdoorn and chartaceous; not exfoliating; remaining fused to Wyk 1990, 1991). Using chemical and morphological mesocarp and epicarp; entire. Seeds 1Ð15; length data, Polhill (1994a,b) and Wyk and Schutte (1995) parallel with fruit length; neither overlapping nor restricted Crotalarieae to the genera without a two- touching; in 1 series. Funiculus less than 0.5 mm long; lipped calyx and transferred those with a two-lipped of 1 length only; thick; straight. Aril absent. calyx to Genisteae, Anarthrophyllum (30.06), Argyrolobium (30.03), Dichilus (30.02), Melolobium Seed 2Ð2.5 ϫ 1.7Ð2 ϫ 1.4Ð1.5 mm; not overgrown; not (30.01), and Sellocharis (30.07), except Lebeckia. They angular; asymmetrical; obliquely cordate or reniform; also more or less inverted the generic order within the compressed; with or without visible radicle and first group according to Wyk and Schutte’s cladistic cotyledon lobes; without umbo on seed faces. Testa not analysis for the genera of Crotalarieae in the narrow adhering to endocarp; glossy; not modified by a bloom; sense. Polhill (1974) monographed Pearsonia. colored; monochrome; brown or tan; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings Pearsonia: P. cajanifolia (C.E.O. Kuntze) R.M. Polhill (CÐ absent. Raphe not visible. Hilum visible or fully E), P. spp. (AÐB). A, Fruit (ϫ 2.8); B, seeds (ϫ 8); CÐ D, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10). 1030 A

1031 Genus: Rothia C.H. Persoon not in groove of raphe; confluent (or nearly so) with hilum; mounded; dissimilar color from testa; darker Phylogenetic Number: 27.02. than testa; reddish brown; not within corona, halo, or rim. Endosperm thin; covering entire embryo; adnate to Tribe: Crotalarieae. testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; Species Studied—Species in Genus: 2 spp.—2 spp. not folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing radicle; entire Fruit a legume; unilocular; 3.5Ð4.5 ϫ 0.15Ð0.2 ϫ 0.15Ð0.17 over radicle; without lobes; with the interface division cm; with persistent or deciduous calyx; with calyx terminating at base of radicle; without margins re- shorter than fruit; without orifice formed by curving of cessed; tan or yellow; inner face flat; glabrous around fruit or fruit segments; straight; not plicate; not twisted; base of radicle. Embryonic axis deflexed; oblique to symmetrical; linear; not inflated; flattened; without length of seed. Radicle linear; deflexed and parallel to beak; tapered at apex; apex aligned with longitudinal cotyledon length; not centered between cotyledons axis of fruit; tapered at base; base aligned with longitu- (radicle outside 1 cotyledon and inside other, therefore dinal axis of fruit; with the apex and base uniform in junctions for each cotyledon different); less than 1/2 texture; membranous; seed chambers externally length of cotyledons. Plumule rudimentary; glabrous. invisible. Fruit margin not constricted; without sulcus; plain. Fruit wings absent. Fruit nonstipitate. Fruit with Distribution: Africa (1 sp.) and Baluchistan to Australia (1 all layers dehiscing; splitting along suture. Dehiscence sp.). of valves along 1 suture; apical and down; active; with valves enrolling (when mature gaping along adaxial suture). Replum invisible. Epicarp dull; monochrome; reddish brown; pubescent and indurate; with hairs appressed; with 1 type of pubescence; with pubescence golden; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent. Endocarp glossy; monochrome; brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to epicarp; entire. Seeds 30Ð40 (estimated); length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; up to 1 mm long; of 1 length only; filiform; contorted. Aril absent.

Seed 1Ð1.4 ϫ 0.8Ð1 ϫ 0.6Ð0.7 mm; not overgrown; not angular; asymmetrical; mitaform; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; conspicuously to faintly mottled or monochrome; with frequent mottles; brown, green, or tan; with purple or tan overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform (may be the smallest in the subfamily); between cotyledon and radicle lobe; recessed; within rim (with or without a black patch around rim). Hilum rim color of testa. Lens Rothia: R. indica (C. Linnaeus) G.C. Druce (CÐE), R. spp. discernible (as large as hilar area (including rim)); less (AÐB). A, Fruits (ϫ 2.5); B, seeds (ϫ 11); CÐD, testa than 0.5 mm in length; with margins curved; circular; (ϫ 50, ϫ 1000); E, embryos (ϫ 18).

1033 Genus: Robynsiophyton R. Wilczek within corona, halo, or rim. Endosperm thin; covering entire embryo; adnate to testa. Cotyledons smooth; both Phylogenetic Number: 27.03. outer faces convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 Tribe: Crotalarieae. degrees from base of radicle; similar at apex; not concealing radicle; entire over radicle; without lobes; Species Studied—Species in Genus: 1 sp.—1 sp. with the interface division terminating at base of radicle; without margins recessed; yellow; inner face Fruit a legume; unilocular; 0.6Ð1 ϫ 0.3Ð0.45 ϫ 0.1Ð0.15 flat; glabrous around base of radicle. Embryonic axis cm; with persistent calyx; with calyx shorter than fruit; deflexed; oblique to length of seed. Radicle linear; without orifice formed by curving of fruit or fruit deflexed and parallel to cotyledon length; centered segments; straight; not plicate; not twisted; symmetri- between cotyledons; 1/2 to nearly length of cotyledons. cal; oblong (ovate), lanceolate (linear), or linear; not Plumule moderately developed; glabrous. inflated; compressed to flattened; without beak; short tapered at apex; apex aligned with longitudinal axis of Distribution: South-central Africa. fruit; rounded at base; base aligned with longitudinal axis of fruit; with the apex and base uniform in texture; Notes: Our observations are based on only one sample membranous; seed chambers externally invisible. Fruit augmented by data from the literature. margin not constricted; without sulcus; plain. Fruit wings absent. Fruit nonstipitate. Fruit with all layers dehiscing; splitting along suture. Dehiscence of valves along apparently 1 suture; assumed apical and down; passive. Replum invisible. Epicarp dull; monochrome; brown; pubescent and indurate; with 1 type of pubescence; pilose or tomentose; with pubescence golden; with simple hairs; pliable; with hair bases plain; without spines; apparently smooth; not veined; not tuberculate; not exfoliating; without cracks. Mesocarp absent or present; thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; chartaceous. Endocarp dull; monochrome; brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp or to epicarp; entire. Seeds 4Ð8; length parallel with or transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 1.5Ð1.7 mm long; of 1 length only; filiform; straight. Aril absent.

Seed 1Ð1.2 ϫ 1Ð1.2 ϫ 0.4Ð0.5 mm; not overgrown; not angular; asymmetrical; mitaform; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; greenish brown; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; between cotyledon and radicle lobe; recessed; within rim. Hilum rim color of testa. Lens discernible; less than 0.5 mm in length; with margins Robynsiophyton: R. vanderystii R. Wilczek (AÐE). A, Fruits straight; wedge-shaped; not in groove of raphe; (ϫ 5.1); B, seeds (ϫ 12.5); CÐD, testa (ϫ 50, ϫ 1000); adjacent to hilum; mounded; same color as testa; not E, embryos (ϫ 20).

1035 Genus: Spartidium A.N. Pomel recessed; within rim. Hilum rim color of testa. Lens discernible; equal to or greater than 0.5 mm in length; Phylogenetic Number: 27.04. 1.4Ð1.8 mm long; with margins straight or curved; oblong; not in groove of raphe; confluent with hilum; Tribe: Crotalarieae. mounded; dissimilar color from testa; darker than testa; brown; not within corona, halo, or rim. Endosperm thin; Species Studied—Species in Genus: 1 sp.—1 sp. covering entire embryo; adnate to testa. Cotyledons smooth; both outer faces convex; both the same Fruit a legume; unilocular; 3Ð4.3 ϫ 0.8Ð1 ϫ 0.08Ð0.17 cm; thickness; both more or less of equal length; not folded; with persistent calyx; with calyx shorter than fruit; margin entire 180 degrees from base of radicle; similar without orifice formed by curving of fruit or fruit at apex; not concealing radicle; split over radicle; with segments; straight; not plicate; not twisted or twisted; lobes; without margins recessed; reddish tan; inner face symmetrical or asymmetrical; oblong; when asym- flat; glabrous around base of radicle. Embryonic axis metrical with 1 straight and 1 curved suture or both deflexed; oblique to length of seed; without a joint sutures nearly straight; widest near middle or D- evident between the radicle and the cotyledons. Radicle shaped; not inflated; flattened; without beak; rounded linear; deflexed and parallel to cotyledon length; at apex; apex aligned or oblique with longitudinal axis centered between cotyledons; less than 1/2 length of of fruit; short tapered at base; base aligned with cotyledons. Plumule rudimentary; glabrous. longitudinal axis of fruit; with the apex and base uniform in texture; membranous; seed chambers Distribution: North Africa (Libya to Algeria and probably externally visible; with the raised seed chambers Morocco). torulose. Fruit margin not constricted; without sulcus; plain. Fruit wings absent. Fruit substipitate. Fruit Notes: Polhill (1981q) noted that this genus is “virtually assumed indehiscent. Replum invisible or visible (valve indistinguishable from Lebeckia” (27.10). However, could irregularly separate from sutures). Epicarp alkaloid data support keeping Spartidium separate from semiglossy; monochrome; reddish brown; glabrous; Lebeckia (Wyk et al. 1989). eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; chartaceous. Endocarp dull; monochrome; tan; smooth and scurfy (smooth around seeds and scurfy between seeds); nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 2Ð6; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 4 mm long; of 1 length only; filiform; hooked. Aril absent.

Seed 3.5Ð4 ϫ 2Ð2.5 ϫ 1.5Ð1.8 mm; not overgrown; not angular or angular (somewhat); asymmetrical; oblong or reniform; terete; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; reddish brown or tan (to reddish); glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible or fully concealed; concealed by funicular remnant; with faboid split; with the lips of the Spartidium: S. saharae (E.S.-C. Cosson & E. Reboul) A.N. faboid split the same color as the rest of the hilum; Pomel (AÐE). A, Fruit (ϫ 1.7); B, seeds (ϫ 8.4); CÐD, punctiform; between cotyledon and radicle lobe; testa (ϫ 50, ϫ 1000); E, embryos (ϫ 4.6).

1037 Genus: Rafnia C.P. Thunberg Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid Phylogenetic Number: 27.05. split the same color as the rest of the hilum; larger than punctiform; 0.5Ð0.7 mm long; with straight outline; Tribe: Crotalarieae. oblong; between cotyledon and radicle lobe; recessed; within rim. Hilum rim color darker (slightly to black) Species Studied—Species in Genus: 9 spp.—ca. 23 spp. than testa. Lens discernible; equal to or greater than 0.5 mm in length; 0.5Ð1 mm long; with margins straight or Fruit a legume; unilocular; 1.5Ð3 ϫ 0.5Ð0.8 ϫ 0.2Ð0.3 cm; curved; narrowly oblong; not in groove of raphe; with persistent calyx; with calyx shorter than fruit; adjacent to hilum; mounded; dissimilar color from without orifice formed by curving of fruit or fruit testa; darker than testa; black or brown (dark to reddish segments; straight; not plicate; not twisted; asymmetri- to greenish); not within corona, halo, or rim. En- cal; lanceolate (linear), linear, or oblong (narrowly to dosperm thin; covering entire embryo; adnate to testa. broadly); when asymmetrical with 1 straight and 1 Cotyledons smooth; both outer faces convex; both the curved suture or both sutures parallelly curved; widest same thickness; both more or less of equal length; not at base or near apex; not inflated; compressed; without folded; margin entire 180 degrees from base of radicle; beak; short tapered at apex; apex aligned with longitu- similar at apex; not concealing radicle; entire over dinal axis of fruit; tapered at base; base aligned with radicle; without lobes; with the interface division longitudinal axis of fruit; with the apex and base terminating at base of radicle; without margins re- uniform in texture; coriaceous; seed chambers exter- cessed; brownish red; inner face flat; glabrous around nally invisible. Fruit margin not constricted; without base of radicle. Embryonic axis deflexed; oblique to sulcus; embellished to plain. Fruit wing present or length of seed. Radicle bulbose; deflexed and parallel to absent; 1; 1Ð2 mm wide; sutural (along upper suture cotyledon width; centered between cotyledons; less and usually best developed near calyx); on 1 suture. than 1/2 length of cotyledons. Plumule rudimentary; Fruit stipitate to substipitate to nonstipitate; with the glabrous. stipe up to 10 mm long. Fruit with all layers dehiscing; splitting along sutures. Dehiscence of valves along both Distribution: South Africa (southwestern Cape Province to sutures; apical and down; active; with valves enrolling Natal). or twisting. Replum invisible. Epicarp dull; monochrome; black or brown (dirty); glabrous; eglandular; Notes: Number of species and distribution taken from Wyk without spines; not smooth; with elevated features; (1991) and Wyk and Schutte (1995). reticulately veined; not tuberculate; lenticular (tannish); not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; nearly spongy or solid; coriaceous. Endocarp dull or glossy; monochrome; brown (to reddish); reticulate or hairy (appressed and silver); nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seed 1 (assumed); assumed length parallel with fruit length. Funiculus less than 0.5 mm long; thick; straight. Aril absent.

Seed 3.7Ð4 ϫ 2.2Ð2.5 ϫ 0.8Ð2 mm; not overgrown; not angular; asymmetrical; reniform; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled and streaked; with frequent mottles; with frequent streaks; reddish brown or tan; with black (grayish) Rafnia: R. amplexicaulis (C. Linnaeus) C.P. Thunberg (CÐ overlay; glabrous; smooth or not smooth; with elevated E), R. spp. (AÐB). A, Fruits (ϫ 2); B, seeds (ϫ 6); CÐD, features; wrinkled; coriaceous. Fracture lines absent. testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

1039 Genus: Aspalathus C. Linnaeus (many); length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less Phylogenetic Number: 27.06. than 0.5 mm long; of 1 length only; thick; straight. Aril absent. Tribe: Crotalarieae. Seed 2Ð3.5 ϫ 1.5Ð3 ϫ 1Ð2 mm; not overgrown; not angular Species Studied—Species in Genus: 24 spp.—ca. 278 spp. to angular; asymmetrical; mitaform, quadrangular, rhombic (sub), or circular (sub); compressed to terete; Fruit a legume or nutlet (a few spp.); unilocular; 0.4Ð3 ϫ with visible radicle and cotyledon lobes; without umbo 0.2Ð6.3 ϫ 0.2Ð0.3 cm; with persistent calyx or decidu- on seed faces. Testa not adhering to endocarp; dull or ous calyx; with calyx longer or shorter than fruit; glossy; not modified by a bloom; colored; monochrome without orifice formed by curving of fruit or fruit or mottled and streaked; with frequent mottles; with segments; straight to curved (slightly); not plicate; not frequent streaks; black, brown (to yellowish or reddish), twisted; asymmetrical or symmetrical; linear, obovate, tan, or white; with black or gray overlay (green, brown, ovate (and rhombic), rhombic (sub), lanceolate, falcate, purple or dark); glabrous; smooth or not smooth; with or rectangular; when asymmetrical with 1 straight and 1 elevated features; warty; coriaceous. Fracture lines curved suture, both sutures parallelly curved, or both absent. Rim absent. Raphe not visible. Hilum visible or sutures unequally curved; widest near middle or D- partially concealed; concealed by funicular remnant; shaped; slightly inflated or not inflated; compressed or with faboid split; with the lips of the faboid split the terete; without beak; long tapered to tapered to short same color as the rest of the hilum; punctiform; tapered to rounded at apex; apex aligned to oblique marginal according to radicle tip or between cotyledon with longitudinal axis of fruit; tapered to short tapered and radicle lobe; recessed; within rim, corona, and halo to rounded (nearly) at base; base aligned to oblique or not within corona, halo, or rim. Hilum corona color with longitudinal axis of fruit; with the apex and base darker than testa. Hilum halo color lighter or darker uniform in texture; coriaceous or ligneous; seed than testa. Hilum rim color of testa. Lens discernible or chambers externally visible or invisible; with the raised not discernible; equal to or greater than 0.5 mm in seed chambers not torulose. Fruit margin not con- length; 0.7 mm long; with margins straight or curved; stricted; without sulcus; plain. Fruit wings absent. Fruit oblong (with or without lighter colored medial line); not stipitate to substipitate to nonstipitate; with the stipe up in groove of raphe; adjacent to hilum; to 0.5 mm from to 7 mm long. Fruit with all layers dehiscing (calyx hilum; slightly mounded; similar color as testa; darker may retard dehiscence for ovate pods shorter than than testa; black or brown (reddish); within halo and calyx); splitting along sutures. Dehiscence of valves corona. Lens corona color darker than testa. Lens halo along 1 suture (for ovate fruits) or both sutures; apical color lighter than testa. Endosperm thin; covering entire and down or basal and up (for ovate fruits); active or embryo; adnate to testa. Cotyledons smooth; both outer passive; with valves twisting. Replum invisible. Epicarp faces convex; both the same thickness; both more or dull; monochrome; black, brown (reddish), or tan; less of equal length; not folded; margin entire 180 glabrous, glabrate, or pubescent and indurate; with 1 degrees from base of radicle; similar at apex; not type of pubescence; villous, tomentose, or sericeous; concealing radicle; entire over radicle; without lobes; with pubescence gray or golden; with pubescence with the interface division terminating at base of uniformly distributed or apical pubescence different radicle; without margins recessed; tan; inner face flat; from basal pubescence; with apical 1/4 tomentose and glabrous around base of radicle. Embryonic axis basal 3/4 glabrous; with simple hairs; pliable; with hair deflexed; oblique to length of seed. Radicle bulbose; bases plain; eglandular; without spines; smooth or not lobe tip straight; deflexed and parallel to cotyledon smooth; with elevated features; reticulately veined; not length not centered between cotyledons (radicle outside tuberculate; rugose; not exfoliating; without cracks. 1 cotyledon and inside other, therefore junctions for Mesocarp thin; surface not veined; 1-layered; without each cotyledon different); less than 1/2 length of balsamic vesicles; without fibers; solid coriaceous to cotyledons to 1/2 to nearly length of cotyledons. ligneous (including subligneous). Endocarp dull; Plumule rudimentary; glabrous. monochrome; brown or tan; smooth; subseptate to nonseptate; with septa thicker than paper, firm; with Distribution: South Africa (southwestern Cape Province to septa eglandular; chartaceous; not exfoliating; remain- Natal). ing fused to mesocarp and epicarp; entire. Seeds 1Ð18

1040 Notes: Dalhgren (1960, 1963a,b, 1965, 1968) monographed Aspalathus in a series of papers. His fruit and seed data were used to enlarge our limited database. In discussing the legume morphology, Dahlgren (1963b, pp. 98Ð105) noted that there is no typical Aspalathus legume, though the short, more or less triangular and compressed legume is most common. His figure 15 is a useful compilation of legume shapes and hair types and patterns. Legumes usually dehisce either actively or passively on the plant. Some species have legumes that remain closed and fall to the ground while closed, acting as a “nutlet”; for example, A. bodkinii H. Bolus and probably A. argyrella P. MacOwan, A. comptonii R.M.T. Dahlgren, and A. villosa C.P. Thunberg. The number of species and distribution used here were taken from Wyk (1991) and Wyk and Schutte (1995).

1041 Aspalathus: A. linearis (N.L. Burman) R.M.T. Dahlgren (CÐE), A. spp. (AÐB). A, Fruits (ϫ 2.1); B, seeds (ϫ 8.4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 12).

1043 Genus: Crotalaria C. Linnaeus neither overlapping nor touching; in 1 series. Funiculus measured; up to 3 mm long; of 1 length only; filiform Phylogenetic Number: 27.07. or thick; S-curved or curved. Aril absent or present (rarely); dry; rim-aril (in section Chrysocalycinae (G. Tribe: Crotalarieae. Bentham) E.G. Baker subsection Stipulosae (E.G. Baker) F.A. Bisby & R.M. Polhill); covering less than Species Studied—Species in Genus: Ca. 200 spp.—ca. 600 1/2 of seed; brown or tan. spp. Seed 1Ð8 ϫ 1Ð6 ϫ 0.75Ð3 mm; not overgrown; not angular Fruit a legume; unilocular; 0.3Ð12 ϫ 0.3Ð1.7 ϫ 1Ð2 cm; to angular; asymmetrical; reniform (and obliquely with deciduous or persistent calyx; with calyx shorter reniform), mitaform, cordate (obliquely), or circular than fruit; without orifice formed by curving of fruit or (sub); compressed or flattened; with visible radicle and fruit segments; straight or curved (slightly); not plicate; cotyledon lobes; with deep, with shallow, or without not twisted; asymmetrical or symmetrical; oblong, hilar sinus; without umbo on seed faces. Testa not elliptic (clavate, oblong, or circular), linear (and linear- adhering to endocarp; usually glossy or dull; not oblong), or circular; when asymmetrical with 1 straight modified by a bloom; colored; monochrome, mottled and 1 curved suture or both sutures parallelly curved; and streaked, or bichrome (primarily dark reddish-black widest near middle or D-shaped; inflated (some species with a white patch over radicle tip and with or without markedly inflated); terete or compressed (especially white patch at junction of radicle and cotyledon lobes species in subgenus Priotropis (R. Wight & G.A.W. or a continuous white patch to radicle tip to radicle Arnott) C.D.F. Meisner); without or with beak; straight; cotyledon junction, C. spectabilis A.W. Roth); with with solid beak the same color and texture as fruit; frequent mottles; with frequent streaks; brown (in rounded to short tapered at apex; apex aligned or combination with most other colors), tan, yellow, oblique with longitudinal axis of fruit; rounded at base; orange, red, black, cream, gray, green, or purple; with base aligned or oblique with longitudinal axis of fruit; black or brown (greenish, yellowish, reddish) overlay; with the apex and base uniform in texture; chartaceous glabrous; smooth or not smooth; with elevated or or coriaceous; seed chambers externally invisible. Fruit recessed features; rugose (faces and/or margins), margin not constricted; with or without sulcus; plain. shagreen, warty, wrinkled, or papillate; punctate Fruit wings absent. Fruit stipitate to substipitate to (minutely punctate in C. stolzii (E.G. Baker) R.M. nonstipitate; with the stipe 0Ð30 mm long. Fruit with all Polhill); coriaceous. Fracture lines absent. Rim absent. layers dehiscing or indehiscent (for a few species); Wings absent. Raphe not visible. Hilum visible, splitting along sutures. Dehiscence of valves along both partially concealed, or fully concealed; concealed by sutures; apical and down; active; with valves enrolling radicle lobe (partially or completely concealed), or twisting. Replum invisible. Epicarp dull; mono- funiculus, or wing; with or without faboid split; with chrome; brown to tan; glabrous or pubescent and the lips of the faboid split the same color as the rest of indurate; with 1 type of pubescence; puberulent, the hilum; larger than punctiform or punctiform; up to 6 tomentose, villous, sericeous, or peltate; with pubes- mm long; with curved outline; elliptic; between cence brown (and including purple and blackish- cotyledon and radicle lobe; recessed; within rim and brown); with pubescence uniformly distributed or halo or rim or not within corona, halo, or rim. Hilum apical pubescence different from basal pubescence; halo color lighter than testa. Hilum rim color darker with apical 1/4 tomentose and basal 3/4 glabrous; with than testa. Lens discernible (not bearing elevated simple hairs; pliable; with hair bases plain; eglandular; features that rest of testa may bear and often with a without spines; smooth or not smooth; with elevated light-colored longitudinal medial line); equal to or features; not veined; not tuberculate; shagreen; not greater than 0.5 mm in length; 0.5Ð1.5 mm long; with exfoliating; without cracks. Mesocarp thin; surface not margins straight or curved; rectangular, wedge-shaped, veined; 1-layered; without balsamic vesicles; without oblong, or circular; not in groove of raphe; adjacent to fibers; solid; coriaceous to ligneous (sub). Endocarp hilum; 1Ð7 mm from hilum; mounded; similar color as dull; monochrome; brown or tan; smooth or hairy; with or dissimilar color from testa; lighter or darker than hairs restricted to sutures, in longitudinal rows, or testa; tan; not within corona, halo, or rim. Endosperm scattered over endocarp; nonseptate; chartaceous; not thick; covering entire embryo; adnate to testa. Cotyle- exfoliating; remaining fused to mesocarp and epicarp; dons smooth; both outer faces convex; both the same entire. Seeds 10Ð80; length parallel with fruit length; thickness; both more or less of equal length; not folded;

1044 margin entire 180 degrees from base of radicle; similar at apex; not concealing radicle; entire over radicle; without lobes; with the interface division terminating at base of radicle; without margins recessed; yellow or tan; inner face flat; glabrous around base of radicle. Embryonic axis deflexed; oblique to length of seed. Radicle linear or bulbose; lobe tip straight, curved, or hooked; deflexed and parallel to cotyledon length; centered between cotyledons; less than 1/2 or 1/2 to nearly length of cotyledons. Plumule rudimentary, or moderately developed; glabrous.

Distribution: Pansubtropical and pantropical with three- quarters of the species in eastern and southern Africa.

Notes: Polhill (1982) monographed the species of this genus in Africa and Madagascar, where three-quarters of the species are endemic. To balance his excellent contribution, we also consulted the following three monographs from the New World and one from Thailand: North America, Windler (1974); Colombia, Bernal (1986); Venezuela, Matos (1978); and Thailand, Niyomdham (1978). Miller (1967) studied the seed morphology and anatomy of “40 species probably endemic to the New World, 21 species from the Old World, and four pantropicals.” Because of synonomy problems, seeds of only 47 different species are keyed and illustrated. Miller’s key seed characters are: hilum open vs. hilum occluded by radicle, seeds small vs. medium, funicular remnant distinct vs. absent, and glossy vs. not glossy. His term “boss” is synonymous with our term “lens.” Windler (1973) noted that like other legume genera, Crotalaria pods have an active ballistic dehiscence. The two valves separate along the “center part of the adaxial suture,” and an “explosive inward and upward move- ment of the lower suture causes the dispersal of the seeds. The valves continue to curl in the same direction, frequently trapping a few seeds in each of the curled valves.” Windler reported that in the laboratory seeds were thrown up to 5 meters. He also noted that move- ment by water and animals accounted for most seed dispersal. Niyomdham (1978) observed that species of Crotalaria can be divided into two seed groups: Reniform group and cordate (with unequal lobes) or mitiform group. The seeds of Crotalaria exhibit an impressive array of monochrome colors, including, but not limited to, brown (light to dark and in combination with red, gray, yellow, orange, or orange-red), tan, black, orange, red, and yellow. Mottled seeds also occur.

1045 Crotalaria: C. juncea C. Linnaeus (CÐE), C. spp. (AÐB). A, Fruits (ϫ 1.6); B, seeds (ϫ 2.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

1047 Genus: Bolusia G. Bentham cotyledon and radicle lobe (in deep hilar sinus); flush; not within corona, halo, or rim. Lens discernible; less Phylogenetic Number: 27.08. than 0.5 mm in length; with margins straight or curved; apparently oblong; not in groove of raphe; adjacent to Tribe: Crotalarieae. hilum; 0.2 mm from hilum; mounded (well developed and blocking hilar sinus); same color as testa; not Species Studied—Species in Genus: 3 spp.—5 spp. within corona, halo, or rim. Endosperm thin; covering entire embryo; adnate to testa. Cotyledons smooth; both Fruit a legume; unilocular; 2.5Ð4 ϫ 0.5Ð1 ϫ 0.5Ð0.6 cm; outer faces convex; both the same thickness; both more with deciduous calyx; without orifice formed by or less of equal length; not folded; margin entire 180 curving of fruit or fruit segments; straight; not plicate; degrees from base of radicle; similar at apex; partially not twisted; asymmetrical; oblong (fusiform or clavate); concealing (barely covering part of radicle near base) or when asymmetrical with both sutures nearly straight; not concealing radicle; entire over radicle; without inflated (like most Crotalaria (27.07) spp.); com- lobes; with the interface division terminating at base of pressed; without beak; short tapered to rounded at apex; radicle; with 1 or both margins recessed; yellow; inner apex aligned with longitudinal axis of fruit; short face flat; glabrous around base of radicle. Embryonic tapered to rounded at base; base aligned with longitudi- axis deflexed; oblique to length of seed. Radicle linear; nal axis of fruit; with the apex and base uniform in lobe tip hooked; deflexed and parallel to cotyledon texture; ligneous; seed chambers externally invisible. length; centered between cotyledons; 1/2 to nearly Fruit margin not constricted; without sulcus; plain. Fruit length of cotyledons. Plumule moderately developed; wings absent. Fruit nonstipitate. Fruit with all layers glabrous. dehiscing; splitting along sutures. Dehiscence of valves along both sutures; apical and down; active; with valves Distribution: Africa (south of the equator). enrolling or twisting. Replum invisible. Epicarp dull; monochrome; tan; glabrous; eglandular; without spines; not smooth; with elevated features; faintly reticulately veined; not tuberculate; faintly wrinkled; not exfoliating; without or with cracks (with age); cracking oblique to fruit length. Mesocarp thin; surface not veined; 1- layered; without balsamic vesicles; without fibers; solid; subligneous. Endocarp glossy; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 5Ð8 (estimated); length parallel with or transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 1Ð1.5 mm long; of 1 length only; filiform; hooked. Aril absent.

Seed 2.5Ð4 ϫ 2.5Ð3 ϫ 1 mm; not overgrown; not angular; asymmetrical; circular (but with deep hilar sinus); compressed; with visible radicle and cotyledon lobes; with shallow hilar sinus; with umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; streaked and mottled or monochrome (when immature); with frequent mottles; with frequent streaks; tan; with brown (ish gray) overlay; glabrous; not smooth; with elevated features; tuberculate (with tiny white tubercules especially along margin of seed); coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum fully concealed; Bolusia: B. rhodesiana A.G. Corbishley (CÐE), B. spp. (AÐ concealed by radicle lobe (hilum in deep sinus) or B). A, Fruit and valve (ϫ 2.1); B, seeds (ϫ 5.2); CÐD, wing; without faboid split; punctiform; between testa (ϫ 50, ϫ 1000); E, embryos (ϫ 9).

1049 Genus: Lotononis (A.-P. de Candolle) C.F. Ecklon & J.M. cotyledon lobes; without umbo on seed faces. Testa not Zeyher adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome, bichrome (especially Phylogenetic Number: 27.09. yellows and purples or brown with black tubercles), or mottled and streaked; with frequent mottles; with Tribe: Crotalarieae. frequent streaks; red (dish to yellowish), yellow (to greenish), or purple (to tannish); with black or purple Species Studied—Species in Genus: 16 spp.—ca. 150 spp. overlay; glabrous; smooth or not smooth; with elevated features; tuberculate (minute and densely to sparsely); Fruit a legume; unilocular; 0.2Ð1.7 ϫ 0.15Ð0.9 ϫ 0.1 cm; coriaceous. Fracture lines absent. Rim absent. Wings with persistent calyx; with calyx longer or shorter than absent. Raphe not visible. Hilum visible; with faboid fruit; without orifice formed by curving of fruit or fruit split; with the lips of the faboid split the same color as segments; straight or curved; not plicate or plicate the rest of the hilum; punctiform; between cotyledon (rarely); not twisted; asymmetrical or symmetrical; and radicle lobe; recessed; within rim. Hilum rim color oblong, falcate, circular, linear, or ovate; when asym- of or darker than testa. Lens discernible; less than 0.5 metrical with both sutures nearly straight or parallelly mm in length; with margins curved; more or less curved; not inflated; flattened; without beak; tapered to circular; not in groove of raphe; adjacent to hilum; 0.1Ð short tapered at apex; apex oblique with longitudinal 0.4 mm from hilum; mounded; same color as, similar axis of fruit; short tapered at base; base aligned with color as, or dissimilar color from testa; lighter or darker longitudinal axis of fruit; with the apex and base than testa; brown; not within corona, halo, or rim. uniform in texture; coriaceous; seed chambers exter- Endosperm thin; covering entire embryo; adnate to nally invisible. Fruit margin not constricted; without embryo. Cotyledons smooth; both outer faces convex; (lower suture flush) or with sulcus (lower suture deeply both the same thickness; both more or less of equal sulcate); plain or embellished. Fruit wing present or length; not folded; margin entire 180 degrees from base absent; 1; 2 mm wide; sutural; on 1 suture. Fruit of radicle; similar at apex; not concealing radicle; entire stipitate to substipitate to nonstipitate; with the stipe up over radicle; without lobes; with the interface division to 10 mm long. Fruit with all layers dehiscing (to terminating at base of radicle; without margins re- tardily dehiscent) or indehiscent; splitting along sutures. cessed; yellow; inner face flat; glabrous around base of Dehiscence of valves along both sutures; assumed radicle. Embryonic axis deflexed; oblique to length of apical and down; active; with valves twisting. Replum seed. Radicle bulbose; deflexed and parallel to cotyle- invisible. Epicarp dull; monochrome; dark brown to don length; centered between cotyledons; 1/2 to nearly black; pubescent and indurate or pubescent but soon length of cotyledons. Plumule rudimentary; glabrous. deciduous; with 1 type of pubescence; pilose, villous, or sericeous; with pubescence golden or gray; with simple Distribution: Southern Africa (ca. 93 percent of species) to hairs; pliable; with hair bases plain; without spines; Mediterranean region and India. smooth or not smooth; with elevated features; reticulately veined; not tuberculate; distinctly to inconspicu- Notes: Buchenroedera C.F. Ecklon & J.M. Zeyher has been ously verrucose-rugose or warty (along upper suture); merged with Lotononis, following Gunn et al. (1992). not exfoliating; without cracks. Mesocarp thin; surface Polhill (1981q) noted that the 20 species of not veined; 1-layered; without balsamic vesicles; Buchenroedera might be “perhaps better included in without fibers; solid; coriaceous. Endocarp dull; Lotononis.” The seeds of Lotononis are remarkably monochrome; reddish brown; smooth; nonseptate; similar in shape, size, and color to seeds of Trifolium chartaceous; not exfoliating; remaining fused to (21.06). Number of species and distribution follow mesocarp and epicarp; entire. Seeds 1Ð30; length Wyk (1991) and Wyk and Schutte (1995). parallel with fruit length; neither overlapping nor touching or touching; in 1 series. Funiculus measured; up to 2.5 mm long; of 2 different lengths (are some of 1 length?); filiform; straight. Aril absent.

Seed 1Ð3 ϫ 1Ð2.8 ϫ 0.5Ð0.7 mm; not overgrown; not Lotononis: L. bainesii J.G. Baker (CÐE), L. spp. (AÐB). A, angular; asymmetrical; mitaform, reniform, or cordate Fruits and valves (ϫ 2.8); B, seeds (ϫ 8.7); CÐD, testa (obliquely); compressed; with visible radicle and (ϫ 50, ϫ 1000); E, embryos (ϫ 15).

1051 Genus: Lebeckia C.P. Thunberg to endocarp; dull; not modified by a bloom; colored; monochrome or mottled and streaked; with frequent Phylogenetic Number: 27.10. mottles; with frequent streaks; tan; with brown (dark reddish to purplish) overlay; glabrous; smooth or not Tribe: Crotalarieae. smooth; with elevated features; faintly warty; coriaceous. Fracture lines absent. Rim absent. Wings absent. Species Studied—Species in Genus: 14 spp.—ca. 35 spp. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of Fruit a legume; unilocular; 1.2Ð6 ϫ 0.2Ð1.3 ϫ 0.25 cm; the hilum; punctiform; between cotyledon and radicle with persistent or deciduous calyx; with calyx shorter lobe; recessed; within rim, halo, and corona or within than fruit; without orifice formed by curving of fruit or rim. Hilum corona color darker than testa. Hilum halo fruit segments; straight to curved; not plicate; not color of testa. Hilum rim color of or darker than testa. twisted; asymmetrical or symmetrical; linear, oblong, or Lens discernible; equal to or greater than 0.5 mm in elliptic (oblong); when asymmetrical with both sutures length; 0.7Ð1 mm long; with margins straight or curved; parallelly to unequally curved; not inflated to inflated; wedge-shaped (elongated and ending or not in circular flattened to terete; without beak; short tapered to area) or circular (above elongated wedge-shape); not in rounded at apex; apex aligned with longitudinal axis of groove of raphe; adjacent to hilum; mounded; dissimi- fruit; tapered to rounded at base; base aligned with lar color from testa; darker than testa; reddish to longitudinal axis of fruit; with the apex and base orangish to dark brown; not within corona, halo, or rim. uniform in texture; membranous to coriaceous; seed Endosperm thick; covering entire embryo; adnate to chambers externally visible; with the raised seed embryo. Cotyledons smooth; both outer faces convex; chambers torulose or not torulose. Fruit margin not both the same thickness; both more or less of equal constricted; without sulcus; plain or embellished. Fruit length; not folded; margin entire 180 degrees from base wing present or absent; 1; 1Ð2 mm wide; sutural; on 1 of radicle; similar at apex; not concealing radicle; entire suture. Fruit stipitate to substipitate; with the stipe up to over radicle; without lobes; with the interface division 30 mm long. Fruit with all layers dehiscing or indehis- terminating at base of radicle; without margins recent; splitting along sutures. Dehiscence of valves along cessed; tan; inner face flat; glabrous around base of both sutures; apical and down; active; with valves radicle. Embryonic axis deflexed; oblique to length of twisting. Replum invisible. Epicarp dull; monochrome; seed. Radicle bulbose; deflexed and parallel to cotyle- reddish brown or tan; glabrous or pubescent and don width to deflexed and parallel to cotyledon length; indurate; with hairs appressed; with 1 type of pubes- centered between cotyledons; less than 1/2 length of cence; with pubescence gray; pliable; with hair bases cotyledons. Plumule rudimentary or moderately plain; eglandular; without spines; not smooth; with developed; glabrous. elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface Distribution: Namibia, Botswana, and South Africa (Cape not veined; 1-layered; without balsamic vesicles; Province). without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth and cobwebby (smooth Notes: Polhill (1981q) noted that Lebeckia is “virtually below seeds and cobwebby between seeds and in areas indistinguishable” from Spartidium (27.04). Number of without seeds); septate or nonseptate; with septa thin species and distribution taken from Wyk (1991) and (tissue paper-like), flexible; with septa eglandular; Wyk and Schutte (1995). coriaceous to chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð8; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 3Ð4 ϫ 1.7Ð2.2 ϫ 1.2Ð1.5 mm; not overgrown; not angular to angular; asymmetrical; obliquely cordate, oblong (reniform), rectangular, or reniform; com- Lebeckia: L. capensis (C. Linnaeus) G.C. Druce (CÐE), L. pressed; with visible (barely) radicle and cotyledon spp. (AÐB). A, Fruits and valves (ϫ 1.6); B, seeds lobes; without umbo on seed faces. Testa not adhering (ϫ 4.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 9).

1053 Genus: Wiborgia C.P. Thunberg (light); glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Phylogenetic Number: 27.11. Hilum visible, fully concealed, or partially concealed; concealed by funicular remnant or wing; with faboid Tribe: Crotalarieae. split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; between cotyledon Species Studied—Species in Genus: 10 spp.—10 spp. and radicle lobe; recessed; within rim. Hilum rim color of or darker than testa (green). Lens discernible; equal Fruit a legume or nutlet; unilocular; 0.7Ð3.2 ϫ 0.4Ð1.8 ϫ to or greater than 0.5 mm in length; 0.8Ð1.5 mm long; 0.15Ð0.5 cm; with persistent or deciduous calyx; with with margins straight or curved; oblong (with narrow calyx shorter than fruit; without orifice formed by extension or full body reaching to hilum); not in groove curving of fruit or fruit segments; straight; not plicate; of raphe; adjacent to hilum; mounded; dissimilar color not twisted; asymmetrical or symmetrical; circular, from testa; darker than testa; brown; not within corona, elliptic, oblong, or ovate; when asymmetrical with both halo, or rim. Endosperm thin; covering entire embryo; sutures parallelly curved or nearly straight; not inflated adnate to testa. Cotyledons smooth; both outer faces or inflated (W. humilis (C.P.Thunberg) R. Dalhgren); convex; both the same thickness; both more or less of flattened, compressed, or terete (because crested); equal length; not folded; margin entire 180 degrees without beak; short tapered or rounded at apex; apex from base of radicle; similar at apex; not concealing aligned with longitudinal axis of fruit; rounded at base; radicle; entire over radicle; without lobes; with the base aligned or oblique with longitudinal axis of fruit; interface division terminating at base of radicle; without with the apex and base uniform in texture; fragile, margins recessed; yellow or tan; inner face flat; thinner than chartaceous like Trifolium (21.06); seed glabrous around base of radicle. Embryonic axis chambers externally visible; with the raised seed deflexed; oblique to length of seed. Radicle bulbose; chambers not torulose. Fruit margin not constricted; deflexed and parallel to cotyledon length; centered without sulcus; embellished. Fruit wing 1; 0.5Ð5 mm between cotyledons; less than 1/2 length of cotyledons. wide (broadest in W. monoptera E.H.F. Meyer and W. Plumule rudimentary; glabrous. sericea C.P. Thunberg); samaroid or sutural; on 1 suture or both sutures. Fruit stipitate or substipitate; Distribution: South Africa (southern and southwestern Cape with the stipe 5 mm long. Fruit indehiscent. Replum Province). invisible. Epicarp dull or glossy; monochrome (though wings may be a different shade); black or brown (to Notes: Dahlgren (1975) monographed the genus and black-brown and with or without patches of brown); illustrated fruits of the species. He noted that fruit with surface texture uniform or not uniform, with characters are important in “distinguishing Wiborgia patches of different texture not restricted to the base from the similar species of Lebeckia” (27.10). Fruits of and apex; glabrous; eglandular; without spines; not most species are typical samaras. However, the fruit of smooth; with elevated features; reticulately veined; not W. humilis (C.P. Thunberg) R. Dahlgren is a stipitate tuberculate; not exfoliating; without cracks. Mesocarp nutlet with no dorsal wing, and the fruits of W. thick; surface not veined; 1-layered; without balsamic leptoptera R. Dahlgren and W. obcordata (B. Bergius) vesicles; without fibers; solid; subligneous or coria- C.P. Thunberg have a distinct upper ridge reminiscent ceous. Endocarp glossy; monochrome; brown; smooth; of the wing in other species. The wing tissue may or nonseptate; chartaceous; not exfoliating; remaining may not be reticulate, but the fruit tissue over the seed fused to mesocarp and epicarp; entire. Seeds 1Ð3; chamber is always reticulate. length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; up to 1 mm long; of 1 length only; filiform; curved. Aril absent.

Seed 3Ð4 ϫ 2.5Ð2.8 ϫ 1.5Ð1.8 mm; not overgrown; not angular to angular (somewhat); asymmetrical; ovate or rectangular; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not Wiborgia: W. mucronata (C. Linnaeus) A.P. Druce (CÐE), adhering to endocarp; dull; not modified by a bloom; W. spp. (AÐB). A, Fruits (ϫ 2.1); B, seeds (ϫ 5.3); CÐ colored or clear; monochrome; salmon brown or orange D, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 7).

1055 Euchresteae (28.01) embryonic axis right angled. Embryonic axis perpendicular to length of seed; radicle linear; radicle with 90 Genus: Euchresta J.J. Bennett degree turn. Radicle centered between cotyledons; less than one half length of cotyledons. Plumule rudimen- Phylogenetic Number: 28.01. tary; glabrous.

Tribe: Euchresteae. Distribution: Northeastern India to southern Japan to Philippines and Malay Islands. Species Studied—Species in Genus: 2 spp.—4 spp. Notes: Ohashi (1978) reported on the taxonomy and Fruit a legume; unilocular; 1.2Ð2.2 ϫ 0.8Ð1 ϫ 0.5Ð0.6 cm; distribution of E. japonica (J.D. Hooker) E.A. von with deciduous corolla; with deciduous calyx without Regal and concluded that E. trifoliolata E.D. Merrill is orifice formed by curving of fruit or fruit segments; a synonym. Therefore, the number of species is four, straight; not plicate; not twisted; symmetrical; oblong; not five. The testa is fragile and difficult to completely not inflated; compressed; without beak; rounded at separate from the endocarp. The cotyledons are free of apex; apex aligned with longitudinal axis of fruit; the testa and rattle when seeds are shaken. Apparently rounded at base; base aligned with longitudinal axis of seeds germinate while still in the moist fruit. Our seed fruit; with the apex and base uniform in texture; fleshy measurements are based on interior fruit measurements (when fresh), drupaceous, or leathery (when dry); seed because the testae were too fragile to measure. chambers externally invisible. Fruit margin not constricted; without sulcus; plain. Fruit wings absent. Fruit stipitate to substipitate; with the stipe 3Ð12 mm long. Fruit indehiscent. Replum invisible. Epicarp dull or glossy; monochrome; violet-black; glabrous; eglandular; without spines; smooth or not smooth; with elevated features; not veined; not tuberculate; wrinkled; not exfoliating; without cracks. Mesocarp thin; 1- layered; without balsamic vesicles; without fibers; spongy; coriaceous. Endocarp dull; monochrome; tan; cobwebby or spongy; nonseptate; not exfoliating; remaining fused to epicarp; entire. Seed 1; length parallel with fruit length. Funiculus less than 0.5 mm long; of 1 length only; straight. Aril absent.

Seed 12Ð20 ϫ 6Ð8 ϫ 3Ð4 mm; not overgrown; not angular; symmetrical; oblong; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; tan; glabrous; smooth; chartaceous. Pleurogram absent. Fracture lines absent. Rim absent. Raphe from hilum to near base of seed and terminating; not bifurcating; darker than testa; black; raised. Hilum visible; without faboid split; punctiform; subapical to radicle tip; flush; not within corona, halo, or rim. Endosperm absent. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; completely concealing radicle; split over radicle; with Euchresta: E. horsfieldii (J.B.L.T. Lescherault de la Tour) lobes; with the interface division terminating at base of J.J. Bennett (DÐF), E. spp. (AÐC). A, Fruits (ϫ 2.1); B, radicle; without margins recessed; red. Cotyledons open fruit (ϫ 3.7); C, seeds (ϫ 3.7); DÐE, testa (ϫ 50, inner face flat; glabrous around base of radicle; ϫ 1000); F, embryos (ϫ 2).

1057 Thermopsideae (29.01Ð29.06) Hilum partially or fully concealed; concealed by funicular remnant, funiculus, or wing; with or without Genus: Ammopiptanthus S.F. Cheng faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; Phylogenetic Number: 29.01. 0.6 mm long; with curved outline; elliptic; between cotyledon and radicle lobe; flush; not within corona, Tribe: Thermopsideae. halo, or rim. Lens discernible; equal to or greater than or less than 0.5 mm in length; 0.5 mm long; with Species Studied—Species in Genus: 2 spp.—2 spp. margins curved; circular; not in groove of raphe; adjacent to hilum; 0.5 mm from hilum; barely Fruit a legume; unilocular; 4Ð8 ϫ 1.5Ð2 ϫ 0.1Ð0.25 cm; mounded; dissimilar color from testa; darker than testa; with deciduous calyx; without orifice formed by brown; not within corona, halo, or rim. Endosperm thin; curving of fruit or fruit segments; straight or curved covering entire embryo; adnate to testa. Cotyledons (slightly); not plicate; not twisted; asymmetrical or smooth; both outer faces convex; both the same symmetrical; oblong; when asymmetrical with both thickness; both more or less of equal length; not folded; sutures nearly straight; not inflated; flattened; without margin entire 180 degrees from base of radicle; similar or with beak; straight or declined; with solid beak the at apex; partially concealing radicle; notched at radicle; same color and texture as fruit; short tapered at apex; without lobes; with the interface division terminating at apex aligned or oblique with longitudinal axis of fruit; base of radicle; without margins recessed; yellow; inner short tapered at base; base aligned with longitudinal face flat; glabrous around base of radicle. Embryonic axis of fruit; with the apex and base uniform in texture; axis deflexed; oblique to length of seed. Radicle linear; chartaceous; seed chambers externally visible. Fruit lobe tip curved; deflexed and parallel to cotyledon margin not constricted or constricted; slightly con- width; centered between cotyledons; less than 1/2 stricted along both margins; without sulcus; plain. Fruit length of cotyledons. Plumule rudimentary; glabrous. wings absent. Fruit stipitate; with the stipe up to 8 mm long. Fruit with all layers dehiscing; splitting along Distribution: Russia to Mongolia (Inner and Outer) and sutures. Dehiscence of valves along both sutures; apical China. and down; passive. Replum invisible. Epicarp dull; monochrome; tan; glabrous; eglandular; without spines; Notes: Yuan and Peng (1990) reevaluated the 6 genera and not smooth; with elevated features; reticulately veined; 47 species in the tribe, and their data confirms the not tuberculate; not exfoliating; without cracks. information reported by Turner (1981). Turner recog- Mesocarp quite thin; surface not veined; 1-layered; nized 46 species and 6 genera in the tribe. Cheng without balsamic vesicles; without fibers; solid; (1959) monographed the genus and illustrated the fruits chartaceous. Endocarp dull; monochrome; tan; smooth; and seeds of both species. Turner (1981) noted “one or nonseptate; chartaceous; not exfoliating; remaining possibly two species,” and Yuan and Peng (1990) fused to mesocarp and epicarp; entire. Seeds 2Ð5; recognized two species, the number which we used. length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent (but indurate funiculus present).

Seed 5Ð8 ϫ 5Ð7 ϫ 1.5Ð1.6 mm; not overgrown; not angular; asymmetrical; reniform (rounded); compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; modified by a bloom; colored; mottled to streaked or monochrome; with frequent mottles; with frequent streaks; brown, tan (greenish), or yellow; with black overlay; glabrous; smooth; coriaceous. Fracture Ammopiptanthus: A. mongolicus (C.J. Maximowicz) S.-H. lines absent. Rim absent. Wings absent. Raphe from Cheng (CÐE), A. spp. (AÐB). A, Fruits (ϫ 1.2); B, seeds hilum through lens and terminating before base of seed; (ϫ 7.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 3). not bifurcating; darker than testa; black; slightly raised.

1059 Genus: Piptanthus R. Sweet overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe from hilum Phylogenetic Number: 29.02. through lens and terminating before base of seed; not bifurcating; darker than testa; brown; flush. Hilum Tribe: Thermopsideae. partially concealed; concealed by funicular remnant; with faboid split; with the lips of the faboid split the Species Studied—Species in Genus: 2 spp.—2–3 spp. same color as the rest of the hilum; larger than punctiform or punctiform; 0.3Ð0.7 mm long; with curved Fruit a legume; unilocular; 7Ð9 ϫ 1Ð1.3 ϫ 0.18Ð0.2 cm; outline; circular; between cotyledon and radicle lobe; with persistent or deciduous calyx; with calyx shorter recessed; within rim. Hilum rim color of or darker than than fruit; without orifice formed by curving of fruit or testa. Lens discernible; less than or equal to or greater fruit segments; straight to curved (to slightly curved); than 0.5 mm in length; 0.5Ð1 mm long; with margins not plicate; not twisted; asymmetrical or symmetrical; curved; circular; not in groove of raphe; adjacent to linear; when asymmetrical with both sutures nearly hilum; 1.3Ð1.5 mm from hilum; mounded; dissimilar straight; not inflated; flattened; without beak; tapered or color from testa; darker than testa; dark brown; not short tapered at apex; apex aligned or oblique with within corona, halo, or rim. Endosperm thin; covering longitudinal axis of fruit; short tapered at base; base entire embryo; adnate to testa (and a sheath around aligned with longitudinal axis of fruit; with the apex radicle). Cotyledons smooth; both outer faces convex; and base uniform in texture; coriaceous; seed chambers both the same thickness; both more or less of equal externally visible. Fruit margin constricted or not length; not folded; margin entire 180 degrees from base constricted; constricted along both margins (not of radicle; similar at apex; not concealing radicle; entire regularly); without sulcus; plain. Fruit wings absent. over radicle; without lobes; with the interface division Fruit stipitate or substipitate; with the stipe 5Ð20 mm terminating at base of radicle; without margins re- long. Fruit with all layers dehiscing or indehiscent; cessed; tan; inner face flat; glabrous around base of splitting along sutures. Dehiscence of valves passive. radicle. Embryonic axis deflexed; oblique to length of Replum invisible. Epicarp dull; monochrome; brown or seed. Radicle bulbose; deflexed and parallel to cotyle- tan (and brown over seeds); pubescent and indurate; don length or width; centered between cotyledons; less with hairs appressed or erect; with 1 type of pubes- than 1/2 length of cotyledons. Plumule moderately cence; tomentose; with pubescence golden (P. developed; glabrous. tomentosa A.R. Franchet) or gray (P. nepalinsis); with pubescence uniformly distributed; with simple hairs; Distribution: India, Nepal, and China. pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; reticulately Notes: Turner (1980), who monographed the genus, veined; not tuberculate; not exfoliating; without cracks. recognized two species, but later, he noted the species Mesocarp quite thin; surface not veined; 1-layered; number as two or three (Turner 1981). We noticed that without balsamic vesicles; without fibers; solid; the area under the hilum is thick and spongy. chartaceous. Endocarp dull; monochrome; dark brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; separating into 1-seeded winged segments. Seeds 2Ð10; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 3 mm long; of 1 length only; thick; straight. Aril absent.

Seed 5Ð7 ϫ 4.3Ð5 ϫ 1.5Ð3.5 mm; not overgrown; not angular; asymmetrical; oblong (with radicle lobe); compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled and streaked; with frequent Piptanthus: P. nepalensis (W.J. Hooker) D. Don (BÐE), P. mottles; with frequent streaks; greenish or reddish spp. (A). A, Fruits (ϫ 1.3); B, seeds (ϫ 5.7); CÐD, testa brown or tan (reddish); with reddish brown or green (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

1061 Genus: Anagyris C. Linnaeus mm in length; 1 mm long; with margins curved; circular; not in groove of raphe; adjacent to hilum; 2 Phylogenetic Number: 29.03. mm from hilum; barely mounded; dissimilar color from testa; darker than testa; reddish brown; not within Tribe: Thermopsideae. corona, halo, or rim. Endosperm thick; covering entire embryo; adnate to testa. Cotyledons smooth; both outer Species Studied—Species in Genus: 2 spp.—1 or 2 spp. faces convex; both the same thickness; both more or less of equal length; not folded; margin entire 180 Fruit a legume; unilocular; up to 13 ϫ 1.5Ð2 ϫ 0.3Ð1.3 cm; degrees from base of radicle; similar at apex; not with deciduous calyx; without orifice formed by concealing radicle; entire over radicle; without lobes; curving of fruit or fruit segments; straight or curved with the interface division terminating at base of (slightly); not plicate; not twisted; symmetrical; broadly radicle; with 1 or both margins recessed; yellow; inner linear; not inflated; flattened or compressed; without or face flat; glabrous around base of radicle. Embryonic with beak; straight; with solid beak the same color and axis deflexed; oblique to length of seed. Radicle linear; texture as fruit; tapered or rounded at apex; apex lobe tip curved; deflexed and parallel to cotyledon aligned with longitudinal axis of fruit (or nearly so); length; centered between cotyledons; less than 1/2 long tapered or rounded at base; base aligned with length of cotyledons. Plumule moderately developed; longitudinal axis of fruit (or nearly so); with the apex glabrous. and base uniform in texture; coriaceous or ligneous (when dry); seed chambers externally visible; with the Distribution: Mediterranean Europe, Asia Minor, northern raised seed chambers not torulose. Fruit margin not Africa, and Canary Islands. constricted; without sulcus; plain. Fruit wings absent. Fruit stipitate; with the stipe up to 10 mm long. Fruit Notes: We are still unsure whether one or two species exist indehiscent. Replum invisible. Epicarp dull; mono- in Anagyris. The other possible species (besides A. chrome; brown; glabrous; eglandular; without spines; foetida) is A. latifolia P.M.A. Broussonet ex C.L. von smooth; not veined; not tuberculate; not exfoliating; Willdenow (Hansen and Sundling 1993). without cracks. Mesocarp thick or thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; spongy or solid; coriaceous or ligneous. En- docarp dull; monochrome; brown; smooth; septate; with septa thin (tissue paper-like), flexible; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 5; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; straight. Aril absent.

Seed 10Ð15 ϫ 7Ð10 ϫ 5Ð6.7 mm; not overgrown; not angular; asymmetrical; elliptic to oblong; compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; monochrome; dark to light brown or tan; glabrous; smooth; osseous. Fracture lines absent. Rim absent. Wings absent. Raphe from hilum through lens and terminating before base of seed; not bifurcating; darker than testa; dark reddish brown; flush. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 2 mm long; with curved or straight outline; elliptic or oblong; marginal accord- Anagyris: A. foetida C. Linnaeus (AÐE). A, Fruits (ϫ 1.1); ing to radicle tip; recessed; within rim. Hilum rim color B, seeds (ϫ 2.5); CÐD, testa (ϫ 50, ϫ 1000); E, of testa. Lens discernible; equal to or greater than 0.5 embryos (ϫ 2).

1063 Genus: Thermopsis R. Brown Seed 3.2Ð6 ϫ 2Ð4 ϫ 1.5Ð2.8 mm; not overgrown; not angular; asymmetrical; reniform (or nearly so); com- Phylogenetic Number: 29.04. pressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to Tribe: Thermopsideae. endocarp; dull or glossy; not modified or modified by a bloom; colored; monochrome or bichrome; greenish or Species Studied—Species in Genus: 17 spp.—23 spp. reddish black, black, brown (yellow, orange, or brown), purple, or orange; glabrous; not smooth or smooth; with Fruit a legume or loment (or a loment segment, or at least elevated features; blistered cuticle (exudate); coria- lomentaceous); unilocular; 3Ð10 ϫ 0.8Ð1 ϫ 0.15Ð0.3 ceous. Fracture lines absent. Rim absent. Wings absent. cm; with persistent calyx; with calyx shorter than fruit; Raphe not visible or visible; from hilum through lens without orifice formed by curving of fruit or fruit and terminating before base of seed; not bifurcating; segments; straight, curved (or slightly curved), or 0.5- color of or darker than testa; brown; raised. Hilum coiled; not plicate; not twisted; asymmetrical or partially concealed; concealed by funicular remnant or symmetrical; linear, oblong, ovate, or elliptic; when wing; with faboid split; with the lips of the faboid split asymmetrical with both sutures parallelly curved or the same color as the rest of the hilum; punctiform; nearly straight; not inflated or inflated (T. inflata J. between cotyledon and radicle lobe; recessed; within Cambessedes); flattened or compressed; without beak; rim. Hilum rim color of testa. Lens discernible or not short tapered or rounded at apex; apex aligned with discernible; less than 0.5 mm in length; with margins longitudinal axis of fruit; short tapered at base; base curved; circular; not in groove of raphe; adjacent to aligned or right angled with longitudinal axis of fruit; hilum; 0.5 mm from hilum; mounded; same color or with the apex and base uniform in texture; coriaceous; similar color as testa; darker than testa; black or brown; seed chambers externally visible; with the raised seed not within corona, halo, or rim. Endosperm thin; chambers not torulose. Fruit margin not constricted; covering entire embryo; adnate to testa or embryo. without sulcus; plain. Fruit wings absent. Fruit Cotyledons smooth; both outer faces convex; both the substipitate or nonstipitate. Fruit with all layers same thickness; 1 longer than other; not folded; margin dehiscing; splitting along sutures. Dehiscence of valves entire 180 degrees from base of radicle; similar at apex; along both sutures; apical and down; passive. Replum completely concealing radicle; entire over radicle; invisible. Loment indehiscent. Loment segments widest without lobes; with the interface division terminating at across seed area; rectangular. Epicarp dull; mono- base of radicle; without margins recessed; tan; inner chrome; brown; glabrous or pubescent and indurate; face flat; glabrous around base of radicle. Embryonic with hairs erect or appressed; with 1 type of pubes- axis deflexed; oblique to length of seed. Radicle linear; cence; puberulent or villous; with pubescence gray; lobe tip straight; deflexed and parallel to cotyledon with pubescence uniformly distributed; with simple width; not centered between cotyledons (radicle outside hairs; pliable; with hair bases plain; eglandular; without 1 cotyledon and inside other, therefore junctions for spines; not smooth; with elevated features; reticulately each cotyledon different); less than 1/2 length of veined; not tuberculate; not exfoliating; without cracks. cotyledons. Plumule moderately developed; glabrous. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Distribution: North America (10 spp.) and Asia (13 spp.). Endocarp dull; monochrome; brown; smooth; nonseptate or septate; with septa thicker than paper, Notes: Larisey (1940b), Isley (1981), and Chen et al. (1994) firm; with septa eglandular; chartaceous; not exfoliat- monographed the North American species, and Barneby ing; remaining fused to mesocarp and epicarp; entire. (1989) treated the species for the intermountain region Seeds 2Ð11; length transverse to fruit length; neither of the western United States. Larisey and Chen et al. overlapping nor touching; in 1 series. Funiculus less used narrow species concepts, and recognized 10 than 0.5 mm long; of 1 length only; thick; straight or species in North America—the number we used for our hooked. Aril dry; when dry rim-aril, or partial rim-aril; species count. Isley and Barneby employed much laciniate, or entire; covering less than 1/2 of seed; broader species concepts. Isley accepted only four without tongue (or flap-like) on lips of 2-lipped rim- species in North America, and Barneby just three. Most aril; white. species have a dehiscent legume, but a few species have lomentaceous fruits and these should be studied. Species producing the latter include T. divaricarpa A.

1064 Nelson, T. gracilis T.J. Howell, and T. rhombifolia (T. Nuttall ex F.T. Pursh) J. Richardson.

1065 Thermopsis: T. lupinoides (C. Linnaeus) J.H.F. Link (CÐE), T. spp. (AÐB). A, Fruits (ϫ 1.4); B, seeds (ϫ 6.6); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

1067 Genus: Baptisia É.P. Ventenat Seed 2.5Ð4.5 ϫ 2Ð3 ϫ 1.6Ð2 mm; not overgrown; not angular; asymmetrical; D-shaped (with inconspicuous Phylogenetic Number: 29.05. radicle lobe); compressed; with visible radicle and cotyledon lobes; without umbo on seed faces. Testa not Tribe: Thermopsideae. adhering to endocarp; dull or glossy; modified by a bloom; colored; monochrome; yellowish to dark brown, Species Studied—Species in Genus: 7 spp.—17 spp. tan, or white; glabrous; smooth or not smooth; with elevated features; tuberculate (glandular exudates) or Fruit a legume; unilocular; 1Ð7 ϫ 0.2Ð2.5 ϫ 0.7Ð2 cm; with blistered cuticle (B. megacarpa A.W. Chapman ex J. persistent or deciduous calyx; with calyx shorter than or Torrey & A. Gray); coriaceous. Fracture lines absent. equal in length to fruit; without orifice formed by Rim absent. Wings absent. Raphe from lens to base of curving of fruit or fruit segments; straight or curved seed and terminating (near base); not bifurcating; (slightly); not plicate; not twisted; symmetrical or darker than testa; black; slightly recessed or flush. asymmetrical; oblong, ovate, circular, or elliptic; when Hilum partially concealed; concealed by funicular asymmetrical with both sutures parallelly curved or 1 remnant; with faboid split; with the lips of the faboid straight and 1 curved suture; widest near middle or D- split the same color as the rest of the hilum; punctiform shaped; usually inflated or not inflated; terete; without or larger than punctiform; 0.3Ð0.4 mm long; with or with beak; straight; with solid beak the same color curved outline; circular; between cotyledon and radicle and texture as fruit; rounded or long tapered at apex; lobe; recessed; within rim or halo (occasionally and apex aligned or oblique with longitudinal axis of fruit; faintly). Hilum halo color darker than testa. Hilum rim long tapered or rounded at base; base aligned with color of testa. Lens discernible or not discernible; less longitudinal axis of fruit; with the apex and base than 0.5 mm in length; with margins curved; circular; uniform in texture; ligneous, coriaceous, or membra- not in groove of raphe; adjacent to hilum; 2 mm from nous (B. megacarpa A.W. Chapman ex J. Torrey & A. hilum; flush; dissimilar color from testa; darker than Gray); seed chambers externally invisible. Fruit margin testa; reddish brown; not within corona, halo, or rim. not constricted; without or with sulcus; plain. Fruit Endosperm thin; covering entire embryo; adnate to wings absent. Fruit stipitate or substipitate; with the testa. Cotyledons smooth; both outer faces convex; both stipe 4Ð13 mm long. Fruit with all layers dehiscing; the same thickness; both more or less of equal length; splitting along sutures. Dehiscence of valves along both not folded; margin entire 180 degrees from base of sutures; apical and down; passive. Replum invisible. radicle; similar at apex; not concealing radicle; entire Epicarp dull or semiglossy; monochrome; dark to over radicle; without lobes; with the interface division yellowish to reddish to purplish brown or black terminating at base of radicle; without margins re- (brownish); glabrous or pubescent and indurate; with cessed; yellow or tan; inner face flat; glabrous around hairs appressed (sub) or erect; with 1 type of pubes- base of radicle. Embryonic axis deflexed; oblique to cence; villous; with pubescence gray; with pubescence length of seed. Radicle linear; lobe tip curved; deflexed uniformly distributed; with simple hairs; pliable; with and parallel to cotyledon width; centered between hair bases plain; eglandular; without spines; smooth or cotyledons; less than 1/2 length of cotyledons. Plumule not smooth; with elevated features; reticulately veined; rudimentary; glabrous. not tuberculate; wrinkled or rugose; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1- Distribution: Eastern Canada, United States, and Mexico. layered; without balsamic vesicles; without fibers; solid; subligneous, coriaceous, or chartaceous. En- Notes: Larisey (1940a) and Isley (1981) monographed docarp dull; monochrome; dark to light brown; smooth; Baptisia. Occasionally, three cotyledons are found in nonseptate; chartaceous; not exfoliating; remaining the seed. fused to mesocarp and epicarp; entire. Seeds 20Ð30 (estimated); length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 3.5 mm long; of 1 length only; thick; straight. Aril absent or present (really funicular remnant); dry; rim-aril; brown. Baptisia: B. alba (C. Linnaeus) É.P. Ventenat (CÐE), B. spp. (AÐB). A, Fruits (ϫ 1.2); B, seeds (ϫ 7.2); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 12).

1069 Genus: Pickeringia T. Nuttall ex J. Torrey & A. Gray raised. Hilum partially or fully concealed; concealed by funicular remnant; with faboid split; with the lips of the Phylogenetic Number: 29.06. faboid split the same color as the rest of the hilum; punctiform; marginal according to radicle tip; recessed; Tribe: Thermopsideae. within rim. Hilum rim color of testa. Lens discernible; less than 0.5 mm in length; with margins curved; Species Studied—Species in Genus: 1 sp.—1 sp. circular; not in groove of raphe; adjacent to hilum; 0.5 mm from hilum; mounded; same color as testa; not Fruit a legume; unilocular; 2Ð5 ϫ 0.4Ð0.5 ϫ 0.2 cm; with within corona, halo, or rim. Endosperm thick; covering persistent calyx; with calyx shorter than fruit; without entire embryo; adnate to embryo. Cotyledons smooth; orifice formed by curving of fruit or fruit segments; both outer faces convex; both the same thickness; both slightly curved; not plicate; not twisted; symmetrical; more or less of equal length; not folded; margin entire linear; not inflated; compressed; with beak; straight; 180 degrees from base of radicle; similar at apex; not with solid beak the same color and texture as fruit; long concealing radicle; entire over radicle; without lobes; tapered at apex; apex aligned or oblique with longitudi- with the interface division terminating at base of nal axis of fruit; long tapered at base; base aligned with radicle; without margins recessed; tan; inner face flat; longitudinal axis of fruit; with the apex and base glabrous around base of radicle. Embryonic axis uniform in texture; coriaceous; seed chambers exter- deflexed; parallel to length of seed; without a joint nally visible or invisible; with the raised seed chambers evident between the radicle and the cotyledons. Radicle not torulose. Fruit margin constricted along both differentiated from cotyledon; linear; deflexed and margins (by abortion of ovules); without sulcus; plain. parallel to cotyledon length; centered between cotyle- Fruit wings absent. Fruit stipitate; with the stipe 5Ð6 dons; less than 1/2 length of cotyledons. Plumule mm long. Fruit with all layers dehiscing; splitting along rudimentary; glabrous. suture. Dehiscence of valves along 1 suture; assumed apical and down; passive. Replum invisible. Epicarp Distribution: United States (north-central California) to dull; monochrome; light brown; pubescent and indu- Mexico (northern Baja California). rate; with hairs appressed; with 1 type of pubescence; with pubescence gray; with pubescence uniformly Notes: Isley (1981) revised the genus. Pickeringia montana distributed; with simple hairs; pliable; with hair bases rarely sets fruits and seeds, hence the lack of material plain; eglandular; without spines; not smooth; with for our plate. Some authors, including Isley but not elevated features; reticulately veined; not tuberculate; Rudd (1968), recognized var. tomentosa, which sets not exfoliating; without cracks. Mesocarp present (other more fruits and seeds. Variety tomentosa cannot be mesocarp characters are assumed); thin; surface not separated from var. montana by the use of morphologi- veined; 1-layered; without balsamic vesicles; without cal characters. fibers; solid; chartaceous. Endocarp dull; monochrome; tan; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð10; length parallel with fruit length; neither overlapping nor touching or touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 3Ð4.2 ϫ 2Ð2.5 ϫ 1Ð1.5 mm; not overgrown; not angular; asymmetrical; rectangular or reniform; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; modified by a bloom; colored; monochrome; brownish black; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings Pickeringia: P. montana T. Nuttall (A), P. montana T. absent. Raphe from hilum through lens and terminating Nuttall var. tomentosa (L.R. Abrams) I.M. Johnston before base of seed; not bifurcating; color of testa; (CÐE), P. spp. (B). A, Fruit (ϫ 3.7); B, seeds (ϫ 11.3); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

1071 Genisteae (30.01Ð30.25) absent. Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by aril; with Genus: Melolobium C.F. Ecklon & J.M. Zeyher faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; between Phylogenetic Number: 30.01. cotyledon and radicle lobe; recessed; within rim. Hilum rim color of, lighter than, or darker than testa. Lens Tribe: Genisteae. discernible; less than 0.5 mm in length; with margins straight or curved; irregular; irregular or key-hole Species Studied—Species in Genus: 8 spp.—ca. 20 spp. shaped; not in groove of raphe; adjacent to hilum; up to 0.2 mm from hilum; mounded; dissimilar color from Fruit a legume; unilocular; 0.8Ð2 ϫ 0.3Ð0.5 ϫ 0.08Ð0.2 cm; testa; darker than testa; black (at least center); not with deciduous calyx; without orifice formed by within corona, halo, or rim. Endosperm thin; covering curving of fruit or fruit segments; straight or curved; entire embryo; adnate to embryo. Cotyledons smooth; not plicate; not twisted; asymmetrical or symmetrical; both outer faces convex; both the same thickness; both linear or oblong (narrowly) to ovate; when asymmetri- more or less of equal length; not folded; margin entire cal with both sutures nearly straight; not inflated; 180 degrees from base of radicle; similar at apex; not compressed; without beak; short tapered at apex; apex concealing radicle; entire over radicle; without lobes; aligned with longitudinal axis of fruit; short tapered at with the interface division terminating at base of base; base aligned with longitudinal axis of fruit; with radicle; without margins recessed; yellow or green; the apex and base uniform in texture; coriaceous; seed inner face flat; glabrous around base of radicle. Embry- chambers externally visible; with the raised seed onic axis deflexed; oblique to length of seed. Radicle chambers torulose. Fruit margin slightly constricted linear; deflexed and parallel to cotyledon length; along both margins; without sulcus; plain. Fruit wings centered between cotyledons; 1/2 to nearly length of absent. Fruit substipitate. Fruit with all layers dehisc- cotyledons. Plumule rudimentary; glabrous. ing; splitting along sutures. Dehiscence of valves along both sutures; apical and down; active; with valves Distribution: Southern Africa. slightly twisting. Replum invisible. Epicarp dull; monochrome or multicolored; mottled; brown or tan; Notes: Traditionally this tribe has been called Genisteae. with brown (reddish) overlay; with mottling over seed Reveal (1997) reported that the name Cytiseae was chambers; pubescent and indurate; with 1 type of published before the name Genisteae. In accordance pubescence; tomentose; with pubescence golden or with the International Code of Botanical Nomenclature gray; with simple hairs; pliable; with hair bases plain; (Greuter et al. 1994), the oldest name for a taxon must eglandular; without spines; not smooth; with elevated be used, so Reveal suggested that this tribe should be features; not veined; not tuberculate; warty; not called Cytiseae. In 1999, however, Reveal (1999) exfoliating; without cracks. Mesocarp thin; surface not reversed himself, so this tribe remains the Genisteae. veined; 1-layered; without balsamic vesicles; without Bisby (1981) summarized the tribe Genisteae, follow- fibers; solid; coriaceous. Endocarp dull; monochrome; ing the excellent study by Polhill (1976). Bisby reddish brown; smooth; subseptate; with septa thin correctly noted that “many species have been moved (tissue paper-like), flexible; with septa eglandular; from one genus to another several times and the chartaceous; not exfoliating; remaining fused to Cytisus-Genista complex has gained a reputation as a mesocarp and epicarp; entire. Seeds 1Ð8; length parallel critical group.” He continued with an in-depth analysis with fruit length; neither overlapping nor touching; in 1 of the tribe. Using chemical and morphological series. Funiculus less than 0.5 mm long; of 1 length characters, Polhill (1994a,b) and Wyk and Schutte only; flattened; straight. Aril present (so tiny that it can (1995) transferred the following five genera from be missed) or absent; dry; tiny tongue-aril; white. Crotalarieae (27) to Genisteae: Anarthrophyllum (30.06), Argyrolobium (30.03), Dichilus (30.02), Seed 2Ð2.5 ϫ 2Ð2.5 ϫ 1.5 mm; not overgrown; not angular; Melolobium, and Sellocharis (30.07). Cristofolini asymmetrical; ovate to circular (sub); compressed; (1997) carried out a cladistic study of the tribe’s without visible radicle and cotyledon lobes; without biogeography and discussed its early evolutionary umbo on seed faces. Testa not adhering to endocarp; history. Polhill (1981q) and Wyk (1991) had Melobium dull; not modified by a bloom; colored; monochrome or in Crotalarieae but later transferred it to the beginning mottled; with frequent mottles; reddish to dark reddish of the Genisteae (Polhill 1994a,b, Wyk and Schutte brown; glabrous; smooth; coriaceous. Fracture lines 1995). 1072 1073 Melolobium: M. decumbens (E.H.F. Meyer) J. Burtt Davy (CÐE), M. spp. (AÐB). A, Fruits (ϫ 2.2); B, seeds (ϫ 7.5); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

1075 Genus: Dichilus A.-P. de Candolle green (restricted to green seeds), or yellow (restricted to green seeds) overlay; glabrous; smooth; coriaceous. Phylogenetic Number: 30.02. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by Tribe: Genisteae. funicular remnant; without faboid split; larger than punctiform; 0.5 mm long; with curved outline; circular; Species Studied—Species in Genus: 5 spp.—5 spp. marginal according to radicle tip or between cotyledon and radicle lobe; flush; within rim or not within corona, Fruit a legume; unilocular; 0.25Ð0.55 ϫ 0.4 ϫ 0.14Ð0.15 halo, or rim. Hilum rim color lighter than testa. Lens cm; with persistent calyx; with calyx shorter than fruit; discernible; equal to or greater than 0.5 mm in length; without orifice formed by curving of fruit or fruit 0.6Ð0.9 mm long; with margins straight or curved; segments; straight to curved (slightly); not plicate; not wedge-shaped (to hourglass-shaped and surrounded by twisted; asymmetrical or symmetrical; oblong to linear tan circular patch, when circular tan with reddish-brown (or narrowly so) or oblong (narrowly); when asym- and hourglass-shaped to wedge-shaped interior) or metrical with both sutures nearly straight; not inflated; circular; not in groove of raphe; adjacent to hilum; 0.1 compressed; without beak; short tapered at apex; apex mm from hilum; mounded; dissimilar color from testa; oblique with longitudinal axis of fruit; tapered at base; lighter than testa; tan and brown (reddish); not within base aligned with longitudinal axis of fruit; with the corona, halo, or rim. Endosperm thick; covering entire apex and base uniform in texture; coriaceous; seed embryo; adnate to embryo. Cotyledons not smooth; chambers externally visible; with the raised seed both outer faces convex; both the same thickness; both chambers not torulose. Fruit margin slightly constricted more or less of equal length; not folded; margin entire along both margins; without sulcus; plain. Fruit wings 180 degrees from base of radicle; similar at apex; absent. Fruit substipitate. Fruit with all layers dehisc- completely concealing radicle; entire over radicle; ing; splitting along sutures. Dehiscence of valves along without lobes; with the interface division terminating at both sutures; apical and down; active; with valves base of radicle; without margins recessed; yellow; inner twisting. Replum invisible. Epicarp dull; monochrome; face flat; glabrous around base of radicle. Embryonic brown; glabrous, pubescent and indurate, or pubescent axis deflexed; oblique to length of seed. Radicle but soon deciduous; with 1 type of pubescence; bulbose; deflexed and parallel to cotyledon length; puberulent (especially along sutures); with pubescence centered between cotyledons; 1/2 to nearly length of gray; with simple hairs; pliable; with hair bases plain; cotyledons. Plumule rudimentary or moderately without spines; not smooth; with elevated features; developed; glabrous. reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1- Distribution: Southern Africa. layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; brown; Notes: Schutte and Wyk (1988) monographed the genus and spongy; septate; with septa thin (tissue paper-like), illustrated its seeds and fruits. They also analyzed the flexible; with septa eglandular; chartaceous; not taxonomic relationships of the species (Schutte and exfoliating; remaining fused to mesocarp and epicarp; Wyk 1990). Polhill (1981q) and Wyk (1991) placed this entire. Seeds 2Ð9; length parallel with fruit length; genus in Crotalarieae but later transferred it to the neither overlapping nor touching; in 1 series. Funiculus beginning of the Genisteae (30) (Polhill 1994a,b, Wyk less than 0.5 mm long; of 1 length only; thick; straight. and Schutte 1995). Aril absent.

Seed 2.5Ð3 ϫ 2Ð2.8 ϫ 1.5Ð1.7 mm; not overgrown; not angular; asymmetrical; circular or oblong; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled and streaked; with frequent or infrequent mottles; with frequent or infrequent streaks; black, Dichilus: D. lebeckioides A.-P. de Candolle (CÐE), D. spp. brown (pale yellow to brown), green (dark to olive), or (AÐB). A, Fruits (ϫ 1.8); B, seeds (ϫ 6.8); CÐD, testa orange; with black (restricted to black seeds), brown, (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

1077 Genus: Argyrolobium C.F. Ecklon & J.M. Zeyher Seed 1.4Ð4 ϫ 1.2Ð2.8 ϫ 1Ð1.3 mm; not overgrown; not angular or angular; asymmetrical; oblong, ovate, Phylogenetic Number: 30.03. quadrangular, reniform, triangular, circular, or mitaform; compressed or terete; with surface smooth; Tribe: Genisteae. with or without visible radicle and cotyledon lobes; with or without external groove between radicle and Species Studied—Species in Genus: 19 spp.—ca. 70 spp. cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar Fruit a legume; unilocular; 0.6Ð8.5 ϫ 0.3Ð0.6 ϫ 0.13Ð0.25 sinus; without umbo on seed faces. Testa not adhering cm; with persistent or deciduous calyx; with calyx to endocarp; glossy or dull; not modified by a bloom; shorter than fruit; without orifice formed by curving of colored; monochrome or mottled and streaked; with fruit or fruit segments; straight or curved (or slightly frequent or infrequent mottles; with frequent streaks; curved); not plicate; not twisted; asymmetrical or reddish to greenish brown, tan (to greenish), green, symmetrical; linear, oblong, ovate, lanceolate, or C- olive, yellow, or black; with brown (dark), purple shaped; when asymmetrical with both sutures nearly (definite marks to “clouds”), or black overlay; glabrous; straight or parallelly curved; not inflated; compressed smooth or not smooth; with elevated features; reticulate or flattened; with or without beak; with solid beak the or wrinkled; coriaceous. Fracture lines absent. Rim same color and texture as fruit; tapered or short tapered absent. Wings absent. Raphe not visible. Hilum visible; at apex; apex aligned or right-angled with longitudinal with faboid split; with the lips of the faboid split the axis of fruit; short tapered at base; base aligned or same color as the rest of the hilum; punctiform or larger oblique with longitudinal axis of fruit; with the apex than punctiform; up to 0.5 mm long; with curved and base uniform in texture; membranous or coria- outline; circular; between cotyledon and radicle lobe, ceous; seed chambers externally visible; with the raised apical at apex of radicle tip, subapical to radicle tip, or seed chambers torulose or not torulose. Fruit margin marginal according to radicle tip; recessed; within rim. not constricted; without sulcus; plain. Fruit wings Hilum rim color of or lighter than testa. Lens discern- absent. Fruit nonstipitate. Fruit with all layers dehisc- ible or not discernible; less than or equal to or greater ing; splitting along sutures. Dehiscence of valves along than 0.5 mm in length; up to 0.5 mm long; with margins both sutures; apical and down; active; with valves straight or curved; triangular or oblong; circular or twisting. Replum invisible. Epicarp dull; monochrome; oblong; not in groove of raphe; adjacent to hilum; 0.2Ð brown or tan; pubescent and indurate or glabrous; with 0.5 mm from hilum; flush or mounded; dissimilar color hairs erect or appressed; with 1 or 2 types of pubes- from or similar color as testa; darker or lighter than cence; pilose, villous, or sericeous; with pubescence testa; brown or tan; not within corona, halo, or rim. gray or golden; with gray hairs on valves and golden Endosperm thick; covering entire embryo; adnate to hairs on sutures; with pubescence uniformly distrib- embryo. Cotyledons not smooth; both outer faces uted; with simple hairs; pliable; with hair bases plain; convex; both the same thickness; both more or less of eglandular; without spines; smooth or not smooth; with equal length; not folded; margin entire 180 degrees elevated features; not veined; not tuberculate; faintly from base of radicle; similar at apex; not concealing wrinkled; not exfoliating; without cracks. Mesocarp radicle; split over radicle; with lobes; with the interface thin; surface not veined; 1-layered; without balsamic division terminating at base of radicle; without margins vesicles; without fibers; solid; coriaceous. Endocarp recessed; tan, yellow, or white; inner face flat; glabrous dull; monochrome; brown or tan; smooth and scurfy; around base of radicle. Embryonic axis deflexed; nonseptate (with slight to well developed scurfy lines oblique to length of seed; without a joint evident between seeds) or subseptate (because of regular between the radicle and the cotyledons. Radicle intrusion of fruit); with septa eglandular; chartaceous; bulbose; lobe tip straight; deflexed and parallel to not exfoliating; remaining fused to mesocarp and cotyledon length; centered between cotyledons; 1/2 to epicarp; entire. Seeds 1Ð14; length oblique to fruit nearly or equaling length of cotyledons. Plumule length; neither overlapping nor touching; in 1 series. rudimentary; glabrous. Funiculus measured or less than 0.5 mm long; up to 1.5 mm long; of 1 length only; flattened or thick; straight Distribution: Mediterranean region to India, highlands or curved. Aril dry; rim-aril or tongue-aril; tan, white, tropical Africa, South Africa, and Madagascar (1 sp.). or black.

1078 Notes: Polhill (1968) monographed the taxa of tropical Africa, Chaudhary (1997) revised the Indian taxa, and Edwards (1997) revised the species of series Racemosae sensu W.H. Harvey (Harvey 1962). Wyk and Schutte (1990) compared Argyrolobium with several Crotalarieae genera of South Africa and concluded that Argyrolobium should be part of Crotalarieae, but they never made the change. Polhill noted that the endocarp often breaks down to form internal partitions.

1079 Argyrolobium: A. biebersteinii P.W. Ball (CÐE), A. spp. (AÐ B). A, Fruits (closed and dehisced) (ϫ 1.2); B, seeds (ϫ 4.7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

1081 Genus: Polhillia C.H. Stirton split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; between cotyledon Phylogenetic Number: 30.04. and radicle lobe; recessed; within rim. Hilum rim color darker than testa. Lens discernible; equal to or greater Tribe: Genisteae. than 0.5 mm in length; 0.7 mm long; with margins straight; wedge-shaped; not in groove of raphe; Species Studied—Species in Genus: 3 spp.—7 spp. adjacent to hilum; mounded; dissimilar color from testa; lighter than or darker than testa; black or tan; not Fruit a legume; unilocular; 1.5Ð4 ϫ 0.3Ð0.6 ϫ 0.2Ð0.3 cm; within corona, halo, or rim. Endosperm thin; covering with persistent calyx; with calyx shorter than fruit; entire embryo; adnate to testa. Cotyledons smooth; both without orifice formed by curving of fruit or fruit outer faces convex; both the same thickness; both more segments; straight or curved (slightly); not plicate or or less of equal length; not folded; margin entire 180 plicate; not twisted; asymmetrical or symmetrical; degrees from base of radicle; similar at apex; not nearly oblong or linear; when asymmetrical with both concealing radicle; entire over radicle; without lobes; sutures nearly straight; not inflated; flattened; without with the interface division terminating at base of beak; short tapered at apex; apex aligned with longitu- radicle; without margins recessed; yellow; inner face dinal axis of fruit; short tapered at base; with the apex flat; glabrous around base of radicle. Embryonic axis and base uniform in texture; chartaceous or ligneous deflexed; oblique to length of seed. Radicle linear; (according to Stirton (1986b)); seed chambers exter- deflexed and parallel to cotyledon length; centered nally visible; with the raised seed chambers torulose. between cotyledons; 1/2 to nearly length of cotyledons. Fruit margin not constricted; without sulcus; plain. Fruit Plumule moderately developed; glabrous. wings absent. Fruit substipitate. Fruit indehiscent. Replum invisible. Epicarp dull; dark brown to black Distribution: South Africa (southwestern Cape Province). (but appearing gray because of hairs); pubescent and indurate; with 1 type of pubescence; villous; with Notes: When he founded the genus, Stirton (1986b) noted pubescence gray; with simple hairs; pliable; with hair that Polhillia “has a natural affinity with Melolobium bases plain; eglandular; without spines; not smooth; and Dichilus,” (30.01) and (30.02) respectively. Polhill with elevated features; not veined; not tuberculate; (1981p) and Wyk (1991) placed this genus in shagreen; not exfoliating; without cracks. Mesocarp Crotalarieae but later transferred it to the beginning of thin; surface not veined; 1-layered; without balsamic the Genisteae (30) (Polhill 1994a,b, Wyk and Schutte vesicles; without fibers; solid; coriaceous. Endocarp 1995). Its number of species and distribution were dull; monochrome; grayish brown; smooth; nonseptate taken from Wyk (1991) and Wyk and Schutte (1995). (but fruit impressed between seeds); chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 4Ð8; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; triangular or thick; straight or curved (slightly). Aril dry; rim-aril; reddish brown.

Seed 2.3Ð3 ϫ 2.3Ð3 ϫ 2Ð2.2 mm; not overgrown; not angular; asymmetrical; obliquely cordate or mitaform; compressed; with visible radicle and cotyledon lobes; with umbo (nearly) on seed faces; with umbo on both faces of seed. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome (if green may be blotchy purple but not mottled and streaked greenish to dark brown or tan); glabrous; smooth or not smooth; with elevated features; wrinkled (based on seed and not testa characters); coriaceous. Fracture lines Polhillia: P. pallens C.H. Stirton (CÐE), P spp. (AÐB). A, absent. Rim absent. Wings absent. Raphe not visible. Fruits (ϫ 2.2); B, seeds (ϫ 7); CÐD, testa (ϫ 50, Hilum fully concealed; concealed by aril; with faboid ϫ 1000); E, embryos (ϫ 10).

1083 Genus: Adenocarpus A.-P. de Candolle compressed or terete; with surface smooth; with or without visible radicle and cotyledon lobes; without Phylogenetic Number: 30.05. external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa Tribe: Genisteae. not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome or mottled and Species Studied—Species in Genus: 12 spp.—15 spp. streaked; with frequent mottles; with frequent streaks; reddish to greenish brown, green, olive, or black; with Fruit a legume; unilocular; 1.5Ð6 ϫ 0.4Ð1 ϫ 0.03Ð0.1 cm; purple or black overlay; glabrous; smooth; coriaceous. with persistent or deciduous calyx; with calyx shorter Fracture lines absent. Rim absent. Wings absent. Raphe than fruit; without orifice formed by curving of fruit or not visible. Hilum partially concealed; concealed by fruit segments; slightly curved or straight; not plicate; aril; with faboid split; with the lips of the faboid split not twisted; asymmetrical or symmetrical; oblong, the same color as the rest of the hilum; larger than or linear, or C-shaped; when asymmetrical with both punctiform; 0.3Ð0.4 mm long; with curved outline; sutures parallelly curved or nearly straight; not inflated; circular; between cotyledon and radicle lobe, apical at compressed or flattened; without or with beak; with apex of radicle tip, or subapical to radicle tip; recessed; solid beak the same color and texture as fruit; short within rim. Hilum rim color of testa. Lens discernible; tapered or rounded at apex; apex aligned or oblique equal to or greater than 0.5 mm in length; up to 1 mm with longitudinal axis of fruit; short tapered or rounded long; with margins straight or curved; triangular; at base; base aligned or oblique with longitudinal axis circular; not in groove of raphe; confluent with or of fruit; with the apex and base uniform in texture; adjacent to hilum; up to 0.4 mm from hilum; mounded chartaceous; seed chambers externally visible; with the or flush; similar color as or dissimilar color from testa; raised seed chambers not torulose. Fruit margin not darker than testa; black; not within corona, halo, or rim. constricted; without sulcus; plain. Fruit wings absent. Endosperm thick; covering entire embryo; adnate to Fruit substipitate. Fruit with all layers dehiscing; embryo. Cotyledons smooth; both outer faces convex; splitting along sutures. Dehiscence of valves along both both the same thickness; both more or less of equal sutures; apical and down; active; with valves twisting length; not folded; margin entire 180 degrees from base (loosely). Replum invisible. Epicarp glossy or dull; of radicle; similar at apex; not concealing radicle; split monochrome (though conspicuous glandular tuberculae over radicle; with lobes; with the interface division may be darker colored); brown to light or dark reddish terminating at base of radicle; without margins re- brown; glabrous or pubescent and indurate; with hairs cessed; white or yellow; inner face flat; glabrous around erect or appressed; with 1 type of pubescence; puberu- base of radicle. Embryonic axis deflexed; oblique to lent, villous, or sericeous; with pubescence gray or length of seed; without a joint evident between the golden; with pubescence uniformly distributed; with radicle and the cotyledons. Radicle bulbose or linear; simple hairs; pliable; with hair bases plain; glandular; lobe tip straight; deflexed and parallel to cotyledon with glandular papillae (long to short); without spines; length; centered between cotyledons; equaling length of not smooth; with elevated features; reticulately veined; cotyledons. Plumule rudimentary; glabrous. not tuberculate; papillose; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; Distribution: Mediterranean region, Canary Islands, highest without balsamic vesicles; without fibers; solid; mountains of tropical Africa, and Syria. coriaceous. Endocarp dull; monochrome; brown; smooth; nonseptate (but may be slightly scurfy between Notes: Gibbs (1967) monographed the genus, and Luaces seeds); chartaceous; not exfoliating; remaining fused to (1972) reported on the seed germination and chromo- mesocarp and epicarp; entire. Seeds 1Ð8; length parallel some counts of the Iberian species. Greinwald et al. with fruit length; neither overlapping nor touching; in 1 (1992) reported on the taxonomic significance of series. Funiculus measured; up to 1 mm long; of 1 alkaloids in Adenocarpus, and Veen et al. (1992) length only; thick; straight or curved. Aril dry; rim-aril reported on the alkaloids of A. hispanicus (J.P.A.P.M. or tongue-aril (somewhat); tan. de Lamarck) A.-P. de Candolle.

Seed 2Ð5.5 ϫ 2Ð5.5 ϫ 1.2Ð2.5 mm; not overgrown; not Adenocarpus: A. decorticans P.E. Boissier (CÐE), A. spp. angular; asymmetrical; quadrangular, mitaform, oblong, (AÐB). A, Fruits (closed and dehisced) (ϫ 1.4); B, seeds ovate, circular (more or less), rectangular, or reniform; (ϫ 4.7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

1085 Genus: Anarthrophyllum G. Bentham partially concealed; concealed by funicular remnant; with faboid split; with the lips of the faboid split the Phylogenetic Number: 30.06. same color as the rest of the hilum; punctiform; apical according to radicle tip but marginal according to seed Tribe: Genisteae. length; recessed; not within corona, halo, or rim. Lens discernible; equal to or greater than 0.5 mm in length; Species Studied—Species in Genus: 13 spp.—15 spp. 0.9 mm long; with margins straight; wedge-shaped; not in groove of raphe; adjacent to hilum; 0.1 mm from Fruit a legume; unilocular; 0.7Ð4 ϫ 0.4Ð0.9 ϫ 0.2Ð0.8 cm; hilum; flush; similar color as testa; darker than testa; with persistent calyx; with calyx shorter than fruit; not within corona, halo, or rim. Endosperm thin; without orifice formed by curving of fruit or fruit covering entire embryo; adnate to testa. Cotyledons segments; straight or curved (slightly); not plicate; not smooth; both outer faces convex; both the same twisted; asymmetrical or symmetrical; lanceolate, thickness; both more or less of equal length; not folded; linear, oblong, ovate, or rhombic; when asymmetrical margin entire 180 degrees from base of radicle; similar with both sutures parallelly to unequally curved; not at apex; not concealing radicle; entire over radicle; inflated; compressed; without or with beak; with solid without lobes; with the interface division terminating at beak the same color and texture as fruit; tapered or base of radicle; without margins recessed; tan; inner short tapered at apex; apex oblique with longitudinal face flat; glabrous around base of radicle. Embryonic axis of fruit; long tapered or tapered at base; base axis parallel (radicle length parallel to cotyledon oblique with longitudinal axis of fruit; with the apex length); parallel to length of seed. Radicle linear; lobe and base uniform in texture; chartaceous or coriaceous; tip straight; deflexed and parallel to cotyledon length; seed chambers externally visible; with the raised seed centered between cotyledons; exceeding length of chambers not torulose. Fruit margin not constricted; cotyledons. Plumule moderately developed; glabrous. without sulcus; plain. Fruit wings absent. Fruit substipitate or nonstipitate. Fruit with all layers Distribution: Chile and Argentina (Andes). dehiscing; splitting along suture. Dehiscence of valves along 1 suture; apical and down; active; with valves Notes: Soraru (1974) monographed the genus and illustrated twisting. Replum invisible. Epicarp dull; monochrome; the external characters of its fruits and seeds. Polhill brown; pubescent and indurate or glabrous to glabrate; (1981q) and Soraru both noted that Anarthrophyllum with 1 type of pubescence; tomentose or villous; with seeds often bulge on the side opposite the hilum (or pubescence gray; with pubescence uniformly distrib- radicle tip). This bulge is a normal legume seed with its uted; with simple hairs; pliable; with hair bases plain; axis parallel to the hilum length shorter than the axis at eglandular; without spines; not smooth; with elevated right angles to the hilum length, that is, the seeds are features; reticulately veined; not tuberculate; not wider than they are long. Polhill continued that exfoliating; without cracks. Mesocarp thin; surface not Anarthophyllum is a “very remarkable genus of veined; 1-layered; without balsamic vesicles; without uncertain affinities, apart from its probable relationship fibers; solid; coriaceous. Endocarp dull; monochrome; with Sellocharis” (30.07). Polhill (1981q) and Wyk brown; smooth; nonseptate; chartaceous; not exfoliat- (1991) had this genus in Crotalarieae but later trans- ing; remaining fused to mesocarp and epicarp; entire. ferred it to the beginning of the Genisteae (30) (Polhill Seeds 1Ð8; length transverse to fruit length; neither 1994a,b, Wyk and Schutte 1995). overlapping nor touching; in 1 series. Funiculus measured; 1.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 2.5Ð5 ϫ 1.5Ð4 ϫ 1.5Ð1.7 mm; not overgrown; not angular; symmetrical (except for hilum); circular, oblong, or ovate; compressed; without visible radicle and cotyledon lobes; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a Anarthrophyllum: A. rigidum (J. Gilles ex W.J. Hooker & bloom; colored; monochrome; reddish brown to brown; G.A.W. Arnott) G.H.E.W. Hieronymus (CÐE), A. spp. glabrous; smooth; coriaceous. Fracture lines absent. (AÐB). A, Fruits and valve (ϫ 1.9); B, seeds (ϫ 4.8); CÐ Rim absent. Wings absent. Raphe not visible. Hilum D, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

1087 Genus: Lupinus C. Linnaeus quadrangular; compressed; with surface smooth; without visible radicle and cotyledon lobes; without Phylogenetic Number: 30.08. hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull or glossy; not modified by a Tribe: Genisteae. bloom; colored or clear; monochrome, mottled and streaked, or bichrome (ivory and reddish-brown); with Species Studied—Species in Genus: 75 spp.—200 spp. frequent mottles; with frequent streaks; dark reddish brown, cream, tan, white, gray (to bluish), pink, or blue; Fruit a legume; unilocular; 2Ð9 ϫ 0.3Ð2 ϫ 0.18Ð0.8 cm; with brown, gray (to bluish), red, or black overlay; with persistent calyx; with calyx shorter than fruit; glabrous; smooth or not smooth; with elevated or without orifice formed by curving of fruit or fruit recessed features; wrinkled, shagreen, or rugose; segments; straight; not plicate; not twisted; asymmetri- concaved; coriaceous. Fracture lines absent. Rim cal or symmetrical; oblong or ovate; when asymmetri- absent. Wings absent. Raphe not visible or visible; from cal with both sutures nearly straight; not inflated; hilum to lens or through lens and terminating before compressed, terete, or flattened; without beak; short base of seed; not bifurcating; color of or darker than tapered at apex; apex aligned with longitudinal axis of testa; reddish brown; recessed. Hilum visible or fruit; long tapered or short tapered at base; base aligned partially concealed; concealed by funicular remnant; with longitudinal axis of fruit; with the apex and base with faboid split; with the lips of the faboid split the uniform in texture; coriaceous, membranous, same color as or lighter than the rest of the hilum and chartaceous, or ligneous; seed chambers externally therefore conspicuous; larger than punctiform; up to 2 visible or invisible; with the raised seed chambers not mm long; with angular, curved, or straight outline; torulose. Fruit margin not constricted; without sulcus; circular, oval, wedge-shaped, triangular (more or less), plain. Fruit wings absent. Fruit substipitate or or oblong; subapical or marginal according to radicle nonstipitate. Fruit with all layers dehiscing; splitting tip; recessed; not within corona, halo, or rim or within along sutures. Dehiscence of valves along both sutures; rim or halo. Hilum halo color of or lighter than testa. apical and down; active; with valves twisting. Replum Hilum rim color of testa. Lens discernible or not invisible. Epicarp dull or glossy; monochrome or discernible; equal to or greater than 0.5 mm in length; multicolored; mottled; brown to dark reddish brown; up to 3 mm long; with margins curved; circular (often with black overlay; pubescent and indurate, glabrous, or with linear center) or elliptic; not in or in groove of pubescent but soon deciduous; with 1 type of pubes- raphe; adjacent to hilum; up to 3 mm from hilum; cence; tomentose, villous, or puberulent; with pubes- mounded; dissimilar color from testa; lighter or darker cence golden or gray; with pubescence uniformly than testa; reddish tan or black; not within corona, halo, distributed; with simple hairs; pliable; with hair bases or rim or within rim (its own). Lens rim color of testa. plain; eglandular; without spines; not smooth; with Endosperm thin; covering entire embryo; adnate to elevated features; reticulately veined; not tuberculate; testa. Cotyledons smooth; both outer faces convex; both not exfoliating; without cracks. Mesocarp thin; surface the same thickness; both more or less of equal length; not veined; 1-layered; without balsamic vesicles; not folded; margin etire 180 degrees from base of without fibers; solid; coriaceous, ligneous (sub), or radicle; similar at apex; not concealing radicle; split chartaceous. Endocarp dull or glossy; monochrome; over radicle; with lobes; with lobes not touching; with brown or tan; spongy; septate, subseptate, or the interface division terminating at base of radicle or in nonseptate; with septa thicker than paper, firm; with radicle tissue; without margins recessed; tan; inner face septa eglandular; chartaceous; not exfoliating or flat or wrinkled; glabrous around base of radicle. exfoliating in part; remaining fused to mesocarp and Embryonic axis deflexed; oblique to length of seed; epicarp; entire. Seeds 1Ð15; length parallel with or without a joint evident between the radicle and the transverse to fruit length; neither overlapping nor cotyledons. Radicle linear or triangular; lobe tip touching or touching; in 2 or more series. Funiculus less straight; deflexed and parallel to cotyledon length; than 0.5 mm long; of 1 length only; thick or triangular; centered between cotyledons; less than 1/2, 1/2 to straight or curved. Aril absent or present; dry; rim-aril; nearly, or equaling length of cotyledons. Plumule white. rudimentary, moderately developed, or well developed; glabrous. Seed 1.5Ð9 ϫ 1.5Ð8 ϫ 1Ð5 mm; not overgrown; not angular or angular; asymmetrical; ovate, circular, oblong, or

1088 Distribution: New World, Mediterranean region, tropical highlands of Africa, and naturalized elsewhere.

Notes: Cristofolini and Chiapella (1977) studied the water- soluble seed proteins of 45 species of Lupinus. Phillips (1955) monographed the perennial lupine species of North America, and Dunn and Gillett (1966) monographed the taxa in Canada. Heyn and Herrnstadt (1977) studied the testae of Old World species. Lawson (1982) edited an overview of the agronomic value, liability, and problems in growing Lupinus species in Australia. Summerfield and Roberts (1985b) also reported on the economic value of the genus. Plitmann (1981) traced the evolutionary history of the Old World lupines.

1089 Lupinus: L. tauris G. Bentham (CÐE), L. spp. (AÐB). A, Fruits (closed and dehisced) (ϫ 1.1); B, seeds (ϫ 1.8); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 2).

1091 Genus: Laburnum P.C. Fabricius and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; Phylogenetic Number: 30.09. without umbo on seed faces. Testa not adhering to endocarp; dull or glossy (somewhat); not modified by a Tribe: Genisteae. bloom; colored; monochrome; dark to bright reddish brown or orange; glabrous; smooth; coriaceous. Species Studied—Species in Genus: 2 spp.—2 spp. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the Fruit a legume; unilocular; 2Ð6 ϫ 0.7Ð1 ϫ 0.15Ð0.4 cm; lips of the faboid split the same color as the rest of the with deciduous or persistent calyx; with calyx shorter hilum; larger than punctiform; up to 0.5 mm long; with than fruit; without orifice formed by curving of fruit or curved outline; circular; between cotyledon and radicle fruit segments; straight or curved (or slightly curved); lobe; recessed; within rim. Hilum rim color of testa. not plicate; not twisted; symmetrical or asymmetrical; Lens discernible; equal to or greater than 0.5 mm in linear or oblong (narrowly); when asymmetrical with length; 0.5Ð1 mm long; with margins straight or curved; both sutures nearly straight; not inflated; flattened or wedge-shaped or elliptic; not in groove of raphe; compressed; without beak; long tapered, tapered, or adjacent to or confluent with hilum; 0.2 mm from short tapered at apex; apex aligned or oblique (slightly) hilum; barely mounded; similar color as testa; reddish with longitudinal axis of fruit; long tapered or tapered brown; not within corona, halo, or rim. Endosperm thin; at base; base aligned or oblique (slightly) with longitu- covering entire embryo; adnate to testa. Cotyledons dinal axis of fruit; with the apex and base uniform in smooth; both outer faces convex; both the same texture; coriaceous, membranous, or ligneous; seed thickness; both more or less of equal length; with both chambers externally visible; with the raised seed folded (barely) or not folded; not sufficiently folded for chambers not torulose or torulose. Fruit margin not inner face to touch itself; portions of inner folded face constricted or constricted; slightly constricted (hardly) unequal; margin entire 180 degrees from base of along both margins; without sulcus; plain. Fruit wings radicle; similar at apex; not concealing radicle; split absent. Fruit substipitate. Fruit with all layers dehiscing over radicle; with lobes; with the interface division (tardily); splitting along sutures. Dehiscence of valves terminating at base of radicle; without margins re- along both sutures; apical and down; passive. Replum cessed; red; inner face flat; glabrous around base of invisible. Epicarp dull; monochrome; dark to reddish radicle. Embryonic axis deflexed; oblique to length of brown or tan; glabrous or pubescent and indurate; with seed; without a joint evident between the radicle and hairs erect or appressed; with 1 type of pubescence; the cotyledons. Radicle bulbose; lobe tip straight; puberulent; with pubescence golden; with pubescence deflexed and parallel to cotyledon length; centered uniformly distributed; with simple hairs; pliable; with between cotyledons; less than 1/2 length of cotyledons. hair bases plain; eglandular; without spines; not Plumule rudimentary; glabrous. smooth; with elevated features; longitudinally veined relative to fruit length; not tuberculate; longitudinally Distribution: Southern Europe. and irregularly wrinkled; exfoliating in part or not exfoliating; without cracks. Mesocarp thin; surface not Notes: Forrester (1979) stated that an average of about veined; 1-layered; without balsamic vesicles; without 3,000 children per summer are poisoned by fruits, fibers; solid; subligneous or coriaceous. Endocarp dull; seeds, and flowers of Laburnum in England. monochrome; reddish brown or tan; smooth; nonseptate or subseptate (because of intrusion of fruit); with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð8; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; up to 1.5 mm long; of 1 length only; thick; curved. Aril absent.

Seed 4Ð5 ϫ 3.5Ð4 ϫ 2.5Ð2.8 mm; not overgrown; not angular or angular (somewhat); asymmetrical; oblong, Laburnum: L. anagyroides F.C. Medikus (CÐE), L. spp. (AÐ reniform, triangular (more or less), or mitaform; B). A, Fruits (closed and dehisced) (ϫ 1.4); B, seeds compressed; with surface smooth; with visible radicle (ϫ 4.8); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

1093 Genus: Hesperolaburnum R.C.J.E. Maire color as the rest of the hilum; punctiform; apical at apex of radicle tip or between cotyledon and radicle lobe; Phylogenetic Number: 30.10. recessed; within rim. Hilum rim color somewhat darker than testa. Lens discernible; equal to or greater than 0.5 Tribe: Genisteae. mm in length; up to 1 mm long; with margins straight or curved; oblong; not in groove of raphe; confluent Species Studied—Species in Genus: 1 sp.—1 sp. with hilum (hilar rim extends out to lens); mounded; similar color as testa; lighter than testa; reddish brown; ϫ ϫ Fruit a legume; unilocular; 4 1 0.5 cm; with deciduous not within corona, halo, or rim. Endosperm thick; or persistent calyx; with calyx shorter than fruit; covering entire embryo; adnate to embryo. Cotyledons without orifice formed by curving of fruit or fruit smooth; both outer faces convex; both the same segments; straight; not plicate; not twisted; asymmetri- thickness; both more or less of equal length; not folded; cal; oblong; when asymmetrical with 1 straight and 1 margin entire 180 degrees from base of radicle; similar curved suture, both sutures unequally curved, or both at apex; not concealing radicle; split over radicle; with sutures parallelly curved; widest near middle or D- lobes; with the interface division terminating at base of shaped; not inflated; compressed; without beak; short radicle; without margins recessed; tan; inner face flat or tapered at apex; apex oblique with longitudinal axis of concave (or one concave and other convex); glabrous fruit; tapered at base; base aligned with longitudinal around base of radicle. Embryonic axis deflexed; axis of fruit; with the apex and base uniform in texture; oblique to length of seed; without a joint evident coriaceous; seed chambers externally visible; with the between the radicle and the cotyledons. Radicle raised seed chambers not torulose. Fruit margin slightly bulbose; lobe tip straight; deflexed and parallel to constricted only on 1 margin; without sulcus; embel- cotyledon length; centered between cotyledons; less lished. Fruit wings absent. Fruit substipitate or than 1/2 length of cotyledons. Plumule rudimentary; nonstipitate. Fruit with all layers dehiscing (to tardily glabrous. so); splitting along sutures. Dehiscence of valves along both sutures; apical and down; passive. Replum Distribution: Morocco (Anti-Atlas mountains). invisible. Epicarp glossy; monochrome; dark reddish brown; glabrous; eglandular; without spines; not smooth; with elevated features; reticulately veined; not tuberculate; warty (tan); not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; pale brown; scurfy; septate; with septa thin (tissue paper-like), flexible; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð3; length transverse to, oblique to, or parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened; curved. Aril absent.

Seed 5.5Ð6 ϫ 4.5Ð5 ϫ 3Ð3.3 mm; not overgrown; not angular; asymmetrical; ovate; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; dark reddish brown; glabrous; smooth; coriaceous. Fracture lines absent. Hesperolaburnum: H. platycarpum (R.C.J.E. Maire) Rim absent. Wings absent. Raphe not visible. Hilum R.C.J.E. Maire (AÐE). A, Valves showing endocarp and fully concealed; concealed by funicular remnant; with epicarp (ϫ 1.7); B, seeds (ϫ 4.3); CÐD, testa (ϫ 50, faboid split; with the lips of the faboid split the same ϫ 1000); E, embryos (ϫ 5).

1095 Genus: Podocytisus P.E. Boissier & T.H.H. von Heldreich seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome; dark Phylogenetic Number: 30.11. reddish brown or mixture of dark and darker reddish brown; glabrous; smooth; coriaceous. Fracture lines Tribe: Genisteae. absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the Species Studied—Species in Genus: 1 sp.—1 sp. faboid split the same color as the rest of the hilum; larger than punctiform; 0.7 mm long; with straight Fruit a legume; unilocular; 2.5Ð4.5 ϫ 1Ð2 ϫ 0.03Ð0.05 cm; outline; oblong; between cotyledon and radicle lobe; with persistent calyx; with calyx shorter than fruit; slightly flush; not within corona, halo, or rim or within without orifice formed by curving of fruit or fruit rim (faint). Hilum rim color of testa. Lens discernible; segments; curved; not plicate; not twisted; asymmetri- less than or equal to or greater than 0.5 mm in length; cal; oblong; when asymmetrical with both sutures 0.5 mm long; with margins curved; circular (with linear nearly straight; not inflated; flattened; without beak; center); not in groove of raphe; confluent with or rounded at apex; apex oblique with longitudinal axis of adjacent to hilum; up to 0.2 mm from hilum; flush; fruit; rounded or short tapered at base; base oblique dissimilar color from testa; lighter than testa; reddish with longitudinal axis of fruit; with the apex and base tan; not within corona, halo, or rim. Endosperm thin; uniform in texture; membranous; seed chambers covering entire embryo; adnate to testa. Cotyledons externally visible; with the raised seed chambers not smooth; both outer faces convex; both the same torulose. Fruit margin not constricted or constricted; thickness; both more or less of equal length; not folded; slightly constricted (and irregularly) only on 1 margin; margin entire 180 degrees from base of radicle; similar without sulcus; embellished. Fruit wing 1; 1.5 mm at apex; not concealing radicle; split over radicle; with wide; sutural; on both valves; on 1 suture. Fruit lobes; with the interface division terminating at base of substipitate. Fruit indehiscent or with all layers dehisc- radicle; without margins recessed; tan; inner face flat; ing (tardily); splitting along suture. Dehiscence of glabrous around base of radicle. Embryonic axis valves along 1 suture; probably medial and up and deflexed; oblique to length of seed; without a joint down; passive. Replum invisible. Epicarp dull; mono- evident between the radicle and the cotyledons. Radicle chrome or multicolored; bichrome (irregularly, no linear; lobe tip straight; deflexed and parallel to pattern) or mottled and streaked (neither true mottling cotyledon length; centered between cotyledons; less nor true streaking); brown (greenish with irregular than 1/2 length of cotyledons. Plumule rudimentary; dark-brown to purple patches), green, purple, or tan glabrous. (greenish); with brown (dark brown to dark purple) overlay; glabrous; eglandular; without spines; not Distribution: Balkans and Turkey (Anatolia). smooth; with elevated features; reticulately veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; chartaceous (or thinner). Endocarp dull or glossy (without “cobwebs”); monochrome; tan; cobwebby; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 2Ð6; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; up to 3 mm long (most of length fused with valve and 1 branched); of 1 length only; flattened; curved. Aril absent.

Seed 4.5Ð5 ϫ 4Ð4.5 ϫ 2Ð2.8 mm; not overgrown; not angular or angular (somewhat); asymmetrical; oblong, circular, or reniform (more or less); compressed; with surface smooth; with visible radicle and cotyledon Podocytisus: P. caramanicus P.E. Boissier & T.H.H. von lobes; without external groove between radicle and Heldreich (AÐE). A, Fruits (ϫ 1.7); B, seeds (ϫ 4.2); cotyledon lobes; without hilar sinus; without umbo on CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

1097 Genus: Cytisophyllum O.F. Lang adnate to embryo. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of Phylogenetic Number: 30.12. equal length; not folded; margin entire 180 degrees from base of radicle; similar at apex; not concealing Tribe: Genisteae. radicle; split over radicle; with lobes; with the interface division terminating at base of radicle; without margins Species Studied—Species in Genus: 1 sp.—1 sp. recessed; tan; inner face flat; glabrous around base of radicle. Embryonic axis deflexed; oblique to length of Fruit a legume; unilocular; 2.5Ð4 ϫ 1 cm; without orifice seed; without a joint evident between the radicle and formed by curving of fruit or fruit segments; straight; the cotyledons. Radicle linear; lobe tip straight; not plicate; not twisted; symmetrical or asymmetrical; deflexed and parallel to cotyledon length; centered linear to oblong; when asymmetrical with both sutures between cotyledons; equaling length of cotyledons. nearly straight; not inflated; compressed; with the apex Plumule rudimentary; glabrous. and base uniform in texture; coriaceous. Fruit margin without sulcus; plain. Fruit wings absent. Fruit with all Distribution: Eastern Spain, southern France, and Italy. layers dehiscing; splitting along sutures. Dehiscence of valves along both sutures; apical and down; active; with Notes: No entire fruits were available for photographing. valves twisting. Replum invisible. Epicarp glossy; monochrome; glabrous; eglandular; without spines; smooth; not veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1- layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; reddish tan; smooth and scurfy; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seed 3-6 (Pignatti 1982); neither overlapping nor touching; in 1 series. Funiculus of 1 length only. Aril fleshy; topknotlike or cupshaped; covering less than 1/2 of seed; yellow.

Seed 4Ð5 ϫ 3Ð4 ϫ 1.2Ð1.7 mm; not overgrown; angular or not angular; asymmetrical; oblong, ovate, circular (more or less), triangular, or irregular (somewhat); compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; colored; monochrome; dark reddish to greenish brown or black; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum partially concealed; concealed by aril; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.6 mm long; with curved outline; elliptic; apical at apex of radicle tip and between cotyledon and radicle lobe; recessed; within rim (inconspicuous). Hilum rim color of testa. Lens discernible; less than 0.5 mm in length; with margins curved; circular; not in groove of raphe; confluent with hilum; flush; dissimilar color Cytisophyllum: C. sessilifolium (C. Linnaeus) O.F. Lang (AÐ from testa; lighter than testa; tan; not within corona, D). A, Seeds (ϫ 4.4); BÐC, testa (ϫ 50, ϫ 1000); D, halo, or rim. Endosperm thick; covering entire embryo; embryos (ϫ 6).

1099 Genus: Petteria C.B. Presl Wings absent. Raphe not visible. Hilum visible, partially concealed, or fully concealed; concealed by Phylogenetic Number: 30.13. funiculus, funicular remnant, radicle lobe, or wing (somewhat); with faboid split; with the lips of the Tribe: Genisteae. faboid split the same color as the rest of the hilum; larger than punctiform; 0.5 mm long; with straight Species Studied—Species in Genus: 1 sp.—1 sp. (more or less) outline; oblong; apical at apex of radicle tip; recessed; within rim (which involves radicle lobe Fruit a legume; unilocular; 2.5Ð5 ϫ 0.5Ð1 cm; with persis- and open on lens side). Hilum rim color of testa. Lens tent calyx; with calyx shorter than fruit; without orifice barely discernible; less than 0.5 mm in length; with formed by curving of fruit or fruit segments; straight or margins straight; tiny wedge-shaped or rhombic (tiny); curved (slightly); not plicate; not twisted; symmetrical not in groove of raphe; adjacent to hilum; 0.5 mm from or asymmetrical; broadly linear or falcate; when hilum; barely mounded; dissimilar color from testa; asymmetrical with 1 straight and 1 curved suture or darker than testa; dark reddish brown; not within both sutures parallelly curved; not inflated; compressed; corona, halo, or rim. Endosperm thick; covering entire without beak; short tapered at apex; apex aligned with embryo; adnate to embryo. Cotyledons smooth; both longitudinal axis of fruit; short tapered at base; base outer faces convex; both the same thickness; both more aligned with longitudinal axis of fruit; with the apex or less of equal length; not folded; margin entire 180 and base uniform in texture; coriaceous; seed chambers degrees from base of radicle; similar at apex; not externally visible; with the raised seed chambers not concealing radicle; split over radicle; with lobes; with torulose. Fruit margin not constricted; without sulcus; the interface division terminating at base of radicle; plain. Fruit wings absent. Fruit substipitate. Fruit with without margins recessed; tan; inner face flat; glabrous all layers dehiscing; splitting along sutures. Dehiscence around base of radicle. Embryonic axis deflexed; of valves along both sutures; apical and down; active; oblique to length of seed; without a joint evident with valves twisting. Replum invisible. Epicarp dull; between the radicle and the cotyledons. Radicle linear; monochrome; dark reddish brown; glabrous; lobe tip straight; deflexed and parallel to cotyledon eglandular; without spines; not smooth; with elevated length; centered between cotyledons; 1/2 to nearly, features; not veined; not tuberculate; wrinkled or equaling, or exceeding length of cotyledons. Plumule shagreen; not exfoliating or exfoliating in part; without rudimentary; glabrous. cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; Distribution: Balkans. coriaceous. Endocarp dull; monochrome; brown; spongy and scurfy (between seeds); nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 8; length parallel with or transverse to fruit length; neither overlapping nor touching or touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; flattened or triangular; straight. Aril dry; tongue-aril (nearly); tan.

Seed 4.5Ð5 ϫ 2.5Ð4.5 ϫ 1.6Ð2.2 mm; not overgrown; not angular or angular; symmetrical; rectangular, oblong, rhombic, or triangular; compressed or terete; with surface smooth; with visible radicle and cotyledon lobes; without or with external groove between radicle and cotyledon lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; monochrome; orangish to greenish and reddish Petteria: P. ramentacea (F.W. Sieber) C.B. Presl (AÐE). A, brown, tan (orangish), green, or orange; glabrous; Fruit (dehisced) and valve (ϫ 2); B, seeds (ϫ 3.7); CÐ smooth; coriaceous. Fracture lines absent. Rim absent. D, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

1101 Genus: Argyrocytisus (R.C.J.E. Maire) D.G. Frodin & V.H. partially concealed; concealed by aril; with faboid split; Heywood ex C. Raymaud with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 0.4 mm long; Phylogenetic Number: 30.14. with curved outline; oval; apical at apex of radicle tip and between cotyledon and radicle lobe; recessed; Tribe: Genisteae. within rim. Hilum rim color darker than testa (slightly to black). Lens discernible; equal to or greater than 0.5 Species Studied—Species in Genus: 1 sp.—1 sp. mm in length; 0.7Ð0.9 mm long; with margins straight; linear; not in groove of raphe; confluent with hilum; Fruit a legume; unilocular; 5.5 ϫ 0.7 cm; without orifice slightly mounded; dissimilar color from testa; lighter formed by curving of fruit or fruit segments; slightly than testa; tan; not within corona, halo, or rim. En- curved; not plicate; not twisted; asymmetrical or dosperm thick; covering entire embryo; adnate to symmetrical; linear; when asymmetrical with both embryo. Cotyledons smooth; both outer faces convex; sutures nearly straight; not inflated; compressed; both the same thickness; both more or less of equal without beak; short tapered at apex; apex aligned with length; not folded; not sufficiently folded for inner face longitudinal axis of fruit; short tapered at base; base to touch itself; margin entire 180 degrees from base of aligned with longitudinal axis of fruit; with the apex radicle; similar at apex; not concealing radicle; split and base uniform in texture; coriaceous; seed chambers over radicle; with lobes; with the interface division externally visible. Fruit margin not constricted; without terminating at base of radicle; without margins re- sulcus; plain. Fruit wings absent. Fruit nonstipitate. cessed; yellow; inner face flat; glabrous around base of Fruit with all layers dehiscing; splitting along sutures. radicle. Embryonic axis deflexed; oblique to length of Dehiscence of valves along both sutures; apical and seed; without a joint evident between the radicle and down; active; with valves twisting. Replum invisible or the cotyledons. Radicle bulbose; lobe tip straight; visible. Epicarp dull; monochrome; pubescent and deflexed and parallel to cotyledon length; centered indurate; with 1 type of pubescence; pilose; with between cotyledons; equaling length of cotyledons. pubescence gray; with pubescence uniformly distrib- Plumule rudimentary; glabrous. uted; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated Distribution: Africa (Atlas Mountains). features; veined or not veined; reticulately veined (faintly near suture); not tuberculate; faintly wrinkled; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; tan; smooth; nonseptate (but fringe of scurfy, light-colored material along ridges between seeds); chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 8; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 0.5 mm long; of 1 length only; thick; straight. Aril dry; tongue-aril; white.

Seed 3.5Ð4 ϫ 3Ð3.5 ϫ 2Ð2.3 mm; not overgrown; not angular; asymmetrical; oblong or rectangular; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled and streaked (both faintly); dark reddish to greenish brown; with brown overlay; Argyrocytisus: A. battandieri (R.C.J.E. Maire) C. Raynaud glabrous; smooth; coriaceous. Fracture lines absent. (AÐE). A, Valves (ϫ 1.5); B, seeds (ϫ 4.5); CÐD, testa Rim absent. Wings absent. Raphe not visible. Hilum (ϫ 50, ϫ 1000); E, embryos (ϫ 6).

1103 Genus: Cytisus R.L. Desfontaines fleshy cupshaped; covering less than 1/2 of seed; when dry rim-aril; brown to reddish brown, tan (to reddish Phylogenetic Number: 30.15. tan), or orange.

Tribe: Genisteae. Seed 3Ð5 ϫ 2Ð4.2 ϫ 1.5Ð2.5 mm; not overgrown; not angular; asymmetrical; oblong, ovate, rectangular, Species Studied—Species in Genus: 13 spp.—35 spp. rhombic, circular, or D-shaped; compressed; with surface smooth; with or without visible radicle and Fruit a legume; unilocular; 1.2Ð7 ϫ 0.4Ð1.3 ϫ 0.1Ð0.3 cm; cotyledon lobes; without or with external groove with persistent or deciduous calyx; with calyx shorter between radicle and cotyledon lobes; with external than fruit; without orifice formed by curving of fruit or groove between radicle and cotyledon lobes same color fruit segments; straight or curved (slightly); not plicate; as testa; without hilar sinus; without umbo on seed not twisted; asymmetrical or symmetrical; linear, faces. Testa not adhering to endocarp; dull; not modi- oblong, or ovate; when asymmetrical with both sutures fied by a bloom; colored; monochrome or mottled and nearly straight; not inflated (more or less) or inflated; streaked; brown to blackish, reddish, or greenish brown, compressed or flattened; without beak; short tapered at tan (to greenish or reddish tan), red, orange, olive, apex; apex aligned or oblique with longitudinal axis of green, or black; with tan overlay; glabrous; smooth; fruit; short tapered at base; base aligned or oblique with coriaceous. Fracture lines absent. Rim absent. Wings longitudinal axis of fruit; with the apex and base absent. Raphe not visible. Hilum visible; with faboid uniform in texture; chartaceous or coriaceous; seed split; with the lips of the faboid split the same color as chambers externally invisible or visible; with the raised the rest of the hilum; larger than punctiform; 0.5 mm seed chambers not torulose. Fruit margin not con- long; with curved outline; circular; apical at apex of stricted or constricted; slightly constricted along both radicle tip and between cotyledon and radicle lobe, margins; without sulcus; plain. Fruit wings absent. Fruit subapical to radicle tip, or marginal according to radicle nonstipitate or substipitate. Fruit with all layers tip; recessed; within rim. Hilum rim color of or darker dehiscing; splitting along sutures. Dehiscence of valves than testa. Lens discernible; less or equal to or greater along both sutures; apical and down; active; with valves than 0.5 mm in length; 0.5 mm long; with margins twisting. Replum invisible. Epicarp dull; monochrome; straight or curved; oblong; not in groove of raphe; brown to dark reddish brown or black; pubescent and confluent with hilum; mounded; similar color as testa; indurate, pubescent but soon deciduous, or glabrous; darker than testa; reddish brown; not within corona, with hairs erect or appressed; with 1 or 2 types of halo, or rim. Endosperm thick; covering entire embryo; pubescence; villous (to restricted to sutures), pilose, or adnate to embryo. Cotyledons smooth; both outer faces tomentose; with pubescence gray, brown, or golden; convex; both the same thickness; both more or less of with long and short gray plain-tipped hairs; with equal length; not folded; margin entire 180 degrees pubescence uniformly distributed; with simple hairs; from base of radicle; similar at apex; not concealing pliable; with hair bases plain; eglandular; without radicle; split over radicle; with lobes; with the interface spines; not smooth; with elevated features; not veined division terminating at base of radicle; without margins or veined; reticulately veined; not tuberculate; faintly recessed; yellow or tan; inner face flat; glabrous around wrinkled; not exfoliating or exfoliating in part; without base of radicle. Embryonic axis deflexed; oblique to or with cracks; cracking oblique to fruit length. Meso- length of seed; without a joint evident between the carp thin; surface not veined; 1-layered; without radicle and the cotyledons. Radicle bulbose or linear; balsamic vesicles; without fibers; solid; coriaceous. lobe tip straight; deflexed and parallel to cotyledon Endocarp dull; monochrome; brown to blackish brown length; centered between coyledons; equaling, 1/2 to or tan; smooth; subseptate or nonseptate; with septa nearly, or exceeding length of cotyledons. Plumule composed of minute fringe of hairs; with septa rudimentary; glabrous. eglandular; chartaceous; not exfoliating or exfoliating in part; remaining fused to mesocarp and epicarp; Distribution: Europe, North Africa, Canary Islands, and entire. Seeds 8Ð14; length transverse to, oblique to, or Asia. parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long Notes: Talavera and Gibbs (1997) added two species to or measured; up to 1 mm long; of 1 length only; Cytisus, and the species count includes them. Polhill triangular; straight or curved. Aril fleshy or dry; when (1994b) accepted Lembotropis A.H.R Grisebach

1105 Cytisus: C. commutatus (H.M. Willkomm) J.I. Briquet (CÐ E), C. spp. (AÐB). A, Valves and fruit (dehisced) (ϫ 1.4); B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 11).

1107 Genus: Chamaecytisus J.H.F. Link circular (more or less), triangular (more or less), or cordate; compressed or terete; with surface smooth; Phylogenetic Number: 30.15A. with or without visible radicle and cotyledon lobes; without external groove between radicle and cotyledon Tribe: Genisteae. lobes; without hilar sinus; with umbo on seed faces. Cuticle wrinkled. Testa not adhering to endocarp; Species Studied—Species in Genus: 11 spp.—30 spp. glossy; not modified by a bloom; colored; monochrome; brown to reddish or greenish brown, tan (to Fruit a legume; unilocular; 2Ð7 ϫ 0.5Ð1.1 ϫ 0.2Ð0.4 cm; greenish tan), olive, or black; glabrous; smooth; with persistent or deciduous calyx; with calyx shorter coriaceous. Fracture lines absent. Rim absent. Wings than fruit; without orifice formed by curving of fruit or absent. Raphe not visible. Hilum partially concealed; fruit segments; straight or curved (slightly); not plicate; concealed by aril; with faboid split; with the lips of the not twisted; asymmetrical or symmetrical; oblong or faboid split the same color as or lighter colored than the falcate (barely); when asymmetrical with both sutures rest of the hilum and therefore conspicuous; larger than nearly straight; not inflated; flattened or compressed; punctiform; up to 1 mm long; with curved outline; without beak; tapered or short tapered at apex; apex circular or elliptic; apical at apex of radicle tip, subapi- aligned or oblique with longitudinal axis of fruit; short cal to radicle tip, or between cotyledon and radicle lobe; tapered at base; base aligned with longitudinal axis of recessed; within rim or halo. Hilum halo color of testa fruit; with the apex and base uniform in texture; (to duller). Hilum rim color of testa. Lens discernible or coriaceous or ligneous; seed chambers externally not discernible; less than or equal to or greater than 0.5 visible or invisible; with the raised seed chambers not mm in length; up to 1 mm long; with margins straight torulose. Fruit margin not constricted; without sulcus; or curved; circular (with light colored line), elliptic, or plain. Fruit wings absent. Fruit substipitate or oblong; not in groove of raphe; confluent with hilum; nonstipitate. Fruit with all layers dehiscing; splitting mounded or flush; same color as or dissimilar color along sutures. Dehiscence of valves along both sutures; from testa; lighter than testa; reddish tan; not within apical and down; active; with valves twisting. Replum corona, halo, or rim. Endosperm thick; covering entire invisible. Epicarp dull; monochrome; brown to reddish embryo; adnate to embryo. Cotyledons smooth; both brown; pubescent and indurate or pubescent but soon outer faces convex; both the same thickness; both more deciduous; with 1 type of pubescence; pilose or or less of equal length; not folded; margin entire 180 tomentose; with pubescence golden; with pubescence degrees from base of radicle; similar at apex; not uniformly distributed; with simple hairs; pliable; with concealing radicle; split over radicle; with lobes; with hair bases plain; eglandular; without spines; smooth or the interface division terminating at base of radicle; not smooth; with elevated features; veined or not without margins recessed; tan; inner face flat; glabrous veined; reticulately veined; not tuberculate; faintly around base of radicle. Embryonic axis deflexed; wrinkled or pusticulate (minutely); not exfoliating, oblique to length of seed; without a joint evident exfoliating, or exfoliating in part; with or without between the radicle and the cotyledons. Radicle linear cracks; cracking oblique or transverse to fruit length. or bulbose; lobe tip straight; deflexed and parallel to Mesocarp thin; surface not veined; 1-layered; without cotyledon length; centered between cotyledons; balsamic vesicles; without fibers; solid; ligneous (or equaling or exceeding length of cotyledons. Plumule subligneous) or coriaceous. Endocarp dull; mono- rudimentary; glabrous. chrome; reddish brown; smooth or scurfy; nonseptate; chartaceous; not exfoliating; remaining fused to Distribution: Europe, Morocco, and Canary Islands. mesocarp and epicarp; entire. Seeds 4Ð13; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; up to 1 mm long; of 1 length only; flattened or triangular; straight. Aril present or absent; fleshy; topknotlike or cupshaped; covering less than 1/2 of seed; reddish brown or tan (to reddish). Chamaecytisus: C. austriacus (C. Linnaeus) J.H.F. Link (CÐ E), C. spp. (AÐB). A, Fruits (closed and dehisced) Seed 2.7Ð5 ϫ 2.2Ð3.5 ϫ 1.5Ð2.7 mm; not overgrown; (ϫ 1.3); B, seeds (ϫ 4.6); CÐD, testa (ϫ 50, ϫ 1000); angular or not angular; asymmetrical; oblong, ovate, E, embryos (ϫ 8).

1109 Genus: Calicotome J.H.F. Link smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon Phylogenetic Number: 30.17. lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy or dull; not Tribe: Genisteae. modified by a bloom; colored; monochrome; brown to light brown to dark reddish brown, tan (to faintly Species Studied—Species in Genus: 2 spp.—2 spp. greenish tan), or yellow; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe Fruit a legume; unilocular; 3Ð5.5 ϫ 0.7Ð0.8 ϫ 0.27 cm; not visible. Hilum visible; with faboid split; with the with persistent or deciduous calyx; with calyx shorter lips of the faboid split the same color as the rest of the than fruit; without orifice formed by curving of fruit or hilum; punctiform or larger than punctiform; 0.3 mm fruit segments; straight or curved (slightly); not plicate; long; with curved outline; circular; apical at apex of not twisted; asymmetrical or symmetrical; linear or radicle tip or between cotyledon and radicle lobe; oblong; when asymmetrical with both sutures nearly recessed; within rim. Hilum rim color of testa. Lens straight; not inflated; compressed; with beak; straight; discernible; equal to or greater than 0.5 mm in length; with solid beak the same color and texture as fruit; 0.7Ð0.8 mm long; with margins straight or curved; tapered or short tapered at apex; apex aligned or oblong or wedge-shaped; oblong; not in groove of oblique with longitudinal axis of fruit; tapered or short raphe; confluent with hilum; mounded; similar color as tapered at base; base aligned with longitudinal axis of testa; darker than testa; reddish brown; not within fruit; with the apex and base uniform in texture; corona, halo, or rim. Endosperm thick; covering entire ligneous; seed chambers externally visible or invisible; embryo; adnate to embryo. Cotyledons smooth; both with the raised seed chambers not torulose. Fruit outer faces convex; both the same thickness; both more margin not constricted; without sulcus; plain (though or less of equal length; not folded; margin entire 180 upper suture thickened and at most with 2 subwings) or degrees from base of radicle; similar at apex; not embellished. Fruit wings absent or present; 2; 1 mm concealing radicle; split over radicle; with lobes; with wide (at most); sutural; on 1 suture. Fruit nonstipitate or the interface division terminating at base of radicle; substipitate. Fruit with all layers dehiscing; splitting without margins recessed; tan or yellow; inner face flat; along sutures. Dehiscence of valves along both sutures; glabrous around base of radicle. Embryonic axis apical and down; active; with valves twisting. Replum deflexed; oblique to length of seed; without a joint invisible. Epicarp glossy; monochrome; dark reddish to evident between the radicle and the cotyledons. Radicle slightly purplish brown; pubescent and indurate; with 1 bulbose; lobe tip straight; deflexed and parallel to type of pubescence; puberulent or villous; with pubes- cotyledon length; centered between cotyledons; 1/2 to cence golden or gray; with pubescence uniformly nearly or equaling length of cotyledons. Plumule distributed; with simple hairs; pliable; with hair bases rudimentary; glabrous. plain; eglandular; without spines; not smooth; with elevated features; not veined; not tuberculate; faintly Distribution: Mediterranean region. wrinkled; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic Notes: The correct spelling of the genus is Calicotome, not vesicles; without fibers; solid; subligneous. Endocarp Calycotome (Gunn 1983, Gunn et al. 1992). dull; monochrome; brown to purplish brown or purple; smooth and scurfy; nonseptate (with well developed scurfy lines between seeds) or subseptate (because of regular intrusion of fruit); chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 2Ð8; length oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform; straight. Aril present or absent; dry; rim-aril or tongue-aril; tan or white.

Seed 3Ð4.2 ϫ 2.6Ð3.3 ϫ 1Ð3 mm; not overgrown; not Calicotome: C. spinosa (C. Linnaeus) J.H.F. Link (CÐE), C. angular or angular; asymmetrical; rectangular, oblong, spp. (AÐB). A, Fruit and valves (ϫ 1.5); B, seeds elliptic, or circular; compressed or terete; with surface (ϫ 6.1); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

1111 Genus: Erinacea M. Adanson absent. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as Phylogenetic Number: 30.18. the rest of the hilum; punctiform; subapical to radicle tip and between cotyledon and radicle lobe; recessed; Tribe: Genisteae. within halo. Hilum halo color darker (greenish) than testa. Lens discernible; less than 0.5 mm in length; with Species Studied—Species in Genus: 1 sp.—1 sp. margins straight or curved; oblong or linear; not in groove of raphe; confluent with hilum; flush; dissimilar ϫ ϫ Fruit a legume; unilocular; 1.5Ð1.8 0.4 0.1 cm; with color from testa; lighter than testa; tan; not within persistent calyx; with calyx shorter than fruit; without corona, halo, or rim. Endosperm thick; covering entire orifice formed by curving of fruit or fruit segments; embryo; adnate to embryo. Cotyledons smooth; both straight; not plicate; not twisted; asymmetrical or outer faces convex; both the same thickness; both more symmetrical; narrowly oblong; when asymmetrical with or less of equal length; not folded; margin entire 180 both sutures nearly straight; not inflated; compressed; degrees from base of radicle; similar at apex; not with beak; straight; with solid beak the same color and concealing radicle; split over radicle; with lobes; with texture as fruit; tapered at apex; apex aligned with the interface division terminating at base of radicle; longitudinal axis of fruit; tapered at base; base aligned without margins recessed; reddish brown; inner face flat; with longitudinal axis of fruit; with the apex and base glabrous around base of radicle. Embryonic axis uniform in texture; chartaceous; seed chambers deflexed; oblique to length of seed; without a joint externally visible; with the raised seed chambers evident between the radicle and the cotyledons. Radicle torulose. Fruit margin not constricted; without sulcus; bulbose; lobe tip straight; deflexed and parallel to plain. Fruit wings absent. Fruit nonstipitate. Fruit with cotyledon length; not centered between cotyledons all layers dehiscing; splitting along sutures. Dehiscence (radicle outside 1 cotyledon and inside other, therefore of valves along both sutures; apical and down; active; junctions for each cotyledon different); equaling length with valves reflexing. Replum invisible. Epicarp dull; of cotyledons. Plumule rudimentary; glabrous. monochrome; reddish brown or black; pubescent and indurate; with hairs appressed or erect; with 1 type of Distribution: Southwestern Europe and northwestern Africa. pubescence; sericeous; with pubescence golden or gray; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; smooth; not veined; not tuberculate; not exfoliating or exfoliating in part; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; chartaceous. Endocarp dull; monochrome; reddish brown or gray; smooth and scurfy; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð3; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; triangular; straight. Aril dry; rim-aril; reddish tan or white.

Seed 3.7 ϫ 3 ϫ 1.5 mm; not overgrown; not angular; asymmetrical; oblong or triangular; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on Erinacea: E. anthyllis J.H.F. Link (AÐE). A, Fruit within seed faces. Testa not adhering to endocarp; glossy; not calyx and corolla and valve (ϫ 2.9); B, seeds (ϫ 6.2); modified by a bloom; colored; monochrome; dark CÐD, testa (ϫ 50, ϫ 1000); E, embryos (upper two reddish to greenish brown or olive; glabrous; smooth; show the outside of both cotyledon-radicle junctions) (ϫ coriaceous. Fracture lines absent. Rim absent. Wings 10).

1113 Genus: Spartium C. Linnaeus punctiform; subapical to radicle tip to apical at apex of radicle tip; recessed; within rim. Hilum rim color of to Phylogenetic Number: 30.19. darker than testa. Lens discernible; less than 0.5 mm in length; with margins straight or curved; circular or Tribe: Genisteae. oblong; not in groove of raphe; confluent with hilum; mounded; similar color as or dissimilar color from Species Studied—Species in Genus: 1 sp.—1 sp. testa; lighter or darker than testa; tan (reddish within dark reddish border) or black; not within corona, halo, Fruit a legume; unilocular; 5Ð7 ϫ 0.5Ð0.8 ϫ 0.2Ð0.3 cm; or rim. Endosperm thin; covering entire embryo; adnate with deciduous calyx; without orifice formed by to testa. Cotyledons smooth; both outer faces convex; curving of fruit or fruit segments; straight; not plicate; both the same thickness; both more or less of equal not twisted; symmetrical; linear; not inflated; flattened; length; not folded; margin entire 180 degrees from base without or with beak; with solid beak the same color of radicle; similar at apex; not concealing radicle; split and texture as fruit; short tapered at apex; apex aligned over radicle; with lobes; with the interface division with longitudinal axis of fruit; short tapered at base; terminating at base of radicle; without margins re- base aligned with longitudinal axis of fruit; with the cessed; pale tan; inner face flat; glabrous around base of apex and base uniform in texture; coriaceous; seed radicle. Embryonic axis deflexed; oblique to length of chambers externally visible; with the raised seed seed; without a joint evident between the radicle and chambers not torulose. Fruit margin not constricted; the cotyledons. Radicle bulbose; lobe tip straight; without sulcus; plain. Fruit nonstipitate. Fruit with all deflexed and parallel to cotyledon length; centered layers dehiscing; splitting along sutures. Dehiscence of between cotyledons; 1/2 to nearly length of cotyledons. valves along both sutures; apical and down; active; with Plumule rudimentary; glabrous. valves twisting. Replum invisible. Epicarp glossy; monochrome; reddish brown; glabrous; eglandular; Distribution: Mediterranean region and widely cultivated. without spines; not smooth; with elevated features; not veined; not tuberculate; faintly wrinkled; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; coriaceous. Endocarp glossy or dull; monochrome; reddish brown or tan (reddish); smooth and scurfy (somewhat between seeds); subseptate; with septa thin (tissue paper-like), flexible; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 10Ð16; length transverse to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril dry; rim-aril or tongue-aril; cream or tan.

Seed 3.5Ð4 ϫ 2.5Ð3 ϫ 1.5Ð1.7 mm; not overgrown; not angular or angular; asymmetrical; oblong, quadrangular, rectangular, circular, D-shaped, or irregular; compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; colored; monochrome; dark to bright reddish brown or purple (almost dark); glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not Spartium: S. junceum C. Linnaeus (AÐE). A, Dehisced fruit visible. Hilum visible; with faboid split; with the lips of and valve (ϫ 1.4); B, seeds (ϫ 6); CÐD, testa (ϫ 50, the faboid split the same color as the rest of the hilum; ϫ 1000); E, embryos (ϫ 8).

1115 Genus: Gonocytisus É. Spach Rim present. Wings absent. Raphe not visible. Hilum partially or fully concealed; concealed by aril or Phylogenetic Number: 30.20. funiculus; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; Tribe: Genisteae. between cotyledon and radicle lobe; recessed; within rim or not within corona, halo, or rim. Hilum rim color Species Studied—Species in Genus: 2 spp.—3 spp. of testa. Lens discernible; less than 0.5 mm in length; with margins straight or curved; more or less oblong or Fruit a legume; unilocular; 1.2Ð1.7 ϫ 0.45Ð0.6 ϫ 0.075Ð circular; not in groove of raphe; adjacent to hilum; 0.5 0.18 cm; with persistent or deciduous calyx; with calyx mm from hilum; flush or recessed; similar color as or shorter than fruit; without orifice formed by curving of dissimilar color from testa; lighter than testa; tan; not fruit or fruit segments; straight; not plicate; not twisted; within corona, halo, or rim. Endosperm thin; covering asymmetrical or symmetrical; oblong or rhombic; when entire embryo; adnate to testa. Cotyledons smooth; both asymmetrical with both sutures nearly straight; not outer faces convex; both the same thickness; both more inflated; compressed; without beak; short tapered at or less of equal length; not folded; margin entire 180 apex; apex aligned or oblique (slightly) with longitudi- degrees from base of radicle; similar at apex; not nal axis of fruit; short tapered at base; base aligned concealing radicle; split over radicle; with lobes; with (slightly) or oblique with longitudinal axis of fruit; with the interface division terminating at base of radicle; the apex and base uniform in texture; ligneous; seed without margins recessed; reddish brown; inner face chambers externally visible; with the raised seed flat; glabrous around base of radicle. Embryonic axis chambers not torulose. Fruit margin not constricted; deflexed; oblique to length of seed; with a joint evident without sulcus; plain. Fruit wings absent. Fruit between the radicle and the cotyledons. Radicle nonstipitate. Fruit with all layers dehiscing; splitting bulbose; lobe tip straight; deflexed and parallel to along sutures. Dehiscence of valves along both sutures; cotyledon length; centered between cotyledons; less apical and down; passive. Replum invisible. Epicarp than 1/2 length of cotyledons. Plumule moderately dull; monochrome; reddish brown; pubescent and developed; glabrous. indurate; with hairs appressed; with 1 type of pubescence; with pubescence gray; with pubescence uni- Distribution: Eastern Mediterranean. formly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; not smooth; with elevated features; obliquely veined relative to fruit length; not tuberculate; faintly wrinkled; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; subligneous. Endocarp dull; monochrome; brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð2; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; filiform or triangular; straight. Aril dry; rim-aril; tan.

Seed 2.5Ð4 ϫ 2Ð3 ϫ 1.7Ð2 mm; not overgrown; not angular or angular; asymmetrical; oblong, reniform, or circular (more or less); compressed; with surface smooth; with visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Testa not adhering to endocarp; glossy; not modified by a bloom; Gonocytisus: G. angulatus (C. Linnaeus) É. Spach (CÐE), colored; monochrome; dark reddish brown or black; G. spp. (AÐB). A, Fruits (closed and dehisced) (ϫ 3.4); glabrous; smooth; coriaceous. Fracture lines absent. B, seeds (ϫ 5.7); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 8).

1117 Genus: Retama C.S. Rafinesque-Schmaltz partially adhering to endocarp; dull; not modified by a bloom; colored; monochrome or mottled; with frequent Phylogenetic Number: 30.21. or infrequent mottles; dark reddish to greenish brown, tan (to greenish or reddish tan), green (light to dark), or Tribe: Genisteae. black; with black overlay; glabrous; smooth; osseous. Fracture lines absent. Rim present. Wings absent. Species Studied—Species in Genus: 4 spp.—4 spp. Raphe not visible. Hilum visible; with faboid split; with the lips of the faboid split the same color as the rest of Fruit a legume; unilocular; 0.7Ð2 ϫ 0.6Ð1 ϫ 0.45Ð1 cm; the hilum; larger than punctiform; up to 1 mm long; with deciduous calyx; without orifice formed by with straight outline; oblong; marginal according to curving of fruit or fruit segments; straight; not plicate; radicle tip; recessed; within rim. Hilum rim color of or not twisted; asymmetrical or symmetrical; ovate to lighter than testa. Lens discernible or not discernible; elliptic or elliptic to ovate; when asymmetrical with 1 equal to or greater than 0.5 mm in length; up to 2.5 mm straight and 1 curved suture or both sutures parallelly long; with margins straight or curved; linear, oblong, curved; widest near middle or D-shaped; not inflated; wedge-shaped, or circular (with a small satellite); not in terete; without or with beak; declined; with solid beak groove of raphe; confluent with or adjacent to hilum; up the same color and texture as fruit; rounded at apex; to 0.3 mm from hilum; flush or recessed; similar color apex oblique with longitudinal axis of fruit; tapered at as or dissimilar color from testa; lighter or darker than base; base aligned with longitudinal axis of fruit; with testa; reddish brown, tan, or black; not within corona, the apex and base uniform in texture; ligneous; seed halo, or rim. Endosperm thick; covering entire embryo; chambers externally invisible. Fruit margin not con- adnate to testa. Cotyledons smooth; both outer faces stricted; without sulcus; plain. Fruit nonstipitate. Fruit convex; both the same thickness; both more or less of indehiscent (to finally incompletely dehiscent along equal length; not folded; margin entire 180 degrees vertical suture) or with all layers dehiscing; splitting from base of radicle; similar at apex; not concealing along suture. Dehiscence of valves along 1 suture; radicle; split over radicle; with lobes; with the interface probably apical and down; passive. Replum invisible. division terminating at base of radicle; without margins Epicarp glossy or dull; monochrome (though some with recessed; tan; inner face flat; glabrous around base of darker areas); brown to dark or dark reddish brown or radicle. Embryonic axis deflexed; oblique to length of tan; glabrous; eglandular; without spines; not smooth or seed; without a joint evident between the radicle and smooth; with elevated features; not veined; not tubercu- the cotyledons. Radicle bulbose; deflexed and parallel late; rugose or wrinkled; not exfoliating; without to cotyledon length; centered beween cotyledons; less cracks. Mesocarp thick or thin; surface not veined; 1- than 1/2 length of cotyledons. Plumule rudimentary; layered; without balsamic vesicles; without fibers; solid glabrous. (tan) or vitreous (dark reddish-brown); ligneous (or subligneous). Endocarp dull or glossy; monochrome (to Distribution: Mediterranean region, northern Africa, Canary darker and lighter when underlain by dark mesocarp); Islands, and Asia (Middle East). brown or tan; spongy and cobwebby; subseptate or nonseptate; with septa thin (tissue paper-like), flexible Notes: This genus is presented under the name Lygos M. or thicker than paper, firm; with septa eglandular; Adanson in Heywood (1968). coriaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1(Ð2); length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; straight. Aril absent.

Seed 6Ð7.5 ϫ 4Ð5.5 ϫ 3Ð4.5 mm; not overgrown; not angular or angular; asymmetrical or symmetrical; oblong, reniform, triangular (more or less), D-shaped, or irregular; compressed, terete, or mounded on 1 side and straight on other side; with surface smooth; without Retama: R. sphaerocarpa (C. Linnaeus) P.E. Boissier (CÐ visible radicle and cotyledon lobes; without hilar sinus; E), R. spp. (AÐB). A, Fruits (ϫ 3.4); B, seeds (ϫ 4); CÐ without umbo on seed faces. Testa not adhering or D, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 5).

1119 Genus: Genista C. Linnaeus overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long or measured; up to 1.5 mm long; of 1 Phylogenetic Number: 30.22. length only; filiform, thick, or triangular; straight, curved, S-curved, or hooked. Aril dry or fleshy; when Tribe: Genisteae. fleshy topknotlike or cupshaped; covering less than 1/2 of seed; when dry rim-aril or tongue-aril; reddish Species Studied—Species in Genus: 33 spp.—87 spp. brown, tan, white, or red (nearly).

Fruit a legume; unilocular; 0.8Ð3 ϫ 0.35Ð0.4 ϫ 0.15Ð0.3 Seed 1.8Ð4.2 ϫ 1.5Ð3 ϫ 1Ð2 mm; not overgrown; not cm; with persistent calyx; with calyx shorter than fruit; angular or angular; asymmetrical; oblong, ovate, without orifice formed by curving of fruit or fruit reniform, circular (more or less), D-shaped, irregular segments; straight or curved (or slightly curved); not (somewhat), quadrangular, rhombic, or triangular; plicate; not twisted; asymmetrical or symmetrical; compressed or terete; with surface smooth; with or oblong, linear, ovate, rhombic, or falcate; when without visible radicle and cotyledon lobes; without or asymmetrical with both sutures parallelly curved; not with external groove between radicle and cotyledon inflated or inflated; compressed, flattened (rarely), or lobes; with external groove between radicle and terete; without beak; tapered or short tapered at apex; cotyledon lobes same color as testa; without hilar sinus; apex aligned, oblique, or right-angled with longitudinal without umbo on seed faces. Testa not adhering to axis of fruit; tapered or short tapered at base; base endocarp; glossy; not modified by a bloom; colored; aligned, oblique, or right angled with longitudinal axis monochrome or mottled and streaked (sometimes both of fruit; with the apex and base uniform in texture; faint); with frequent mottles; with frequent streaks; coriaceous; seed chambers externally visible or brown to yellowish, greenish or reddish brown, yellow invisible; with the raised seed chambers not torulose or (to brownish to greenish yellow), green, olive, or black; torulose. Fruit margin not constricted; without sulcus; with brown or black overlay; glabrous; smooth; plain. Fruit wings absent. Fruit substipitate or coriaceous. Fracture lines absent. Rim absent. Wings nonstipitate. Fruit with all layers dehiscing or indehis- absent. Raphe not visible. Hilum visible or fully cent; splitting along sutures. Dehiscence of valves along concealed (occasionally); concealed by funiculus or both sutures; apical and down; active; with valves funicular remnant; with faboid split; with the lips of the twisting. Replum invisible. Epicarp glossy or dull; faboid split the same color as the rest of the hilum; monochrome; dark to light reddish brown; glabrous or punctiform or larger than punctiform; 0.3 mm long; pubescent and indurate; with hairs erect or appressed; with curved outline; circular; apical at apex of or with 1 type of pubescence; tomentose or villous; with subapical to radicle tip; recessed; within rim. Hilum rim pubescence golden or gray; with pubescence uniformly color of testa. Lens discernible or not discernible; less distributed or with apical pubescence different from than or equal to or greater than 0.5 mm in length; 0.5 basal pubescence; with apical 1/3Ð1/2 pubescent and mm long; with margins straight or curved; elliptic or basal 1/2Ð2/3 glabrous (G. cephalantha É. Spach); with oblong; not in groove of raphe; adjacent to or confluent simple hairs; pliable; with hair bases plain; eglandular; with hilum; 0.1Ð0.2 mm from hilum; mounded or flush; without spines; not smooth; with elevated features; similar color as testa; not within corona, halo, or rim. veined or not veined; obliquely veined relative to fruit Endosperm thick or thin; covering entire embryo; length or reticulately veined; not tuberculate; wrinkled; adnate to embryo or testa. Cotyledons smooth; both exfoliating in part or not exfoliating; without cracks. outer faces convex; both the same thickness; both more Mesocarp thin; surface not veined; 1-layered; without or less of equal length; not folded; margin entire 180 balsamic vesicles; without fibers; solid; coriaceous. degrees from base of radicle; similar at apex; not Endocarp dull; monochrome; reddish brown; smooth concealing radicle; split over radicle; with lobes; with and scurfy (around seeds); with hairs scattered over the interface division terminating at base of radicle; endocarp (minute except along sutures where much without margins recessed; tan, yellow, or white; inner longer); subseptate or nonseptate; with septa thin (tissue face flat; glabrous around base of radicle. Embryonic paper-like), flexible; with septa eglandular; axis deflexed; oblique to length of seed; without a joint chartaceous; not exfoliating; remaining fused to evident between the radicle and the cotyledons. Radicle mesocarp and epicarp; entire. Seeds 1Ð7; length linear or bulbose; lobe tip straight; deflexed and parallel transverse, oblique, or parallel with fruit length; neither to cotyledon length; centered between cotyledons; 1/2

1120 to nearly, equaling, or exceeding length of cotyledons. Plumule rudimentary; glabrous.

Notes: Gibbs (1966) revised the genus, and Gibbs and Dingwall (1971) revised its segregate Teline F.K. Medikus, now recombined with Genista.

1121 Genista: G. germanica C. Linnaeus (CÐE), G. spp. (AÐB). A, Fruits (dehisced) (ϫ 1.5); B, seeds (ϫ 4); CÐD, testa (ϫ 50, ϫ 1000); E, embryos (ϫ 10).

1123 Genus: Echinospartum (É. Spach) J.P. Fourreau of cuticle); coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Hilum visible; Phylogenetic Number: 30.23. with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform; apical Tribe: Genisteae. at apex of radicle tip; slightly recessed; within rim. Hilum rim color of testa. Lens not discernible. En- Species Studied—Species in Genus: 2 spp.—3 spp. dosperm thick; covering entire embryo; adnate to testa. Cotyledons smooth; both outer faces convex; both the Fruit a legume; unilocular; 1Ð1.5 ϫ 0.3Ð0.5 ϫ 0.1Ð0.2 cm; same thickness; both more or less of equal length; not with persistent calyx; with calyx longer than fruit; folded; margin entire 180 degrees from base of radicle; without orifice formed by curving of fruit or fruit similar at apex; not concealing radicle; split over segments; straight; not plicate; not twisted; asymmetri- radicle; with lobes; with the interface division terminat- cal; oblong; when asymmetrical with 1 straight and 1 ing at base of radicle; without margins recessed; reddish curved suture or both sutures parallelly curved; widest tan; inner face flat; glabrous around base of radicle. near middle or D-shaped; not inflated; compressed; Embryonic axis deflexed; oblique to length of seed; with beak; straight; with solid beak the same color and without a joint evident between the radicle and the texture as fruit; tapered at apex; apex aligned with cotyledons. Radicle somewhat linear; lobe tip straight; longitudinal axis of fruit; tapered at base; base aligned deflexed and parallel to cotyledon length; centered with longitudinal axis of fruit; with the apex and base between cotyledons; 1/2 to nearly length of cotyledons. uniform in texture; ligneous; seed chambers externally Plumule moderately developed; glabrous. invisible. Fruit margin not constricted; without sulcus; embellished. Fruit wings absent. Fruit nonstipitate. Distribution: Southwestern Europe. Fruit with all layers dehiscing; splitting along sutures. Dehiscence of valves along both sutures; apical and Notes: We are following Gunn et al. (1992) for the author of down; active; with valves enrolling. Replum invisible. this genus rather than Bisby (1981). Epicarp dull; monochrome; brown; pubescent and indurate; with 1 type of pubescence; villous; with pubescence golden; with pubescence uniformly distributed; with simple hairs; pliable; with hair bases plain; eglandular; without spines; smooth; not veined; not tuberculate; not exfoliating; without cracks. Mesocarp thick; surface not veined; 1-layered; without balsamic vesicles; without fibers; solid; subligneous. Endocarp glossy; monochrome; reddish tan; smooth and scurfy; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 1 or 2; length oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; curved. Aril dry; rim- aril; black.

Seed 2.5Ð2.7 ϫ 2Ð2.5 ϫ 1.7Ð2 mm; not overgrown; angular or not angular; asymmetrical; oblong, ovate, or D- shaped; compressed or terete; with surface smooth; without visible radicle and cotyledon lobes; without hilar sinus; without umbo on seed faces. Cuticle wrinkled (E. horridum (J.L.M. Vahl) W.H.P. Rothmaler). Testa not adhering to endocarp; dull; not modified by a bloom; colored; mottled and streaked; Echinospartum: E. lusitanicum (C. Linnaeus) W.H.P. with frequent mottles; with frequent streaks; dark Rothmaler (CÐE), E. spp. (AÐB). A, Fruits (dehisced) reddish brown; with black overlay; glabrous; smooth or (ϫ 2.9); B, seeds (ϫ 6.6); CÐD, testa (ϫ 50, ϫ 1000); not smooth; with elevated features; wrinkled (because E, embryos (ϫ 12).

1125 Genus: Stauracanthus J.H.F. Link minute); with frequent mottles; with frequent streaks; reddish to greenish brown or green; with purple Phylogenetic Number: 30.24. overlay; glabrous; smooth; coriaceous. Fracture lines absent. Rim absent. Wings absent. Raphe not visible. Tribe: Genisteae. Hilum visible or fully concealed; concealed by aril; with faboid split; with the lips of the faboid split the Species Studied—Species in Genus: 2 spp.—2 spp. same color as the rest of the hilum; punctiform or larger than punctiform; 0.3 mm long; with straight outline; Fruit a legume; unilocular; 0.8Ð2.5 ϫ 0.5 cm; with persis- oblong; apical at apex of radicle tip and between tent calyx; with calyx shorter than fruit; without orifice cotyledon and radicle lobe; recessed; within rim. Hilum formed by curving of fruit or fruit segments; straight; rim color of testa (or nearly so). Lens discernible; equal not plicate; not twisted; asymmetrical; linear or oblong to or greater than 0.5 mm in length; 0.8 mm long; with (to linear); when asymmetrical with 1 straight and 1 margins straight or curved; oblong; not in groove of curved suture or both sutures nearly straight; widest raphe; confluent with hilum; mounded; similar color as near middle or D-shaped (somewhat); not inflated; testa; darker than testa; reddish brown; not within compressed; without beak; rounded at apex; apex corona, halo, or rim. Endosperm thin; covering entire aligned with longitudinal axis of fruit; short tapered at embryo; adnate to testa. Cotyledons smooth; both outer base; base oblique with longitudinal axis of fruit; with faces convex; both the same thickness; both more or the apex and base uniform in texture; coriaceous; seed less of equal length; not folded; margin entire 180 chambers externally invisible. Fruit margin not con- degrees from base of radicle; similar at apex; not stricted; without sulcus; plain. Fruit wings absent. Fruit concealing radicle; split over radicle; with lobes; with nonstipitate. Fruit with all layers dehiscing; splitting the interface division terminating at base of radicle; along sutures. Dehiscence of valves along both sutures; without margins recessed; tan; inner face flat; glabrous assumed apical and down; passive. Replum invisible. around base of radicle. Embryonic axis deflexed; Epicarp dull; monochrome; reddish brown; glabrous or oblique to length of seed; without a joint evident pubescent but soon deciduous; with hairs appressed; between the radicle and the cotyledons. Radicle with 1 type of pubescence; with pubescence golden bulbose; lobe tip straight; deflexed and parallel to (barely golden, primarily colorless); with pubescence cotyledon length; centered between cotyledons; 1/2 to uniformly distributed; with simple hairs; pliable; with nearly length of cotyledons. Plumule rudimentary; hair bases plain; eglandular; without spines; smooth; glabrous. not veined; not tuberculate; not exfoliating; without cracks. Mesocarp thin; surface not veined; 1-layered; Distribution: Southwestern Europe and northwestern Africa. without balsamic vesicles; without fibers; solid; coriaceous. Endocarp dull; monochrome; reddish brown; smooth; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seeds 2Ð6; length oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus measured; 1 mm long; of 1 length only; flattened; curved. Aril present (but apparently easily knocked off); fleshy; topknotlike; covering less than 1/2 of seed; brown or tan.

Seed 2.8Ð3.5 ϫ 2Ð2.5 ϫ 1.2Ð1.8 mm; not overgrown; not angular or angular; asymmetrical; oblong or rectangular (more or less); compressed; with surface smooth; with visible radicle and cotyledon lobes; without external Stauracanthus: S. boininii (P.B. Webb) G.A. da S.F. groove between radicle and cotyledon lobes; without Sampaio (A), S. genistoides (F.d’A. Brotero) G.A. da hilar sinus; without umbo on seed faces. Testa not S.F. Sampaio (CÐE), S. spp. (B). A, Inner and outer adhering to endocarp; glossy; not modified by a bloom; valve surfaces (ϫ 5.7); B, seeds (ϫ 6.3); CÐD, testa colored; monochrome or streaked and mottled (large or (ϫ 50, ϫ 1000); E, embryos (ϫ 14).

1127 Genus: Ulex C. Linnaeus visible radicle and cotyledon lobes; without or with (faint) external groove between radicle and cotyledon Phylogenetic Number: 30.25. lobes; with external groove between radicle and cotyledon lobes same color as testa; without hilar sinus; Tribe: Genisteae. without umbo on seed faces. Testa not adhering to endocarp; glossy or dull; not modified by a bloom; Species Studied—Species in Genus: 7 spp.—20 spp. colored; monochrome; reddish to greenish brown, tan (greenish), or green; glabrous; smooth; coriaceous. Fruit a legume; unilocular; 0.8Ð2 ϫ 0.4Ð0.45 ϫ 0.14Ð0.15 Fracture lines absent. Rim absent. Wings absent. Raphe cm; with persistent calyx; with calyx shorter than fruit; not visible. Hilum fully concealed; concealed by aril; without orifice formed by curving of fruit or fruit with faboid split; with the lips of the faboid split the segments; straight; not plicate; not twisted; symmetrical same color as the rest of the hilum; larger than puncti- or asymmetrical; linear (oblong), oblong, ovate, or form; 0.5Ð1 mm long; with curved outline; elliptic or rhombic; when asymmetrical with both sutures nearly oval; apical at apex of radical tip, subapical to radicle straight; not inflated; compressed or terete; without or tip, or between cotyledon and radicle lobe; recessed; with beak; straight; with solid beak the same color and within rim. Hilum rim color lighter (greenish-tan) or texture as fruit; tapered at apex; apex aligned with darker (dark brown) than testa. Lens discernible; equal longitudinal axis of fruit; short tapered at base; base to or greater than 0.5 mm in length; 0.7Ð1 mm long; aligned or oblique with longitudinal axis of fruit; with with margins straight or curved; oblong (with or the apex and base uniform in texture; coriaceous, without central groove) or key-hole shaped (more or ligneous, or fragile, thinner than chartaceous like less); not in groove of raphe; confluent with hilum; Trifolium (23.07); seed chambers externally visible or flush or recessed; same or similar color as testa; lighter invisible. Fruit margin not constricted; without sulcus; or darker than testa; reddish brown; not within corona, plain. Fruit wings absent. Fruit substipitate or halo, or rim. Endosperm thick; covering entire embryo; nonstipitate. Fruit with all layers dehiscing; splitting adnate to embryo. Cotyledons smooth; both outer faces along sutures. Dehiscence of valves along both sutures; convex; both the same thickness; both more or less of apical and down; active; with valves twisting. Replum equal length; not folded; margin entire 180 degrees invisible. Epicarp dull; monochrome or multicolored; from base of radicle; similar at apex; not concealing bichrome (upper portion darker than lower portion); radicle; split over radicle; with lobes; with the interface reddish brown; with surface texture uniform or not division terminating at base of radicle; without margins uniform, with patches of different texture not restricted recessed; tan; inner face flat; glabrous around base of to the base and apex; pubescent and indurate; with 1 radicle. Embryonic axis deflexed; oblique to length of type of pubescence; villous; with pubescence more or seed; without a joint evident between the radicle and less golden or gray; with pubescence uniformly the cotyledons. Radicle bulbose; lobe tip straight; distributed; with simple hairs; pliable; with hair bases deflexed and parallel to cotyledon length; centered plain; eglandular; without spines; smooth; not veined; between cotyledons; 1/2 to nearly or equaling length of not tuberculate; not exfoliating; without cracks. cotyledons. Plumule rudimentary or moderately Mesocarp thin; surface not veined; 1-layered; without developed; glabrous. balsamic vesicles; without fibers; solid; coriaceous or ligneous (sub). Endocarp dull; monochrome; reddish Distribution: Native to Western Europe and northern Africa; brown; smooth; nonseptate; chartaceous; not exfoliat- introduced elsewhere. ing; remaining fused to mesocarp and epicarp; entire. Seeds 1Ð6; length parallel with or oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus less than 0.5 mm long; of 1 length only; thick; curved. Aril fleshy or dry; topknotlike; rim-aril; brown, tan, cream, or ivory.

Seed 2Ð3 ϫ 1.3Ð3.5 ϫ 1Ð2.7 mm; not overgrown; not Ulex: U. parviflorus P.A. Pourret de Figeac (CÐE), U. spp. angular or angular; asymmetrical; oblong, ovate, (AÐB). A, Dehisced fruits with one in calyx and bracts rectangular, triangular, circular, or irregular; com- (ϫ 2.6); B, seeds (ϫ 6); CÐD, testa (ϫ 50, ϫ 1000); E, pressed or terete; with surface smooth; with or without embryos (ϫ 10).

#1. Subfamily:/ 1. Caesalpinioideae/ 2. Mimosoideae/ 3. Faboideae/ #2. Phylogenetic number:/ #3. Tribe: / 1. Caesalpinieae/ 2. Cassieae/ 3. Cercideae/ 4. Detarieae/ 5. Amherstieae/ #4. Tribe: / 1. Parkieae/ 2. Mimozygantheae/ 3. Mimoseae/ 4. Acacieae/ 5. Ingeae/ #5. Tribe: / 1. Swartzieae/ 2. Sophoreae/ 3. Dipteryxeae/ 4. Dalbergieae/ 5. Abreae/ 6. Amorpheae/ 7. Millettieae/ 8. Robinieae / 9. Indigofereae/ 10. Phaseoleae/ 11. Desmodieae/ 12. Psoraleeae/ 13. Loteae/ 14. Aeschynomeneae/ 15. Adesmieae/ 16. Galegeae/ 17. Carmichaelieae/ 18. Hedysareae/ 19. Fabeae/ 20. Cicereae/ 21. Trifolieae/ 22. Brongniartieae/ 23. Bossiaeeae/ 24. Mirbelieae/ 25. Podalyrieae/ 26. Hypocalypteae/ 27. Crotalarieae/ 28. Euchresteae/ 29. Thermopsideae/ 30. Genisteae/ #6. Subtribe:/ #7. Group:/

FRUIT CHARACTERS #10. Fruit / 1. internal and external morphology recorded/ 2. external morphology only recorded/ 3. internal morphology only recorded/ 4. morphology not recorded/ #11. Fruit a / 1. legume / 2. loment (or a loment segment) / 3. nutlet / #12. Fruit / 1. unilocular/ 2. bilocular/ #13. Fruit / cm long/ #14. Fruit / cm wide/ #15. Fruit /cm thick/ #16. Fruit / 1. length less than twice as long as width/ 2. 2-9 times longer than wide/ 3. more than 9 times longer than wide/ 4. wider than long/ #17. Fruit with / 1. persistent androecial sheath/ 2. deciduous androecial sheath/ #18. Fruit with / 1. persistent corolla/ 2. deciduous corolla/ #19. Fruit with / 1. keel/ 2. standard/ 3. various petals/ #20. Fruit with / 1. persistent calyx/ 2. deciduous calyx/ #21. Fruit with calyx / 1. longer than fruit/ 2. equal in length to fruit/ 3. shorter than fruit/ #22. Fruit / 1. with orifice formed by curving of fruit or fruit segments <1 or more in either coiled or straight fruit>/ 2. without orifice formed by curving of fruit or fruit segments/ #23. Fruit / 1. 0.5-coiled/

1131 2. 1-coiled/ 3. 1.5-coiled/ 4. 2-coiled/ 5. 3-coiled/ 6. 4-coiled/ 7. 5- to 10-coiled/ 8. contorted/ 9. curved / 10. S-curved/ 11. straight/ #24. Fruit / 1. plicate/ 2. not plicate/ #25. Fruit / 1. twisted/ 2. not twisted/ #26. Fruit / 1. symmetrical/ 2. asymmetrical/ #27. Fruit / 1. C-shaped / 2. circular / 3. coiled / 4. didymous / 5. dolabriform / 6. elliptic / 7. falcate / 8. fusiform / 9. irregularly fusiform / 10. harp-shaped / 11. irregular / 12. lanceolate / 13. linear / 14. moniliform / 15. oblanceolate / 16. obliquely oblanceolate / 17. oblong / 18. obovate / 19. obliquely obovate / 20. ovate / 21. obliquely ovate / 22. quadrangular / 23. rectangular / 24. reniform / 25. rhombic / 26. S-shaped / 27. samaroid / 28. triangular / #28. Fruit when asymmetrical with / 1. 1 straight and 1 curved suture/ 2. both sutures nearly straight/

1132 3. both sutures parallelly curved/ 4. both sutures unequally curved/ #29. Fruit / 1. widest near apex/ 2. widest near middle or D-shaped/ 3. widest at base/ 4. narrowing in several places, resembling Desmodium (9.09) fruit/ 5. narrowest near middle, B-shaped/ 6. narrowing slightly once or twice on one side/ #30. Fruit / 1. inflated / 2. not inflated/ #31. Fruit / 1. compressed/ 2. cruciform/ 3. flattened/ 4. mounded on 1 side and straight on other side/ 5. quadrangular/ 6. subtriangular/ 7. terete / #32. Fruit / 1. with beak/ 2. without beak/ #33. Fruit / 1. straight/ 2. declined/ 3. hooked/ 4. coiled/ #34. Fruit with / 1. papery fragile beak up to 1 cm long/ 2. white-bristle beak up to 3 mm long/ 3. solid beak the same color and texture as fruit/ #35. Fruit / 1. abruptly long acuminate at apex/ 2. blunt at apex/ 3. cordate at apex/ 4. emarginate at apex/ 5. rounded at apex/ 6. long tapered at apex / 7. tapered at apex / 8. short tapered at apex / 9. truncate at apex/ #36. Fruit apex / 1. aligned with longitudinal axis of fruit/ 2. oblique with longitudinal axis of fruit/ 3. right-angled with longitudinal axis of fruit/ 4. almost reaching longitudinal axis of fruit/ 5. exceeding (crossing) longitudinal axis of fruit/ #37. Fruit / 1. cordate at base/ 2. emarginate at base/

1133 3. rounded at base/ 4. long tapered at base / 5. tapered at base / 6. short tapered at base / 7. truncate at base/ #38. Fruit base / 1. aligned with longitudinal axis of fruit/ 2. oblique with longitudinal axis of fruit/ 3. right angled with longitudinal axis of fruit/ #39. Fruit with the apex and base / 1. uniform in texture/ 2. differing in texture/ #40. Fruit / 1. upper 1/2 hardened (coriaceous, leathery, or cartilaginous) and lower 1/2 membraneous/ 2. upper 1/2 inflated and reticulate over seed cavity and lower 1/2 adnate and wrinkled to scurfy over seed cavity/ 3. upper 3/4 barely inflated, reticulate, and pubescent and lower 1/4 not inflated, reticulate, or pubescent/ 4. upper 1/4-2/3 firm and/or pubescent and lower 3/4-1/3 fragile and glabrous/ #41. Fruit / 1. chartaceous/ 2. coriaceous / 3. drupaceous/ 4. fleshy / 5. fragile, thinner than chartaceous, like Trifolium/ 6. leathery/ 7. ligneous / 8. membranous/ #42. Fruit seed chambers externally / 1. visible/ 2. invisible/ #43. Fruit with the raised seed chambers / 1. torulose/ 2. not torulose/ #44. Fruit margin / 1. constricted/ 2. not constricted/ #45. Fruit margin / 1. constricted along both margins/ 2. slightly constricted along both margins/ 3. constricted only on 1 margin/ 4. slightly constricted only on 1 margin/ 5. constricted on 1 margin and slightly constricted on the other margin/ #46. Fruit margin / 1. with sulcus/ 2. without sulcus/ #47. Fruit margin / 1. embellished/ 2. plain/ #48. Fruit margin with /

1134 1. flange(s)/ 2. fringe/ 3. prickles/ 4. ridge(s)/ 5. spines/ 6. thickened sutural areas/ 7. wing(s)/ #49. Fruit wing(s) / 1. present/ 2. absent/ #50. Fruit wing(s) / #51. Fruit wing(s) / mm wide/ #52. Fruit wing(s) / 1. samaroid/ 2. valvular / 3. sutural / 4. continuous wing around fruit/ #53. Fruit wing(s) / 1. apical / 2. basal / #54. Fruit wing(s) on / 1. 1 valve/ 2. both valves/ #55. Fruit wing(s) on / 1. 1 suture/ 2. both sutures/ #56. Fruit / 1. stipitate <5 mm or more long>/ 2. substipitate <0.1-5 mm long>/ 3. nonstipitate / #57. Fruit with the stipe / mm long/

FRUIT DEHISCENCE #58. Fruit / 1. with all layers dehiscing / 2. with epicarp and mesocarp dehiscing and endocarp not dehiscing/ 3. with epicarp and mesocarp breaking near center of valve and endocarp dehiscing along suture/ 4. indehiscent/ #59. Fruit / 1. splitting along suture(s)/ 2. separating from suture(s) and along loment segment margins/ 3. fenestrating, opening by a coiling strip of tissue/ 4. opening by deterioration of delicate strip of tissue along inner suture or both sutures/ 5. with epicarp and mesocarp splitting along suture, endocarp lomented, forming an envelope around each seed, with a flat winglike part/ 6. with epicarp and mesocarp splitting along suture, endocarp entire, forming an envelope around the seeds with flat winglike areas/

1135 #60. Dehiscence of valves along / 1. 1 suture/ 2. both sutures/ #61. Dehiscence of valves / 1. apical and down/ 2. medial and up and down/ 3. basal and up/ #62. Dehiscence of valves / 1. active/ 2. passive/ #63. Dehiscence of valves with valves / 1. breaking/ 2. coiling/ 3. enrolling/ 4. reflexing/ 5. revolute/ 6. twisting/ #64. Replum / 1. visible / 2. invisible / #65. Loment / 1. an intact article / 2. a partial article / #66. Loment / 1. dehiscing along 1 suture/ 2. dehiscing along both sutures/ 3. indehiscent/ #67. Loment segments (articles) / 1. conspicuous/ 2. inconspicuous/ #68. Loment segments (articles) /mm long/ #69. Loment segments widest across

/ 1. seed area/ 2. each end/ #70. Loment segments with / 1. all essentially similar in shape/ 2. upper 1 different shape than middle one(s)/ 3. lower 1 different shape than middle one(s)/ 4. upper 1 different shape than lower 1/ #71. Loment segments

/ 1. circular/ 2. curved/ 3. D-shaped/ 4. elliptic/ 5. hippocrepiform / 6. linear/ 7. oblong/ 8. ovate/ 9. quadrangular/

1136 10. rectangular/ 11. trapezoid/ 12. triangular/

EPICARP #72. Epicarp / 1. dull/ 2. glaucous/ 3. glossy/ 4. semiglossy/ #73. Epicarp / 1. monochrome/ 2. multicolored/ #74. Epicarp / 1. bichrome/ 2. mottled/ 3. streaked/ #75. Epicarp / 1. black/ 2. brown/ 3. gray/ 4. green/ 5. orange/ 6. pink/ 7. purple/ 8. red/ 9. tan/ 10. yellow/ #76. Epicarp with / 1. black overlay/ 2. brown overlay/ 3. gray overlay/ 4. green overlay/ 5. orange overlay/ 6. pink overlay/ 7. purple overlay/ 8. red overlay/ 9. tan overlay/ 10. yellow overlay/ #77. Epicarp mottling color combination / 1. constant/ 2. variable/ #78. Epicarp with / 1. mottling over seed chambers/ 2. without mottling over seed chambers/ #79. Epicarp with surface texture / 1. uniform/ 2. not uniform, with patches of different texture not restricted to the base and apex/ #80. Epicarp / 1. glabrous/

1137 2. glabrate/ 3. pubescent and indurate/ 4. pubescent but soon deciduous/ #81. Epicarp with hairs / 1. erect/ 2. appressed/ #82. Epicarp with / 1. 1 type of pubescence/ 2. 2 types of pubescence/ 3. 3 types of pubescence/ #83. Epicarp / 1. hirsute/ 2. peltate / 3. pilose/ 4. puberulent/ 5. sericeous/ 6. strigose/ 7. tomentose/ 8. velutinous/ 9. villous/ #84. Epicarp with pubescence / 1. black/ 2. brown/ 3. golden/ 4. gray / 5. gray-brown/ 6. red/ 7. tan/ 8. white/ 9. yellow/ #85. Epicarp with / 1. longitudinal bands of lighter and darker brown/ 2. appressed dark brown hairs and scattered erect gray hairs intermixed/ 3. appressed dark brown hairs and scattered erect white hairs intermixed/ 4. long appressed brown hairs and short and shorter white hairs intermixed/ 5. golden glandular hairs and short-pilose reddish-brown intermixed/ 6. erect golden hairs and white hooked or not hooked hairs/ 7. golden hooked hairs and gray plain hairs/ 8. gray and black hairs intermixed/ 9. long and short gray plain-tipped hairs/ 10. gray hairs on valves and golden hairs on sutures/ 11. long and short white hairs intermixed/ 12. long and short yellow hairs intermixed/ 13. long and short golden to brown hairs intermixed/ 14. long white and short golden hairs intermixed/ 15. contiguous areas of golden and white hairs/ #86. Epicarp / 1. with pubescence uniformly distributed/ 2. with apical pubescence different from basal pubescence/

1138 3. denser near sutures, sparse centrally/ #87. Epicarp with / 1. apical 1/4 tomentose and basal 3/4 glabrous/ 2. apical 3/4 tomentose and basal 1/4 glabrous/ 3. apical 1/2 crinkly tomentose and basal 1/2 densely villose with straight hairs/ 4. apical 1/3-1/2 pubescent and basal 1/2-2/3 glabrous/ 5. apical 3/4 glabrous and basal 1/4 pilose/ 6. stipe (only) sericeous / #88. Epicarp with / 1. simple hairs/ 2. glandular hairs/ 3. oleaginous hairs/ 4. complex hairs/ #89. Epicarp with / 1. bristle-like hairs/ 2. plumose hairs/ 3. scales/ 4. setae/ 5. stellate hairs/ 6. T-shaped hairs/ #90. Epicarp / 1. stiff/ 2. pliable/ #91. Epicarp with hair bases / 1. swollen/ 2. plain/ #92. Epicarp / 1. antrorse/ 2. retrorse/ 3. straight / #93. Epicarp / 1. straight at apex/ 2. coiled at apex/ 3. curved at apex/ 4. hooked at apex/ #94. Epicarp / 1. glandular/ 2. eglandular/ #95. Epicarp with glandular / 1. disks/ 2. dots/ 3. hairs/ 4. papillae/ 5. setae/ #96. Epicarp / 1. distributed over entire fruit/ 2. limited to a portion of fruit/ #97. Epicarp / 1. upper 1/4-2/3 glandular and lower 3/4-1/3 eglandular/ 2. upper 1/2 glandular and lower 1/2 eglandular/

1139 3. upper 2/3 glandular and lower 1/3 eglandular/ 4. upper 3/4 glandular and lower 1/4 eglandular/ #98. Epicarp / 1. with spines <= prickles>/ 2. without spines/ #99. Epicarp with spines / 1. persistent/ 2. broken off and their bases evident/ #100. Epicarp with spines / 1. same color as the rest of the fruit/ 2. (or their basal remanent) a different color from the rest of the fruit/ #101. Epicarp / 1. smooth/ 2. not smooth/ #102. Epicarp with / 1. elevated features/ 2. recessed features/ #103. Epicarp / 1. veined/ 2. not veined/ #104. Epicarp / 1. longitudinally veined relative to fruit length/ 2. obliquely veined relative to fruit length/ 3. transversely veined relative to fruit length/ 4. reticulately veined / 5. irregularly veined/ #105. Epicarp / 1. tuberculate/ 2. not tuberculate/ #106. Epicarp with / 1. spongy tubercles on each valve/ 2. solid tubercles on each valve/ #107. Epicarp / 1. blistered/ 2. concentric whorls like a fingerprint/ 3. dotted/ 4. faveolate/ 5. glandular dotted / 6. knobbed/ 7. lenticular/ 8. muricate/ 9. papillose/ 10. pusticulate/ 11. raised reticulate/ 12. ribbed/ 13. rugose/ 14. scaly/ 15. scurfy/ 16. shagreen/ 17. striate/

1140 18. subvesicular/ 19. tessellate/ 20. tuberculate/ 21. verrucose-rugose/ 22. warty/ 23. wrinkled/ #108. Epicarp / 1. glandularly punctate/ 2. grooved/ 3. pitted/ 4. punctate/ 5. slitted obliquely/ #109. Epicarp / 1. exfoliating/ 2. exfoliating in part/ 3. checking/ 4. not exfoliating/ #110. Epicarp / 1. with cracks/ 2. without cracks/ #111. Epicarp cracking / 1. longitudinal to fruit length/ 2. oblique to fruit length/ 3. transverse to fruit length/ 4. irregular/

MESOCARP #112. Mesocarp / 1. present/ 2. absent/ #113. Mesocarp / 1. thick/ 2. thin/ 3. trace/ #114. Mesocarp surface / 1. uniformly veined/ 2. veined over seed chamber and inconspicuously veined on wing/ 3. not veined/ #115. Mesocarp / 1. 1-layered/ 2. 2-layered/ 3. 3-layered/ #116. Mesocarp / 1. with balsamic vesicles/ 2. without balsamic vesicles/ #117. Mesocarp / 1. with fibers/ 2. without fibers/ #118. Mesocarp / 1. with reniform canals/ 2. without reniform canals/

1141 #119. Mesocarp / 1. firm-walled open empty cells/ 2. fibrous throughout / 3. fleshy / 4. glassy beads/ 5. mealy / 6. spongy / 7. vitreous / 8. solid / #120. Mesocarp with fibers / 1. over mealy tissue/ 2. over solid layer/ 3. over spongy tissue/ 4. over vitreous layer/ 5. embedded in mealy tissue over solid layer/ 6. below solid or compacted fibrous layer/ #121. Mesocarp with / 1. empty space (with or without spongy tissue) within solid layer/ 2. honeycomb layer over solid layer/ 3. solid layer over solid layer/ 4. solid layer over spongy layer/ 5. solid layer over vitreous layer/ 6. spongy layer over solid layer/ 7. veins over solid layer/ 8. vitreous layer over solid layer/ #122. Mesocarp with / 1. solid layer over spongy layer over solid layer/ 2. solid layer over 2 distinct spongy layers/ 3. solid layer over 2 distinct solid layers/ 4. solid layer over vitreous layer over solid layer/ 5. spongy layer over vitreous over solid layer/ 6. vitreous layer over 2 distinct solid layers/ #123. Mesocarp / 1. chartaceous/ 2. coriaceous/ 3. fleshy/ 4. ligneous /

ENDOCARP #124. Endocarp / 1. present/ 2. absent/ 3. concealed by adnate testa/ 4. concealed by fleshy mesocarp/ #125. Endocarp / 1. dull/ 2. glossy/ #126. Endocarp / 1. opaque/ 2. translucent/ #127. Endocarp /

1142 1. bichrome/ 2. monochrome/ 3. mottled/ 4. streaked/ #128. Endocarp / 1. black/ 2. brown/ 3. gray/ 4. green/ 5. purple/ 6. orange/ 7. red/ 8. tan/ 9. white/ 10. yellow/ #129. Endocarp with / 1. mottling/ 2. mottling over seed chambers/ 3. mottling above and below seed chambers/ #130. Endocarp with / 1. streaking/ 2. streaking over seed chambers/ 3. streaking above and below seed chambers/ #131. Endocarp with / 1. black overlay/ 2. brown overlay/ 3. gray overlay/ 4. green overlay/ 5. pink overlay/ 6. purple overlay/ 7. red overlay/ 8. tan overlay/ 9. yellow overlay/ #132. Endocarp / 1. cobwebby/ 2. cracked/ 3. fibrous/ 4. floury-filamentous/ 5. hairy/ 6. pithy/ 7. pulpy/ 8. reticulate/ 9. rugose/ 10. scurfy/ 11. smooth/ 12. spongy/ 13. veined/ 14. vitreous/ 15. transversely wrinkled/ #133. Endocarp / 1. with adhering pieces of testa/

1143 2. without adhering pieces of testa/ #134. Endocarp with hairs / 1. restricted to sutures/ 2. in longitudinal rows/ 3. scattered over endocarp/ 4. surrounding seed chambers/ #135. Endocarp / 1. septate/ 2. subseptate/ 3. nonseptate/ #136. Endocarp with septa / 1. thin (tissue paper-like), flexible/ 2. thicker than paper, firm/ 3. composed of minute fringe of hairs/ #137. Endocarp with septa / 1. glandular/ 2. eglandular/ #138. Endocarp / 1. chartaceous/ 2. coriaceous/ 3. ligneous / 4. osseous/ 5. pulpy/ #139. Endocarp / 1. exfoliating/ 2. exfoliating in part/ 3. not exfoliating/ #140. Endocarp / 1. remaining fused to mesocarp and epicarp/ 2. separating from mesocarp/ 3. separating with mesocarp from epicarp/ #141. Endocarp / 1. remaining fused to epicarp/ 2. separating from epicarp/ #142. Endocarp / 1. without wings/ 2. with wing(s) extending into epicarp/ 3. with wing(s) not extending into epicarp/ #143. Endocarp / 1. entire/ 2. separating into 1-seeded winged segments/

SEEDS IN FRUIT #144. Seed(s) / #145. Seed(s) length / 1. parallel with fruit length/ 2. oblique to fruit length/ 3. transverse to fruit length/ #146. Seed(s) / 1. overlapping/ 2. touching/

1144 3. neither overlapping nor touching/ #147. Seed(s) in / 1. 1 series/ 2. 2 or more series/

FUNICULUS #148. Funiculus / 1. / 2. <0.5 mm long or more>/ #149. Funiculus / mm long/ #150. Funiculus of / 1. 1 length only/ 2. 2 different lengths/ #151. Funiculus / 1. filiform/ 2. flattened/ 3. partially filiform and partially thick/ 4. thick/ 5. triangular/ #152. Funiculus / 1. anvil-shaped/ 2. coiled/ 3. contorted/ 4. convoluted/ 5. curved/ 6. S-curved/ 7. hooked/ 8. plicate/ 9. straight/ 10. triangular/

ARIL #153. Aril / 1. present/ 2. absent/ #154. Aril / 1. fleshy/ 2. dry/ #155. Aril when fleshy / 1. annular/ 2. caplike/ 3. cupshaped/ 4. expanded funiculus/ 5. flat from apex to near base/ 6. hippocrepiform rim-aril/ 7. hooked/ 8. horseshoe shaped/ 9. irregular/ 10. knotty/ 11. leaflike and attached to marginal hilum/

1145 12. 2-lipped rim-aril/ 13. marginal around seed/ 14. marginal hilar/ 15. spherical with prominent apical depression/ 16. topknotlike/ #156. Aril / 1. crenate/ 2. entire/ 3. fimbriate/ 4. laciniate/ #157. Aril covering / 1. less than 1/2 of seed/ 2. 1/2 to nearly all of seed/ #158. Aril when dry / 1. caplike/ 2. cupshaped/ 3. hippocrepiform rim-aril/ 4. hooded/ 5. 2-lipped rim-aril/ 6. rim-aril/ 7. partial rim-aril/ 8. tongue-aril/ #159. Aril / 1. crenate/ 2. entire/ 3. fimbriate/ 4. laciniate/ #160. Aril covering / 1. less than 1/2 of seed/ 2. 1/2 to nearly all of seed/ #161. Aril / 1. with tongues (or flap-like) on lips of 2-lipped rim-aril/ 2. without tongue (or flap-like) on lips of 2-lipped rim-aril/ #162. Aril with / 1. 1 tongue or flap on 1 lip of 2-lipped rim-aril/ 2. 2 tongues or flaps, 1 on each lip of 2-lipped rim-aril/ #163. Aril / 1. black/ 2. brown/ 3. cream/ 4. gray/ 5. green/ 6. ivory/ 7. olive/ 8. orange/ 9. pink/ 10. red/ 11. tan/ 12. white/ 13. yellow/

1146 SEED CHARACTERS #164. Seed / 1. internal and external morphology recorded/ 2. external morphology only recorded/ 3. internal morphology only recorded <>/ 4. morphology not scored/ #165. Seed / mm long/ #166. Seed / mm wide/ #167. Seed / mm thick/ #168. Seed / 1. overgrown, 1 seed filling entire fruit cavity/ 2. not overgrown/ #169. Seed / 1. angular/ 2. not angular/ #170. Seed / 1. symmetrical/ 2. asymmetrical/ #171. Seed / 1. bilobed, cicerlike / 2. C-shaped/ 3. circular/ 4. cordate/ 5. D-shaped/ 6. elliptic/ 7. falcate/ 8. hippocrepiform/ 9. irregular/ 10. lanceolate/ 11. linear/ 12. mitaform / 13. oblong/ 14. obovate/ 15. ovate/ 16. pyriform/ 17. quadrangular/ 18. rectangular/ 19. reniform/ 20. rhombic/ 21. samaroid/ 22. trapezoid/ 23. triangular / #172. Seed / 1. terete <1:1 ratio including subterete>/ 2. quadrangular <1:1 ratio>/ 3. compressed / 4. flattened / 5. mounded on 1 side and straight on other side/

1147 #173. Seed with surface / 1. grooved/ 2. ridged/ 3. smooth/ 4. wrinkled/ #174. Seed with grooves / 1. longitudinal/ 2. oblique/ 3. reticulate/ 4. transverse/ #175. Seed / 1. with visible radicle and cotyledon lobes/ 2. without visible radicle and cotyledon lobes/ #176. Seed / 1. with external groove between radicle and cotyledon lobes/ 2. without external groove between radicle and cotyledon lobes/ #177. Seed with external groove between radicle and cotyledon lobes / 1. same color as testa/ 2. lighter in color than testa/ #178. Seed / 1. with deep hilar sinus/ 2. with shallow hilar sinus/ 3. without hilar sinus/ #179. Seed / 1. with umbo on seed faces/ 2. without umbo on seed faces/ #180. Seed with umbo on / 1. 1 face of seed/ 2. both faces of seed/ #181. Seed / 1. with medial ridge on each face / 2. without medial ridge on each face/

CUTICLE #182. Cuticle / 1. exfoliating/ 2. not exfoliating/ #183. Cuticle / 1. inflated/ 2. not inflated/ #184. Cuticle inflated around / 1. hilum/ 2. margin of seed/ #185. Cuticle / 1. wrinkled/ 2. not wrinkled/

TESTA #186. Testa / 1. present/ 2. absent /

1148 #187. Testa / 1. with pieces of adhering epicarp/ 2. without pieces of adhering epicarp/ #188. Testa / 1. partially adhering to endocarp/ 2. not adhering to endocarp/ 3. completely adhering to endocarp/ #189. Testa / 1. free from endocarp/ 2. fused to endocarp, at most a transparent brown tissue/ #190. Testa / 1. dull/ 2. glaucous/ 3. glossy/ 4. velvet/ #191. Testa / 1. not modified by a bloom/ 2. modified by a bloom/ #192. Testa / 1. colored/ 2. clear / #193. Testa / 1. bichrome <2 different colored areas>/ 2. monochrome/ 3. mottled/ 4. streaked/ #194. Testa with / 1. frequent mottles / 2. infrequent mottles <1-3 marks per face>/ #195. Testa with / 1. frequent streaks / 2. infrequent streaks <1-3 marks per face>/ #196. Testa / 1. black/ 2. blue/ 3. brown/ 4. cream/ 5. gray/ 6. green/ 7. ivory/ 8. olive/ 9. orange/ 10. pink/ 11. purple/ 12. red/ 13. scarlet/ 14. tan/ 15. white/ 16. yellow/ #197. Testa with / 1. black overlay/

1149 2. brown overlay/ 3. gray overlay/ 4. green overlay/ 5. orange overlay/ 6. pink overlay/ 7. purple overlay/ 8. red overlay/ 9. tan overlay/ 10. yellow overlay/ #198. Testa / 1. glabrous/ 2. minutely pubescent/ #199. Testa / 1. smooth/ 2. not smooth/ #200. Testa with / 1. elevated features/ 2. recessed features/ #201. Testa / 1. bearing endocarp remnants/ 2. bearing a raised line/ 3. blistered cuticle/ 4. corrugate/ 5. echinate/ 6. papillate/ 7. powdery and not rubbing off/ 8. pustulate/ 9. reticulate/ 10. with 1 longitudinal ridge on each face/ 11. ridged longitudinally more than once/ 12. transversely ridged/ 13. rugose/ 14. shagreen/ 15. tessoroid/ 16. tuberculate/ 17. veined/ 18. verrucose/ 19. warty/ 20. wrinkled/ #202. Testa / 1. concaved/ 2. cracked/ 3. large depressions on each face/ 4. grooved/ 5. pitted with large concatenated pits/ 6. pitted with small separate pits/ 7. pitted with stomata in the bottom of the pits/ 8. punctate/ 9. striate/ #203. Testa / 1. chartaceous/ 2. coriaceous /

1150 3. osseous / #204. Pleurogram >/ 1. present/ 2. absent/ #205. Pleurogram >/ %/ #206. Pseudopleurogram >/ 1. present/ 2. absent/ #207. Pseudopleurogram >/ %/ #208. Fracture lines / 1. present/ 2. absent/ #209. Fracture lines / 1. concentric/ 2. irregular/ 3. longitudinal/ 4. reticulate/ 5. transverse/ #210. Rim / 1. present/ 2. absent/ #211. Rim wing-like / 1. at 1 end of seed/ 2. at both ends of seed/ 3. around seed/ 4. along 1 side of seed/ #212. Wing(s) / 1. present/ 2. absent/ #213. Wing(s) / 1. at 1 end/ 2. at both ends/ 3. continuous around seed/

RAPHE #214. Raphe / 1. visible/ 2. not visible/ #215. Raphe from / 1. hilum through lens to base of seed and terminating/ 2. hilum through lens to base of seed and bifurcating/ 3. hilum to lens/ 4. hilum to near base of seed and terminating/ 5. hilum through lens and base of seed to point opposite hilum/ 6. hilum through lens and terminating before base of seed/ 7. hilum through base of seed and up the other side/ 8. lens to base of seed and terminating/

1151 9. lens to base of seed and bifurcating/ #216. Raphe / 1. bifurcating at base of seed with each arm going up antiraphe side turning (U-shaped) down and approaching bifurcation/ 2. not bifurcating/ #217. Raphe / 1. color of testa/ 2. lighter than testa/ 3. darker than testa/ #218. Raphe / 1. black/ 2. brown/ 3. purple / 4. tan/ #219. Raphe / 1. raised/ 2. flush/ 3. recessed/

HILUM #220. Hilum / 1. present / 2. absent/ #221. Hilum / 1. visible / 2. partially concealed/ 3. fully concealed/ #222. Hilum concealed by / 1. aril/ 2. aril remnant/ 3. funiculus/ 4. funicular remnant/ 5. fusion to endocarp/ 6. radicle lobe/ 7. wing/ #223. Hilum / 1. with faboid split/ 2. without faboid split/ #224. Hilum with the lips of the faboid split / 1. the same color as the rest of the hilum/ 2. lighter colored than the rest of the hilum and therefore conspicuous/ #225. Hilum / 1. punctiform / 2. larger than punctiform/ #226. Hilum / mm long/ #227. Hilum with / 1. angular outline/ 2. curved outline/ 3. straight outline / #228. Hilum /

1152 1. circular/ 2. elliptic/ 3. fusiform/ 4. heart-shaped/ 5. oval/ #229. Hilum / 1. irregular/ 2. triangular/ 3. V-shaped/ 4. wedge-shaped/ #230. Hilum / 1. oblong/ 2. linear/ #231. Hilum / 1. apical at apex of radicle tip / 2. subapical to radicle tip/ 3. apical according to radicle tip but marginal according to seed length/ 4. marginal according to radicle tip / 5. between cotyledon and radicle lobe/ #232. Hilum / 1. raised/ 2. flush/ 3. recessed/ #233. Hilum / 1. within corona/ 2. within halo/ 3. within rim/ 4. not within corona, halo, or rim/ #234. Hilum corona color / 1. of testa/ 2. lighter than testa/ 3. darker than testa/ #235. Hilum halo color / 1. of testa/ 2. lighter than testa/ 3. darker than testa/ #236. Hilum rim color / 1. of testa/ 2. lighter than testa/ 3. darker than testa/

LENS #237. Lens / 1. discernible/ 2. not discernible/ #238. Lens / mm long/ #239. Lens with margins / 1. straight/ 2. curved/ #240. Lens / 1. diamond-shaped/

1153 2. irregular/ 3. linear/ 4. oblong/ 5. rectangular/ 6. rhombic/ 7. square/ 8. triangular/ 9. wedge-shaped/ #241. Lens / 1. circular/ 2. elliptic/ 3. hourglass or dumbbell-shaped/ 4. irregular/ 5. key-hole shaped/ 6. 2 circular mounds separated by groove/ 7. 2 oblong mounds separated by groove/ 8. oblong/ 9. ovate/ 10. punctiform/ #242. Lens / 1. in groove of raphe/ 2. not in groove of raphe/ #243. Lens / 1. confluent with hilum / 2. adjacent to hilum / 3. 180 degrees from hilum/ 4. 270 degrees from hilum/ #244. Lens /mm from hilum/ #245. Lens / 1. mounded/ 2. flush/ 3. recessed/ #246. Lens / 1. same color as testa/ 2. similar color as testa/ 3. dissimilar color from testa/ #247. Lens / 1. lighter than testa/ 2. darker than testa/ #248. Lens / 1. black/ 2. brown / 3. gray/ 4. green/ 5. orange/ 6. purple / 7. red/ 8. tan/ 9. white/ 10. yellow/

1154 #249. Lens / 1. within corona/ 2. within halo/ 3. within rim/ 4. not within corona, halo, or rim/ #250. Lens corona color / 1. of testa/ 2. lighter than testa/ 3. darker than testa/ #251. Lens halo color / 1. of testa/ 2. lighter than testa/ 3. darker than testa/ #252. Lens rim color / 1. of testa/ 2. lighter than testa/ 3. darker than testa/

ENDOSPERM #253. Endosperm / 1. present/ 2. absent/ #254. Endosperm / 1. thick/ 2. thin/ 3. trace/ #255. Endosperm / 1. pluglike and resembling tip of radicle/ 2. not pluglike and not resembling tip of radicle/ #256. Endosperm / 1. covering entire embryo/ 2. covering at least 1/2 of embryo, but not entire embryo/ 3. restricted to region of embryo/ #257. Endosperm adnate to / 1. testa/ 2. embryo /

COTYLEDONS #258. Cotyledons / 1. smooth/ 2. not smooth/ #259. Cotyledons / 1. convoluted/ 2. dimpled once/ 3. dotted/ 4. glandular dotted (yellow latex-like substance inside)/ 5. 1-3 grooves on each face/ 6. 4-6 grooves on each face/ 7. 5-7-branched grooves (from veins of testa) on each face/ 8. with midrib/ 9. pitted/

1155 10. glandular pitted/ 11. glandular punctate/ 12. rugose/ 13. ruminate/ 14. sulcate/ 15. veined/ 16. wrinkled/ #260. Cotyledons / 1. outer face of one cotyledon flat and other cotyledon convex/ 2. outer face of one cotyledon flat and other cotyledon concave/ 3. both outer faces convex/ 4. outer face of one cotyledon concave and other cotyledon convex/ 5. both outer faces flat/ 6. with 2 outer faces on each cotyledon, one flat and the other convex/ #261. Cotyledons / 1. one thicker than the other/ 2. both the same thickness/ #262. Cotyledons / 1. 1 longer than other/ 2. both more or less of equal length <>/ #263. Cotyledons / 1. with only 1 folded/ 2. with both folded/ 3. not folded/ #264. Cotyledons / 1. sufficiently folded for inner face to touch itself/ 2. not sufficiently folded for inner face to touch itself/ #265. Cotyledons portions of inner folded face / 1. equal/ 2. unequal/ #266. Cotyledons margin / 1. entire 180 degrees from base of radicle/ 2. not entire 180 degrees from base of radicle/ #267. Cotyledons / 1. bearing flaps/ 2. notched/ 3. wavy/ #268. Cotyledons / 1. similar at apex/ 2. differing at apex (1 concealed by overarching radicle and other auriculate and concealing radicle)/ #269. Cotyledons / 1. completely concealing radicle/ 2. partially concealing radicle/ 3. not concealing radicle/ #270. Cotyledons / 1. entire over radicle/ 2. notched at radicle/ 3. split over radicle/ 4. 1 cotyledon scooped out to accommodate plicate radicle and other cotyledon entire/

1156 #271. Cotyledons / 1. with lobes/ 2. without lobes/ #272. Cotyledons with lobes / 1. overlapping/ 2. touching (auriculate)/ 3. not touching/ #273. Cotyledons / 1. with basal groin formed by lobes/ 2. without basal groin formed by lobes/ #274. Cotyledons with the interface division terminating / 1. at base of radicle/ 2. in radicle tissue/ #275. Cotyledons / 1. with margin(s) recessed/ 2. without margins recessed/ #276. Cotyledons with / 1. 1 margin recessed/ 2. both margins recessed / #277. Cotyledons recessed on / 1. same side as hilum (terminal radicle)/ 2. same side as radicle/ 3. side opposite from radicle/ #278. Cotyledons / 1. brown/ 2. green/ 3. orange/ 4. pink/ 5. red/ 6. tan/ 7. white/ 8. yellow/ #279. Cotyledons inner face / 1. flat/ 2. concave/ 3. wavy/ 4. with central ridge on 1 and central groove on other/ 5. wrinkled/ 6. glandular dotted (with yellow latex-like substance inside)/ #280. Cotyledons / 1. glabrous around base of radicle/ 2. pubescent around base of radicle/

EMBRYONIC AXIS #281. Embryonic axis / 1. deflexed / 2. oblique / 3. parallel / 4. right angled / 5. S-curved / 6. straight /

1157 #282. Embryonic axis / 1. oblique to length of seed/ 2. parallel to length of seed/ 3. perpendicular to length of seed/ 4. plicate to length of seed/ #283. Embryonic axis / 1. with a joint evident between the radicle and the cotyledons/ 2. without a joint evident between the radicle and the cotyledons/

RADICLE #284. Radicle / 1. differentiated from cotyledon/ 2. not differentiated from cotyledon/ #285. Radicle / 1. bulbose/ 2. linear/ 3. triangular/ 4. truncate/ #286. Radicle lobe tip / 1. straight/ 2. curved/ 3. hooked/ #287. Radicle / 1. deflexed and parallel to cotyledon length/ 2. deflexed and parallel to cotyledon width/ 3. oblique to cotyledons/ 4. with 90 degree turn/ 5. with 180 degree turn/ 6. with 360 degree turn/ 7. straight with embryonic axis / #288. Radicle / 1. centered between cotyledons/ 2. not centered between cotyledons (radicle outside 1 cotyledon and inside other, therefore junctions for each cotyledon different)/ #289. Radicle / 1. less than 1/2 length of cotyledons/ 2. 1/2 to nearly length of cotyledons/ 3. equaling length of cotyledons/ 4. exceeding length of cotyledons/

PLUMULE #290. Plumule / 1. rudimentary/ 2. moderately developed/ 3. well developed/ #291. Plumule / 1. glabrous/ 2. pubescent/

DISTRIBUTION #292. / #293. Distribution in world /

1158 1. New World/ 2. Old World/ 3. pantropical/ 4. pansubtropical/ 5. pan warm temperate/ #294. Distribution in world / 1. worldwide crop/ 2. New World crop/ 3. Old World crop/ #295. New World / 1. Canada/ 2. United States / 3. Mexico/ 4. West Indies/ 5. Central America/ 6. South America/ #296. Old World / 1. Europe/ 2. Mediterranean/ 3. Russia/ 4. Asia/ 5. China/ 6. Japan/ 7. Africa/ 8. Madagascar/ 9. Indian Ocean/ 10. India/ 11. Indochina/ 12. Indonesia/ 13. Australia/ 14. New Zealand/ 15. Pacific/ 16. New Guinea/ 17. Fiji/ 18. Hawaii/ 19. Korea/

NOTES #297. /

LEGENDS #298. / #299. / #300. / #301. / #302. / #303. /

1159 Literature Cited

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Ohashi, H. 1982a. The taxonomic position of Tadehagi Pandey, A.K., and Jha, S.S. 1989. Seed structure in rodgeri (Leguminosae-Desmodieae). J. Jap. Bot. 57(9):8Ð Aeschynomene (Fabaceae). J. Jap. Bot. 64:108Ð116. 12. Pankhurst, R.J. 1988. An interactive program for the Ohashi, H. 1982b. The taxonomic position of the genus construction of identification keys. Taxon 37(3):747Ð755. Murtonia (Leguminosae). J. Jap. Bot. 57:225Ð231. Pankhurst, R.J. 1991. Practical Taxonomic Computing. Ohashi, H. 1988. Desmodium (Leguminosae) of the Cambridge University Press, Cambridge, England. Hawaiian islands. Sci. Rep. Taihoku Univ. 39:171Ð189. Pankhurst, R.J. 1995. PANKEY - Programs for the Identifi- Ohashi, H. 1997a. Aphyllodium (DC.) Gagnep. instead of cation and Descriptions of Plants and Animals. Exeter Dicerma DC. (Leguminosae-Papilionoideae: Desmodieae). Software, Setauket, NY. Taiwania 42(2):142Ð148. Pauwels, L. 1983. Révision du genre africain Eminia Taub. Ohashi, H. 1997b. The taxonomic position of Dendrolobium (Fabaceae). Bull. Jard. Bot. État. 53(1/2):153–160. cumingianum Benth. (Leguminosae-Papilionoideae: Desmodieae). Taiwania 42(2):149Ð153. Peltier, M.A.G. 1970. Singularites de las graine et de la plantulé chez Phylloxylon ensifolium H. Baill. Ohashi, H. 1997c. Three species of Dendrolobium in (Papilionacées). Adansonia 10(Ser.2):533–535. Malesia, Australia and Fiji (Leguminosae-Papilionoideae: Desmodieae). Taiwania 42(2):135Ð141. Peltier, M.A.G. 1977. Genus Disynstemon. Adansonia (Ser. 2)17:201Ð203. Ohashi, H., R.M. Polhill, and B.G. Schubert. 1981. Tribe 9. Desmodieae (Benth.) Hutch. (1964). In R.M. Polhill and Phillips, L.L. 1955. A revision of the perennial species of P.H. Raven, eds., Advances in Legume Systematics, vol. 2, Lupinus of North America. Res. Stud. State Coll. Wash. pt. 1, pp. 292Ð300. Proc. Int. Legume Conf., 1978, Kew, 23:161Ð224. England. Minis. Agric., Fisheries and Food, Richmond, England. Pignatti, S. 1982. Flora d’Italia. Volume I. Edagricole, Bologna, Italy. Ohashi, H., T. Nemoto, and T.-L. Wu. 1996. The taxonomic position of Trifidacanthus (Leguminosae). J. Jap. Bot. Pittier, H. 1917. The middle American species of 71:57Ð66. Lonchocarpus. Contr. U.S. Natl. Herb. 20:37Ð93.

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Oliveira, M. de L.A.A. de. 1990. Adições para gênero Plitmann, U. 1981. Evolutionary history of the Old World Desmodium Desvaux (Leguminosae: Faboideae) no Rio lupines. Taxon 30:430Ð437. Grande do Sul, Brasil. Iheringia, Bot. 40:77Ð87. Podlech, D. 1983. Zur taxonomie und nomenklature der Ott, E. 1978. Revision der Sekt. Chronopus Bge. der tragacanthoiden astragali. Mitt. Bot. Staatssamml. München gattung Astragalus L. Phanerog. Monogr. 9:1Ð142. 19:1Ð23.

1174 Podlech, D. 1984. Revision von Astragalus L. Sect. Polhill, R.M. 1981e. Tribe 5. Abreae (Wight & Arn.) Hutch. Herpocaulos Bunge. Mitt. Bot. Staatssamml. München (1964). In R.M. Polhill and P.H. Raven, eds., Advances in 20:441Ð449. Legume Systematics, vol. 2, pt. 1, pp. 243Ð244. Proc. Int. Legume Conf., 1978, Kew, England. Minis. Agric., Fisher- Podlech, D. 1988. Revision von Astragalus L. Sect. Caprini ies and Food, Richmond, England. DC. (Leguminosae). Mitt. Bot. Staatssamml. München 25:1Ð924. Polhill, R.M. 1981f. Tribe 8. Indigofereae (Benth.) Rydb. (1923). In R.M. Polhill and P.H. Raven, eds., Advances in Podlech, D. 1990. Revision von Astragalus L. sect. Legume Systematics, vol. 2, pt. 1, pp. 289Ð291. Proc. Int. Platyglottis Bunge (Leguminosae). Mitt. Bot. Staatssamml. Legume Conf., 1978, Kew, England. Minis. Agric., Fisher- München 29:541–572. ies and Food, Richmond, England.

Podlech, D. 1993. Revision von Astragalus L. Sektion Polhill, R.M. 1981g. Tribe 15. Adesmieae (Benth.) Hutch. Eremophysa Bunge und Eremophysopsis Gontsch. (1964). In R.M. Polhill and P.H. Raven, eds., Advances in (Leguminosae). Sendtnera 1:45Ð64. Legume Systematics, vol. 2, pt. 1, pp. 355Ð356. Proc. Int. Legume Conf., 1978, Kew, England. Minis. Agric., Fisher- Podlech, D., and E. Kozik. 1983. Revision von Astragalus ies and Food, Richmond, England. L. Sektion Theiochrus Bunge. Mitt. Bot. Staatssamml. München 19:351–362. Polhill, R.M. 1981h. Tribe 16. Galegeae (Bronn) Torrey & Gray (1838). In R.M. Polhill and P.H. Raven, eds., Ad- Polhill, R.M. 1968. Argyrolobium Eckl. & Zeyh. vances in Legume Systematics, vol. 2, pt. 1, pp. 357Ð363. (Leguminosae) in tropical Africa. Kew Bull. 22:145Ð168. Proc. Int. Legume Conf., 1978, Kew, England. Minis. Agric., Fisheries and Food, Richmond, England. Polhill, R.M. 1974. A revision of Pearsonia (Leguminosae: Papilionoideae). Kew Bull. 29:383Ð410. Polhill, R.M. 1981i. Tribe 17. Carmichaelieae Hutch. (1964). In R.M. Polhill and P.H. Raven, eds., Advances in Polhill, R.M. 1976. Genisteae (Adans.) Benth. and related Legume Systematics, vol. 2, pt. 1, pp. 364Ð366. Proc. Int. tribes (Leguminosae). Bot. Syst. 1:143Ð368. Legume Conf., 1978, Kew, England. Minis. Agric., Fisher- ies and Food, Richmond, England. Polhill, R.M. 1981a. Papilionoideae. In R.M. Polhill and P.H. Raven, eds., Advances in Legume Systematics, vol. 2, Polhill, R.M. 1981j. Tribe 18. Hedysareae DC. (1825). In pt. 1, pp. 191Ð208. Proc. Int. Legume Conf., 1978, Kew, R.M. Polhill and P.H. Raven, eds., Advances in Legume England. Minis. Agric., Fisheries and Food, Richmond, Systematics, vol. 2, pt. 1, pp. 367Ð370. Proc. Int. Legume England. Conf., 1978, Kew, England. Minis. Agric., Fisheries and Food, Richmond, England. Polhill, R.M. 1981b. Tribe 2. Sophoreae Sprengel (1818). In R.M. Polhill and P.H. Raven, eds., Advances in Legume Polhill, R.M. 1981k. Tribe 19. Loteae DC. (1825). In R.M. Systematics, vol. 2, pt. 1, pp. 213Ð230. Proc. Int. Legume Polhill and P.H. Raven, eds., Advances in Legume System- Conf., 1978, Kew, England. Minis. Agric., Fisheries and atics, vol. 2, pt. 1, pp. 371Ð374. Proc. Int. Legume Conf., Food, Richmond, England. 1978, Kew, England. Minis. Agric., Fisheries and Food, Richmond, England. Polhill, R.M. 1981c. Tribe 3. Dipteryxeae Polhill. In R.M. Polhill and P.H. Raven, eds., Advances in Legume System- Polhill, R.M. 1981l. Tribe 20. Coronilleae (Adans.) Boiss. atics, vol. 2, pt. 1, pp. 231Ð232. Proc. Int. Legume Conf., (1872). In R.M. Polhill and P.H. Raven, eds., Advances in 1978, Kew, England. Minis. Agric., Fisheries and Food, Legume Systematics, vol. 2, pt. 1, pp. 375. Proc. Int. Richmond, England. Legume Conf., 1978, Kew, England. Minis. Agric., Fisher- ies and Food, Richmond, England. Polhill, R.M. 1981d. Tribe 4. Dalbergieae Bronn ex DC. (1825). In R.M. Polhill and P.H. Raven, eds., Advances in Legume Systematics, vol. 2, pt. 1, pp. 233Ð242. Proc. Int. Legume Conf., 1978, Kew, England. Minis. Agric., Fisher- ies and Food, Richmond, England.

1175 Polhill, R.M. 1981m. Tribe 25. Mirbelieae (Benth.) Polhill, Polhill, R.M., and M. Sousa. 1981. Tribe 7. Robinieae stat. nov. In R.M. Polhill and P.H. Raven, eds., Advances in (Benth.) Hutch. (1964). In R.M. Polhill and P.H. Raven, Legume Systematics, vol. 2, pt. 1, pp. 391Ð392. Proc. Int. eds., Advances in Legume Systematics, vol. 2, pt. 1, pp. Legume Conf., 1978, Kew, England. Minis. Agric., Fisher- 283Ð288. Proc. Int. Legume Conf.,1978, Kew, England. ies and Food, Richmond, England. Minis. Agric., Fisheries and Food, Richmond, England.

Polhill, R.M. 1981n. Tribe 26. Bossiaeeae (Benth.) Hutch. Porter, C.L. 1956. The genus Peteria (Leguminosae). (1964). In R.M. Polhill and P.H. Raven, eds., Advances in Rhodora 58:344Ð354. Legume Systematics, vol. 2, pt. 1, pp. 393Ð395. Proc. Int. Legume Conf., 1978, Kew, England. Minis. Agric., Fisher- Potokina, E.K. 1997. Vicia sativa L. aggregate (Fabaceae) in ies and Food, Richmond, England. the flora of former USSR. Genet. Resources Crop Evol. 44:199Ð209. Polhill, R.M. 1981o. Tribe 27. Podalyrieae Benth. (1837). In R.M. Polhill and P.H. Raven, eds., Advances in Legume Potter, D. 1992. Economic botany of Sphenostylis Systematics, vol. 2, pt. 1, pp. 396Ð397. Proc. Int. Legume (Leguminosae). Econ. Bot. 46(3):262Ð275. Conf., 1978, Kew, England. Minis. Agric., Fisheries and Food, Richmond, England. Potter, D., and J.J. Doyle. 1994. Phylogeny and systematics of Sphenostylis and Nesphostylis (Leguminosae: Polhill, R.M. 1981p. Tribe 28. Liparieae (Benth.) Hutch. Phaseoleae) based on morphological and chloroplast DNA (1964). In R.M. Polhill and P.H. Raven, eds., Advances in data. Syst. Bot. 19(3):389Ð406. Legume Systematics, vol. 2, pt. 1, p. 398. Proc. Int. Legume Conf., 1978, Kew, England. Minis. Agric., Fisheries and Pramanik, A., and Thothathri, K. 1986. On the status of Food, Richmond, England. Desmodiastrum Prain (Fabaceae). J. Indian Bot. Soc. 65:373Ð379. Polhill, R.M. 1981q. Tribe 29. Crotalarieae (Benth.) Hutch. (1984). In R.M. Polhill and P.H. Raven, eds., Advances in Purseglove, J.W. 1968. Tropical Crops, Dicotyledons 1. Legume Systematics, vol. 2, pt. 1, pp. 399Ð402. Proc. Int. Longmans, Green and Co., London. Legume Conf., 1978, Kew, England. Minis. Agric., Fisher- ies and Food, Richmond, England. Ramírez, N. 1995. Revisión taxonómica del género Alexa Moq. (Fabaceae, Sophoreae). Ann. Missouri Bot. Gard. Polhill, R.M. 1982. Crotalaria in Africa and Madagascar. 82:549Ð569. A.A. Balkema, Rotterdam, Netherlands. Rauschert, S. 1982. Nomina nova generica et combinationes Polhill, R.M. 1994a. Classification of the Leguminosae. In novae spermatophytorum et pteridophytorum. Taxon F.A. Bisby, J. Buckingham, and J.B. Harborne, eds., 31(3):554Ð563. Phytochemical Dictionary of the Leguminosae, pp. xxxvÐ xlviii. Chapman & Hall, London. Reer, U., and D. Podlech. 1986. Die Europäischen vertreter des gattung Astracantha Podl. (Leguminosae). Mitt. Bot. Polhill, R.M. 1994b. Complete synopsis of legume genera. Staatssamml. München 22:513–569. In F.A. Bisby, J. Buckingham, and J.B. Harborne, eds., Phytochemical Dictionary of the Leguminosae, pp. xlixÐ Reis, M.S., and P.S. Martins. 1989a. Caracterização lvii. Chapman & Hall, London. morfológica do fruto de espécies de Stylosanthes e controle de produção mensal de sementes. Revista Ceres 36:336– Polhill, R.M. 1997. Introduction to the Leguminosae, 347. legumes in street and gardens. Bot. Mag. 14(4):176Ð183. Reis, M.S., and P.S. Martins. 1989b. Efeito do dimorfismo Polhill, R.M., and P.H. Raven, eds. 1981. Advances in do fruto de espécies de Stylosanthes sobre a germinação das Legume Systematics, vol. 2, pt. 1. Internatl. Legume Conf., sementes. Revista Ceres 36:347Ð356. Kew, Proc. 1978. Minis. Agric., Fisheries and Food, Richmond, England. Resslar, P.M. 1980. A review of the nomenclature of the genus Arachis L. Euphytica 29:813Ð817.

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1184 Scientific Name Index

Abrus M. Adanson (5.01) ...... 270 Afgekia W.G. Craib (Millettieae) ...... 292 A. aureus R. Viguier (not illustrated) A. sericea W.G. Craib (AÐE, fruits, seeds, testa, A. canescens J.G. Baker (not illustrated) embryos) A. fruticulosus N. Wallich ex R. Wight & G.A.W. Arnott (not illustrated) Aganope F.A.W. Miquel (Millettieae) ...... 296 A. precatorius C. Linnaeus (CÐE, testa, embryos) A. impressa (S.T. Dunn) R.M. Polhill (CÐE, testa, embryos) Acanthoscelides obtectus, a bruchid, see Phaseolus, p. 606 Airyantha R.K. Brummitt (2.19) ...... 164 Acmispon C.S. Rafinesque-Schmaltz (13.07A)...... 734 A. borneensis (D. Oliver) R.K. Brummitt (CÐE, A. americanus (T. Nuttall) P.A. Rydberg (AÐB, testa, embryos) fruits) Aldina S.F.L. Endlicher (1.04) ...... 92 Acosmium H.W. Schott (2.01) ...... 118 A. latifolia R. Spruce ex G. Bentham (AÐE fruits, A. dasycarpum (J.R.T. Vogel) G.P. Yakovlev (not seed, embryos, embryo magnified) illustrated) A. nitens (J.R.T. Vogel) G. Bentham (CÐE, testa, Alexa C.H.B.A. Moquin-Tandon (2.12A) ...... 144 embryos) A. confusa H. Pittier (CÐE, testa, embryos)

Adenocarpus A.-P. de Candolle (30.05) ...... 1084 Alhagi F. Gagnepain (16.21) ...... 862 A. decorticans P.E. Boissier (CÐE, testa, embryos) A. maurorum F.C. Medikus (CÐE, testa, embryos) A. hispanicus (J.P.A.P.M. de Lamarck) A.-P. de. Candolle (not illustrated) Alistilus N.E. Brown (10.62) ...... 578 A. bechuanicus N.E. Brown (A, fruit) Adenodolichos H.A.T. Harms (10.83) ...... 634 A. magnificus B. Verdcourt (not illustrated) A. punctatus (M. Micheli) H.A.T. Harms (CÐE, testa, embryos) Almaleea M.D. Crisp & P.H. Weston (24.24) ...... 1002 A. subumbellata (W.D. Hooker) M.D. Crisp & Adesmia A.-P. de Candolle (15.01) ...... 810 P.E. Weston (CÐE, testa, embryos) A. aromatica A. Burkart (H, plumose fruit hair) A. bicolor (J.L.M. Poiret) A.-P. de Candolle (not Alophotropis (H.F. Jaubert & É. Spach) A.A. Grossheim, illustrated) see Vavilovia, p. 908 A. capitellata A. Burkart (G, simple fruit hairs) A. incana E. de Vogel (CÐE, testa, embryos) Alysicarpus A.N. Desvaux (11.18) ...... 676 A. lihuelensis A. Burkart (F, bristlelike fruit hair) A. rugosus (C.L. von Willdenow) A.-P. de Candolle (CÐE, testa, embryos) Aenictophyton A. Lee (23.10) ...... 958 A. vaginalis (C. Linnaeus) A.-P. de Candolle A. reconditum A.T. Lee (AÐE, fruits, seeds in situ, (not illustrated) testa, embryos) Amburana C.A.W. Schwacke & P.H.W. Taubert Aeschynomene C. Linnaeus (14.09) ...... 772 (1.15) ...... 116 A. aspera C. Linnaeus (not illustrated) A. cearensis (F.F. Allemäo e Cysneiro) A.C. Smith A. fluminensis J. Velloso de Miranda (not (BÐE, seeds, testa, embryos) illustrated) A. indica C. Linnaeus (not illustrated) Amicia K.S. Kunth (14.19) ...... 792 A. parviflora M. Micheli (not illustrated) A. medicaginea A.H.R. Grisebach (BÐF, fruits, A. virginica (C. Linnaeus) N.L. Britton, E.E., seed, testa, embryos) Sterns, & J.F. Poggenberg (CÐE, testa, A. zygomeris A.-P. de Candolle (not illustrated) embryos)

1185 Ammodendron F.E.L. von Fischer ex A.-P. de Candolle Angylocalyx P.H.W. Taubert (2.13) ...... 146 (2.46) ...... 218 A. oligophyllus (J.G. Baker) E.G. Baker A. karelinii F.E.L. von Fischer & C.A. von Meyer (C, embryos) var. conollyi (A.A. von Bunge) G.P. Yakovlev (CÐE, testa, embryos) Antheroporum F. Gagnepain (Millettieae) ...... 298 A. pierrei F. Gagnepain (AÐG, fruit, seed, testa, Ammopiptanthus S.F. Cheng (29.01) ...... 1058 embryos, cotyledon with glands, magnification A. mongolicus (C.J. Maximowicz) S.-H. Cheng of inner cotyledon surface with glands) (CÐE, testa, embryos) Anthyllis C. Linnaeus (13.02) ...... 718 Amorpha C. Linnaeus (6.04) ...... 280 A. hermanniae C. Linnaeus (CÐE, testa, embryos) A. croceolanata P.W. Watson (CÐE, testa, embryos) A. tetraphylla C. Linnaeus (not illustrated) A. vulneraria C. Linnaeus (not illustrated) Amphicarpaea S. Elliott ex T. Nuttall (10.44) ...... 536 A. bracteata (C. Linnaeus) M.L. Fernald (CÐE, Antopetitia A. Richard (13.16) ...... 750 testa, embryos) A. abyssinica A. Richard (AÐE, fruits and fruit segments, seeds, testa, embryos) Amphimas J.B.L. Pierre ex H.A.T. Harms (2.11) ...... 140 A. pterocarpoides H.A.T. Harms (CÐE, testa, Aotus J.E. Smith (24.21) ...... 996 embryos) A. ericoides (É.P. Ventenat) G. Don (CÐE, testa embryos) Amphithalea C.F. Ecklon & C.L.P. Zeyher (25.02) ...... 1012 A. phyliocides G. Bentham (not illustrated), A. cuneifolia C.F. Ecklon & C.L.P. Zeyher (BÐE, see Gompholobium, p. 960 seed, testa, embryos) A. ericifolia (C. Linnaeus) C.F. Ecklon & J.M. Aphyllodium (A.-P. de Candolle) F. Gagnepain Zeyher (not illustrated) (11.08) ...... 650 A. biarticulatum (C. Linnaeus) F. Gagnepain. Anagyris C. Linnaeus (29.03) ...... 1062 (AÐE, fruits, seeds, testa, embryos) A. foetida C. Linnaeus (AÐE, fruits, seeds, testa, embryos) Apios P.C. Fabricius (10.07)...... 450 A. latifolia P.M.A. Broussonet ex C.L. A. americana F.C. Medikus (A, CÐE, fruits, testa, von Willdenow (not illustrated) embryos)

Anarthrophyllum G. Bentham (30.06)...... 1086 Apoplanesia C.B. Presl (6.01) ...... 274 A. rigidum (J. Gilles ex W.J. Hooker & G.A.W. A. paniculata C.B. Presl (AÐE, fruits with Arnott) G.H.E.W. Hieronymus (CÐE, testa, star-shaped calyx, seeds, testa, embryos, embryos) hilum-lens view)

Andira A.L. de Jussieu (4.04) ...... 234 Apurimacia H.A.T. Harms (Millettieae) ...... 300 A. fraxinifolia G. Bentham (CÐE, testa, embryos) A. michelii (H.H. Rusby) H.A.T. Harms (CÐE, A. inermis (O.P. Swartz) K.S. Kunth ex A.-P. de testa, embryos) Candole subsp. grandiflora (J.B.A. Guillemin & G.S. Perrottet) R.M. Polhill (not illustrated) Arachis C. Linnaeus (14.26) ...... 808 A. inermis (O.P. Swartz) K.S. Kunth ex A.-P. A. hypogaea C. Linnaeus (CÐE, testa, embryos) de Candole subsp. inermis (not illustrated) A. inermis (O.P. Swartz) K.S. Kunth ex A.-P. Argyrocytisus (R.C.J.E. Maire) D.G. Frodin & V.H. de Candole subsp. rooseveltii (E.A.J. Heywood ex C. Raymaud (30.14) ...... 1102 Wildeman) R.M. Polhill (not illustrated) A. battandieri (R.C.J.E. Maire) C. Raynaud (AÐE, valves, seeds, testa, embryos)

1186 Argyrolobium C.F. Ecklon & J.M. Zeyher (30.03) ...... 1078 A. sect. Phyllolobium A.A. von Bunge A. ser. Racemosae sensu W.H. Harvey (not (not illustrated) illustrated) A. sect. Platyglottis A.A. von Bunge A. biebersteinii P.W. Ball (CÐE, testa, embryos) (not illustrated) A. sect. Sesamei A.-P. de Candolle Arthrocarpum I.B. Balfour (14.22) ...... 2, 798 (not illustrated) A. gracile I.M. Balfour (AÐB, fruits) A. sect. Skythropos N.D. Simpson A. somalense J.O.D. Hillcoat & J.B. Gillett (not illustrated) (not illustrated) A. sect. Theiochrus A.A. von Bunge (not illustrated) Arthroclianthus H.E. Baillon (11.03) ...... 640 A. crassicarpus T. Nuttall (DÐF, testa, A. pseudo-malaccensis E. Vieillard ex A.T. embryos) Brongnairt & J.A.A. Gris (CÐE, testa, A. shelkovnikovii A.A. Grossheim embryos) (not illustrated)

Aspalathus C. Linnaeus (27.06) ...... 1040 Ateleia (A.-P. de Candolle) G. Bentham (1.13) ...... 112 A. argyrella P. MacOwan (not illustrated) A. herbert-smithii H.F. Pittier (CÐE, testa, A. bodkinii H. Bolus (not illustrated) embryos) A. comptonii R.M.T. Dahlgren (not illustrated) A. linearis (N.L. Burman) R.M.T. Dahlgren (CÐE, Austrodolichos B. Verdcourt (10.59) ...... 572 testa, embryos) A. errabundus (M. Schott) B. Verdcourt (AÐF, A. villosa C.P. Thunberg (not illustrated) fruits, seeds, testa, embryos)

Astracantha D. Podlech (16.16) ...... 850 Austrosteenisia R. Geesink (Millettieae) ...... 302 A. echinus (A.-P. de Candolle) D. Podlech (A, fruit) A. blackii (F. von Mueller) R. Geesink (AÐD, A. gossypina (F.E.L. von Fischer) D. Podlech (CÐE, fruits, seed, testa) testa, embryos) Baphia A. Afzelius ex C. Loddiges (2.18) ...... 160 Astragalus C. Linnaeus (16.15) ...... 844 B. capparidifolia J.G. Baker (CÐE, testa, embryos) A. sect. Acanthophace A.A. von Bunge (not illustrated) Baphiastrum H.A.T. Harms (2.22) ...... 170 A. sect. Aegacantha A.A. von Bunge B. boonei (E.A.J. Wildeman) P. Vermeulen ex (not illustrated) E.A.J. Wildeman (CÐE, testa, embryos) A. sect. Alopecuroidei A.-P. de Candolle B. brachycarpum H.A.T. Harms (A, fruits) (not illustrated) A. sect. Caprini A.-P. de Candolle (not illustrated) Baphiopsis G. Bentham ex J.G. Baker (1.05)...... 94 A. sect. Chlorostachys A.A. von Bunge B. parviflora G. Bentham ex J.G. Baker (AÐE, (not illustrated) fruit, seeds, testa, embryos) A. sect. Chronopus A.A. von Bunge (not illustrated) A. sect. Dasyphyllium A.A. von Bunge Baptisia É.P. Ventenat (29.05) ...... 1068 (not illustrated) B. alba (C. Linnaeus) É.P. Ventenat (CÐE, testa, A. sect. Eremophysa A.A. von Bunge embryos) (not illustrated) B. megacarpa A.W. Chapman ex J. Torrey & A. sect. Eremophysopsis N.F. Gontscharov A. Gray (not illustrated) (not illustrated) A. sect. Herpocaulos A.A. von Bunge Barbieria A.P. de Candolle (10.16A) ...... 476 (not illustrated) B. pinnata (C.H. Persoon) H.E. Baillon (AÐE, A. sect. Laxiflori C. Agerer-Kirchhoff fruits, seeds, testa, embryos) (not illustrated) A. sect. Megalocystis A.A. von Bunge Baukea G.C.W. Vatke (10.76), see Rhynchosia, p. 626 (not illustrated)

1187 Behaimia A.H.R Grisebach (Millettieae) ...... 304 Bossiaea É.P. Ventenat (23.06) ...... 950 B. cubensis A.H.R. Grisebach (AÐE, fruits, seeds, B. cinerea R. Brown (CÐE, testa, embryos) testa, embryos) Bowdichia K.S. Kunth (2.28) ...... 180 Belairia A. Richard (14.04) ...... 762 B. virgilioides K.S. Kunth (BÐE, seeds, testa, B. mucronata A.H.R. Grisebach (AÐE, fruits, seed, embryos) testa, embryos) Bowringia J.G. Champion ex G. Bentham (2.21) ...... 168 Beliceodendron C.L. Lundell, see Lecointea, p. 102 B. callicarpa J.G. Champion (A, CÐE, fruits, testa, B. tango (P.C. Standley) C.L. Lundell (not embryos) illustrated), see Lecointea, p. 102 Brachypterum R. Wight & G.A.W. Arnott (not illustrated), Bergeronia M. Micheli (Millettieae) ...... 306 see Derris, p. 324 B. sericea M. Micheli (AÐE, fruits, seed, testa, embryos) Brachysema R. Brown (24.16) ...... 988 B. bracteolosus F.H.J. von Mueller (not illustrated) Biserrula C. Linnaeus (16.15A) ...... 848 B. lanceolata C.D.F. Meissner (CÐE, testa, B. pelecinus C. Linnaeus (AÐE, fruits, seeds, testa, embryos) embryos) B. pelecinus subsp. leiocarpa (A. Richard) J.B. Brongniartia K.S. Kunth (22.01) ...... 934 Gillett (not illustrated) B. intermedia S. Moricand (CÐE, testa, embryos)

Bituminaria L. Heister ex P.C. Fabricius (12.02)...... 700 Brya P. Browne (11.01) ...... 636 B. bituminosa (C. Linnaeus) C.H. Stirton (AÐE, B. ebenus (C. Linnaeus) A.-P. de Candolle (AÐE, fruits, embryo and two fruits functioning as fruits, seeds, testa, embryos) seeds, exocarp, embryos) Bryaspis P. Duvigneaud (14.14) ...... 784 Bobgunnia J.H. Kirkbride & J.H. Wiersema (1.01A) ... 84 B. lupulina (L.D. Planchon ex G. Bentham) B. fistuloides (H.A.T. Harms) J.H. Kirkbride & J.H. P. Duvigneaud (AÐE, articles and fruit within Wiersema (not illustrated) bracts, seeds, testa, embryos) B. madagascariensis (A.N. Desvaux) J.H. Kirkbride & J.H. Wiersema (AÐE, partial Burkilliodendron A.R.K.R. Sastry (Millettieae) ...... 2 fruits, fruit in transection, and fruit in parti longitudinal section; seeds, testa, embryos) Burtonia R. Brown, see Gompholobium, p. 960 B. simplicifolia F. von Mueller & R. Tate Bocoa J.B.C.F. Aublet (1.02) ...... 86 (not illustrated), see Gompholobium, p. 960 B. mollis (G. Bentham) R.S. Cowan (B, DÐE, fruits and seeds, testa) Butea W. Roxburgh ex C.L. von Willdenow (10.04) .... 444 B. prouacensis J.B.C.F. Aublet (not illustrated) B. monosperma (J.B.A.P.M. de Lamarck) P.H.W B. viridiflora (A. Ducke) R.S. Cowan (F, embryos) Taubert (CÐE, testa, embryos)

Bolusafra C.E.O. Kuntze (10.75) ...... 618 Cadia P. Forsskål (2.36) ...... 196 B. bituminosa (C. Linnaeus) C.D.F. Meisner (AÐF, C. purpurea (G. Piccioli) W. Aiton (AÐE, fruits, fruits, seeds, testa, embryos) seeds, testa, embryos)

Bolusanthus H.A.T. Harms (2.37) ...... 198 Cajanus A.-P. de Candolle (10.73) ...... 612 B. speciosus (H. Bolus) H.A.T. Harms (AÐE, fruits, C. cajan (C. Linnaeus) C.F. Millspaugh (CÐE, seeds, testa, embryos) testa, embryos) C. goensis J.M. Dalziel (not illustrated) Bolusia G. Bentham (27.08) ...... 1048 B. rhodesiana A.G. Corbishley (CÐE, testa, embryos)

1188 Calia J.L. Berlandier (2.43), see Sophora, p. 214 Carmichaelia R. Brown (17.05) ...... 878 C. exsul H.F. Mueller (not illustrated) Calicotome J.H.F. Link (30.17) ...... 1110 C. uniflora T. Kirk (CÐE, testa, embryos) C. spinosa (C. Linnaeus) J.H.F. Link (CÐE, testa, embryos) Carrissoa E.G. Baker (10.79)...... 2

Callerya Endlicher (Millettieae)...... 308 Cascaronia A.H.R Grisebach (4.16) ...... 264 C. cinera (G. Bentham) A.M. Schot (CÐD, testa) C. astragalina A.H.R. Grisebach (AÐD, fruits, C. speciosa (J.G. Champion) A.M. Schot (E, seed, testa) embryos) Castanospermum A. Cunningham ex W.J. Hooker Callistachys É.P. Ventenat (24.12)...... 982 (2.12) ...... 142 C. lanceolata É.P. Ventenat (AÐE, fruits with C. australe A. Cunningham & C. Fraser ex calyx, seeds, testa, embryos) W.J. Hooker (AÐE, fruits, seed, testa, embryos) Calophaca F.E.L. von Fischer ex A.-P. de Candolle (16.12) ...... 840 Caulocarpus E.G. Baker, see Tephrosia, p. 380 C. wolgarica (C. Linnaeus f.) F.E.L. von Fischer ex A.-P. de Candolle (CÐE, testa, embryos) Centrolobium C.F.P. von Martius ex G. Bentham (4.12) ...... 254 Calopogonium A.N. Desvaux (10.45) ...... 540 C. paraense E.L.R. Tulasne (BÐF, seed in situ, C. galactoides (K.S. Kunth) W.B. Hemsley (CÐE, seeds, testa, embryos) testa, embryos) C. parceanum J. Hill (A, fruits) C. mucunoides A.N. Desvaux (not illustrated) Centrosema (A.-P. de Candolle) G. Bentham (10.14) .. 466 Calpurnia E.H.F. Meyer (25.09) ...... 1026 C. molle Martius ex G. Bentham (not illustrated) C. aurea (W. Aiton) G. Bentham (CÐE, testa, C. plumieri (J.P.F. Turpin ex C.H. Persoon) embryos) G. Bentham (B, valves) C. pubescens G. Bentham (not illlustrated) Camoensia F.M.J. Welwitsch ex G. Bentham (2.39) .... 202 C. virginianum (C. Linnaeus) G. Bentham (DÐF, C. maxima G. Bentham (AÐB, DÐF, a piece of a testa, embryos) fruit, external and internal views, testa, embryos) Chadsia W. Bojer (Millettieae) ...... 312 Camptosema W.J. Hooker & G.A.W. Arnott (10.24) ... 492 C. andravinensis H.E. Baillon (AÐE, fruit, seeds, C. rubicundum W.J. Hooker & G.A.W. Arnott testa, embryos) (CÐE, testa, embryos) Chaetocalyx A.-P. de Candolle (14.07) ...... 768 Campylotropis A.A. von Bunge (11.23) ...... 688 C. brasiliensis (J.R.T. Vogel) G. Bentham (E, C. macrocarpa (A.A. von Bunge) A. Rehder embryos) (CÐE, testa, embryos) C. latisiliqua (J.L.M. Poiret) G. Bentham ex W.B. Hemsley (CÐD, testa) Canavalia A.-P. de Candolle (10.21) ...... 486 C. septentrionalis J. Sauer (CÐE, testa, embryos) Chamaecytisus J.H.F. Link (30.15A) ...... 1108 C. austriacus (C. Linnaeus) J.H.F. Link (CÐE, Candolleodendron R.S. Cowan (1.03) ...... 90 testa, embryos) C. brachystachyum (A.-P. de Candolle) R.S. Cowan (AÐF, fruits, seeds, testa, embryos) Chapmannia J. Torrey & A. Gray (14.24) ...... 802 C. floridana J. Torrey & A. Gray (AÐE, articles Caragana P.C. Fabricius (16.11) ...... 838 and incomplete fruits, seeds, testa, embryos) C. densa V.L. Komarov (CÐE, testa, embryos)

1189 Chesneya J. Lindley ex S.F.L. Endlicher (16.14) ...... 842 C. fairchildiana R.A. Howard (CÐE, testa, C. rytidosperma H.F. Jaubert & É. Spach (CÐE, embryos) testa, embryos) C. polystachya G. Bentham (not illustrated), and see Barbieria, p. 476 Chordospartium T.F. Cheeseman (17.03) ...... 874 C. stevensonii T.F. Cheeseman (AÐE, fruits, seeds, Clitoriopsis R. Wilczek (10.17)...... 2 testa, embryos) Cochlianthus G. Bentham (10.08) ...... 454 Chorizema J.J.H. de Labillardière (24.10) ...... 976 C. gracilis G. Bentham (AÐD, fruit, seed, testa) C. ilicifolium J.J.H. de Labillardière (CÐE, testa embryos) Codariocalyx J.C. Hasskarl (11.10) ...... 656 C. retrorsum J.M. Taylor & M.D. Crisp C. gyroides (W. Roxburgh ex J.H.F. Link) (not illustrated) J.K. Hasskarl (CÐE, testa, embryos)

Christia C. Moench (11.17)...... 674 Coelidium (B.C. Vogel) G.W. Walpers (25.03) ...... 1014 C. obcordata (J.L.M. Poiret) R.C. Bakhhuizen C. muraltioides G. Bentham (C, seed) van den Brink (CÐE, testa, embryos) C. pageae H.M.L. Bolus (B, DÐF, seed, testa, embryos) Chrysoscias E.H.F. Meyer (10.78) ...... 624 C. argentea (C.P. Thunberg) C.A. Smith (A, CÐE, Collaea A.-P. de Candolle (10.26) ...... 496 fruits, testa, embryos) C. speciosa (J.L.A. Loiseleur-Deslongchamps) C. parviflora E.H.F. Meyer (B, seed) A.-P. de Candolle (CÐE, testa, embryos)

Cicer C. Linnaeus (20.01) ...... 910 Cologania K.S. Kunth (10.43) ...... 534 C. arietinum C. Linnaeus (CÐE, testa, embryos) C. ovalifolia K.S. Kunth (CÐE, testa, embryos)

Cladrastis C.S. Rafinesque-Schmaltz (2.41)...... 208 Colutea C. Linnaeus (16.05) ...... 826 C. lutea (F.A. Michaux) K.H.E.L. Koch (CÐE, C. orientalis P. Miller (CÐE, testa, embryos) testa, embryos) Corallospartium J.B. Armstrong (17.04) ...... 876 Clathrotropis (G. Bentham) H.A.T. Harms (2.26) ...... 176 C. crassicaule (J.D. Hooker) J.B. Armstrong C. brachypetala (E.L.R. Tulasne) A. Kleinhoonte (AÐE, fruits, seeds, testa, embryos) (CÐE, testa, embryos) C. nitida (G. Bentham) H.A.T. Harms (A, fruits) Cordyla J. de Loureiro (1.06) ...... 96 C. africana J. de Loureiro (BÐH, seed with Cleobulia C.F.P. von Martius ex G. Bentham (10.20) .. 484 endocarp functioning as testa, seed without C. leiantha G. Bentham (E, embryos) endocarp, endocarp, embryos, embryo C. multiflora C.F.P. von Martius ex G. Bentham magnified) (BÐD, seeds, testa) C. haraka R. Capuron (not illustrated)

Clianthus D.C. Solander ex J. Lindley (16.01) ...... 814 Coronilla C. Linnaeus (13.11)...... 738 C. formsus (G. Don) N.C. Ford & J.W. Vickery C. emerus C. Linnaeus (not illustrated), (not illustrated), and see Swainsona, p. 818 see Hippocrepis, p. 742 C. puniceus (G. Don) D.C. Solander ex J. Lindley C. scorpioides (C. Linnaeus) W.D.J. Koch (AÐE, broken fruit, seeds, testa, embryos) (CÐE, testa, embryos)

Clitoria C. Linnaeus (10.16) ...... 472 Coursetia A.-P. de Candolle (8.08) ...... 412 C. subgen. Clitoria (not illustrated) C. caribaea (N. von Jacquin) M. Lavin (CÐE, C. sect. Neurocarpum (A.N. Desvaux) J.G. Baker testa, embryos) (not illustrated) Craibia H.A.T. Harms & S.T. Dunn (Millettieae) ...... 314 C. brownii S.T. Dunn (CÐE, testa, embryos)

1190 Cranocarpus G. Bentham (11.02) ...... 638 Cyclocarpa A. Afzelius ex J.G. Baker (14.16)...... 788 C. martii G. Bentham (AÐD, fruit, seeds, testa) C. stellaris A. Afzelius ex J.G. Baker (AÐG, C. mezii P.H.W. Taubert (not illustrated) articles and fruit, article with marginal dehiscence, segment with seed dispersing, seeds, Craspedolobium H.A.T. Harms (Millettieae)...... 316 testa, embryos) C. schochii H.A.T. Harms (AÐE, fruits, seeds, testa, embryos) Cyclolobium G. Bentham (Millettieae) ...... 318 C. blanchetianum E.L.R. Tulasne (BÐE, seed, Cratylia C.F.P. von Martius ex G. Bentham (10.25) .... 494 testa, embryos) C. mollis C.F.P. von Martius ex G. Bentham (CÐE, testa, embryos) Cyclopia É.P. Ventenat (25.05) ...... 1020 C. galioides (B. Bergius) A.-P. de Candolle (A, Crotalaria C. Linnaeus (27.07) ...... 1044 CÐE, valves, testa, embryos) C. sect. Chrysocalycinae (G. Bentham) E.G. Baker subsection Stipulosa (E.G. Baker) F.A. Cymbosema G. Bentham (10.19)...... 482 Bisby & R.M. Polhill (not illustrated) C. roseum G. Bentham (AÐE, fruits, seeds, testa C. subgen. Priotropis (R. Wight & G.A.W. embryos) Arnott) C.D.F. Meisner (not illustrated) C. juncea C. Linnaeus (CÐE, testa, embryos) Cytisophyllum O.F. Lang (30.12) ...... 1098 C. spectabilis A.W. Roth (not illustrated) C. sessilifolium (C. Linnaeus) O.F. Lang (AÐD, C. stolzii (E.G. Baker) R.M. Polhill (not illustrated) seeds, testa, embryos)

Cruddasia D. Prain (10.13) ...... 464 Cytisopsis H.F. Jaubert & É. Spach (13.01) ...... 716 C. craibii C. Niyomdham (not illustrated) C. pseudocytisus (P.E. Boissier) Fertig (AÐE, C. insignis D. Prain (AÐE, fruits, seeds, testa, Fruit and valve, seeds, testa, embryos) embryos) C. laotica (F. Gagnepain) C. Niyomdham Cytisus R.L. Desfontaines (30.15) ...... 1104 (not illustrated) C. commutatus (H.M. Willkomm) J.I. Briquet C. pinnata (E.D. Merrill) C. Niyomdham (CÐE, testa, embryos) (not illustrated) Dahlstedtia G.O.A. Malme (Millettieae) ...... 2, 320 Cullen F.C. Medikus (12.01) ...... 696 D. pinnata (G. Bentham) G.O.A. Malme (AÐC, C. americanum (C. Linnaeus) P.A. Rydberg fruit, fruits, seed) (CÐE, exocarp, embryos) C. glandulosa (C. Linnaeus) J.W. Grimes Dalbergia C. Linnaeus f. (4.05) ...... 236 (not illustrated) D. arbutifolia A. Ducke (CÐE, testa, embryos)

Cupulanthus J. Hutchinson, see Brachysema, p. 988, and Dalbergiella E.G. Baker (Millettieae) ...... 322 Gompholobium, p. 960 D. nyasae E.G. Baker (CÐE, testa, embryos) C. bracteolosus (F.H.J. von Mueller) D. welwitschii (E.G. Baker) E.G. Baker J. Hutchinson (not illustrated), see Brachysema, (not illustrated) p. 988 Dalea C. Linnaeus (6.08) ...... 288 Cyamopsis A.-P. de Candolle (9.06) ...... 428 D. leporina (W. Aiton) A.A. Bullock C. seneganlensis J.B.A. Guillemin & G.S. Perrottet (not illustrated) (not illustrated) D. purpurea É.P. Ventenat (CÐE, testa, C. tetragonoloba (C. Linnaeus) P.H.W. Taubert embryos) (CÐE, testa, embryos) D. urceolota E.L. Greene (not illustrated)

Cyathostegia (G. Bentham) R.W. Schery (1.14) ...... 114 Dalhousiea N. Wallich ex G. Bentham (2.20) ...... 166 C. matthewsii (G. Bentham) R.W. Schery (AÐE, D. bracteata (W. Roxburgh) R.C. Graham ex fruits, seeds, testa, embryos) G. Bentham (CÐE, testa, embryos)

1191 Daviesia J.E. Smith (24.04) ...... 968 Dicerma A.-P. de Candolle, see Aphyllodium, p. 650 D. acicularis J.E. Smith (CÐE, testa, embryos) Dichilus A.-P. de Candolle (30.02) ...... 1076 Decorsea R. Viguier (10.54) ...... 560 D. lebeckioides A.-P. de Candole (CÐE, testa, D. schlechteri (H.A.T. Harms) B. Verdcourt embryos) (AÐE, fruits, seeds, testa, embryos) Dicraeopetalum H.A.T. Harms (2.33)...... 190 Dendrolobium (R. Wight & G.A.W. Arnott) D. capuronianum (M. Peltier) G.P. Yakovlev G. Bentham (11.06) ...... 646 (CÐE, testa, embryos) D. cumingianum G. Bentham (not illustrated) D. dispermum (B. Hayata) A.K. Schindler Dillwynia J.E. Smith (24.25) ...... 1004 (not illustrated) D. floribunda J. Smith (CÐE, testa, embryos) D. lanceolatum (S.T. Dunn) A.K. Schindler (CÐE, testa, embryos) Dioclea K.S. Kunth (10.18) ...... 478 D. rostratum (A.K. Schindler) A.K. Schindler D. multiflora (J. Torrey & A. Gray) C.T. Mohr (not illustrated) (not illustrated) D. reflexa J.D. Hooker (CÐE, testa, embryos) Derris J. de Loureiro (Millettieae) ...... 324 D. sect. Brachypterum (R. Wight & G.A.W. Diphyllarium F. Gagnepain (10.37) ...... 522 Arnott) G. Bentham (not illustrated) D. mekongense F. Gagnepain (AÐE, fruits, seeds, D. sect. Derris (not illustrated) testa, embryos) D. sect. Dipteroderris G. Bentham (not illustrated) D. sect. Paraderris F.A.W. Miquel (not illustrated) Diphysa N. von Jacquin (14.03) ...... 760 D. robusta (W. Roxburgh ex A.-P. de Candolle) D. minutifolia J.N. Rose (E, embryos) G. Bentham (CÐE, testa, embryos) D. robinioides G. Bentham (CÐD, testa) D. trifoliata J. de Loureiro (not illustrated) Diplotropis G. Bentham (2.27) ...... 178 Desmodiastrum (D. Prain) A. Pramanik & D. peruviana J.F. Macbride (BÐF, seeds, testa, K. Thothathri (11.18A) ...... 678 embryos, embryo magnified) D. belgaumense (W. Wight) A. Pramanik & D. purpurea (L.C.M. Richard) G.J.H. Amshoff K. Thothathri (AÐB, fruit, seeds) (GÐH, embryos, embryo magnified)

Desmodium A.N. Desvaux (11.09)...... 652 Dipogon F.M. Liebmann (10.63) ...... 580 D. subgen. Sagatia (O.P. Swartz) A.-P. D. lignosus (C. Linnaeus) B. Verdcourt (AÐE, de Candolle (not illustrated) fruit, seeds, testa, embryos) D. auricomum R. Graham (not illustrated) D. heterophyllum (C.L. von Willdenow) A.-P. Dipterocarpus C. Gaertner (Dipterocarpaceae), see de Candolle (not illustrated) Apoplanesia, p. 274 D. microphyllum (C.P. Thunberg) A.-P. de Candolle (not illustrated) Dipteryx J.C.D. von Schreber (3.01) ...... 220 D. oojeinensis (W. Roxburgh) H. Ohasi D. alata (J.R.T. Vogel) P.H.W. Taubert (not illustrated) (not illustrated) D. styracifolium (P. Osbeck) E.D. Merrill D. punctata (J. Blake) G.J.H. Amshoff (CÐE, (not illustrated) testa, embryos) D. uncinatum (N. von Jacquin) A.-P. D. trifoliolata A. Ducke (not illustrated) de Candolle (CÐE, testa, embryos) Discolobium G. Bentham (14.17) ...... 790 Dewevrea M. Micheli (Millettieae) ...... 328 D. psoraleifolium G. Bentham (BÐF, fruit diagram D. bilabata M. Micheli (AÐE, fruits, seeds, showing coiling, seed, testa, embryos) testa, embryos) D. pulchellum G. Bentham (not illustrated)

1192 Disynstemon R. Viguier (Millettieae)...... 330 Ebenus C. Linnaeus (18.07) ...... 892 D. paullinioides (J.C. Baker) M.A.G. Peltier (AÐE, E. cretica C. Linnaeus (DÐF, testa, embryos) fruit, seed, testa, embryos) Echinospartum (É. Spach) J.P. Fourreau (30.23) ...... 1124 Dolichopsis E. Hassler (10.69) ...... 598 E. horridum (J.L.M. Vahl) W.H.P. Rothmaler D. paraguariensis E. Hassler (AÐE, fruits, seeds, (not illustrated) testa, embryos) E. lusitanicum (C. Linnaeus) W.H.P. Rothmaler (CÐE, testa, embryos) Dolichos C. Linnaeus (10.64) ...... 582 D. gululu E.A.J. De Wildeman (CÐE, testa, Eleiotis A.-P. de Candolle (11.21)...... 684 embryos) E. monophylla (N.L. Burman) A.-P. de Candolle. D. monticola C.F.P. von Martius ex G. Bentham (AÐE, fruits, seeds, testa, embryos) (not illustrated), see Dolichopsis, p. 598 Eligmocarpus R. Capuron (Caesalpinioideae: Cassieae) Dolichovigna B. Hayata, see Vigna, p. 588 E. cynometroides R. Capuron, see Discolobium, p. 790 Dorycnium P. Miller (13.06) ...... 728 D. sect. Bonjeania (H.G.L.R. Reichenbach) Eminia P.H.W. Taubert (10.29)...... 502 P.E. Boissier (not illustrated) E. antennulifera (J.G. Baker) P.H.W. Taubert D. sect. Dorycnium (not illustrated) (CÐE, testa, embryos) D. hirsutum (C. Linnaeus) N.C. Seringe (CÐE, testa, embryos) Endomallus pellitus F. Gagnepain (not illustrated), see Cajanus, p. 612 Dorycnopsis P.E. Boissier (13.05) ...... 726 E. spirei F. Gagnepain (not illustrated), D. abyssinica (A. Richard) V.N. Tikhomirov & see Cajanus, p. 612 D.D. Sokoloff (not illustrated), and see Vermifrux, p. 736 Endosamara R. Geesink (Millettieae) ...... 332 D. gerardii (C. Linnaeus) P.E. Boissier (AÐE, E. racemosa (W. Roxburgh) R. Geesink (AÐB, fruits, seeds, testa, embryos) fruits, seeds)

Droogmansia E.A.J. De Wildeman (11.14) ...... 666 Eremosparton F.E.L. von Fischer & B. Meyer D. pteropus (J.G. Baker) E.A.J. De Wildemann (16.09) ...... 834 var. whytei (A.K. Schindler) B. Verdcourt E. aphyllum F.E.L. von Fischer & B. Meyer (CÐE, testa, embryos) (CÐE, testa, embryos)

Dumasia A.-P. de Candolle (10.42) ...... 532 Erichsenia W.B. Hemsley (24.05) ...... 2 D. truncata P.F.B. von Siebold & J.G. Zuccarini (CÐE, testa, embryos) Erinacea M. Adanson (30.18) ...... 1112 D. villosa A.-P. de Candolle (A, fruits) E. anthyllis J.H.F. Link (AÐE, fruit within calyx and corolla and valve, seeds, testa, embryos) Dunbaria R. Wight & G.A.W. Arnott (10.74) ...... 616 D. punctata (R. Wight & G.A.W. Arnott) G. Eriosema (A.-P. de Candolle) H.G.L.R. Reichenbach .. Bentham (CÐE, testa, embryos) (10.81) ...... 630 E. floribundum G. Bentham (CÐE, testa, embryos) Dussia C.W.L. Krug & I. Urban ex P.H.W. Taubert E. glaziovii H.A.T. Harms (not illustrated) (2.24) ...... 174 D. lehmannii H.A.T. Harms (CÐE, testa, embryos) Errazurizia R.A. Philippi (6.05) ...... 282 E. rotundata (E.O. Wooton) R.C. Barneby (CÐE, Dysolobium (G. Bentham) D. Prain (10.50) ...... 550 testa, embryos) D. grande N. Wallich ex (G. Bentham) D. Prain (CÐE, testa, embryos)

1193 Erythrina C. Linnaeus (10.01)...... 434 Fordia W.B. Hemsley (Millettieae) ...... 334 E. stricta W. Roxburgh (CÐE, testa, embryos) F. splendidissima (C.L. von Blume ex F.A.W. E. suberosa W. Roxburgh (not illustrated) Miquel) J.R.M. Buijsen (CÐE, testa, embryos)

Etaballia G. Bentham (2.09) ...... 136 Galactia P. Browne (10.27)...... 498 E. dubia (K.S. Kunth) V.E. Rudd (AÐE, fruits, G. jussiaeana K.S. Kunth (CÐE, testa, embryos) seed, testa, embryos) Galega C. Linnaeus (16.20) ...... 860 Euchilopsis F.H.J. von Mueller (24.20) ...... 994 G. officinalis C. Linnaeus (CÐE, testa, embryos) E. linearis (G. Bentham) F.H.J. von Mueller (AÐE, fruits and valve, seed, testa) Gastrolobium R. Brown (24.13) ...... 984 G. bilobum R. Brown (CÐE, testa, embryos) Euchresta J.J. Bennett (28.01) ...... 1056 E. japonica (J.D. Hooker) E.A. von Regal Geissaspis R. Wight & G.A.W. Arnott (14.13)...... 782 (not illustrated) G. cristata R. Wight & G.A.W. Arnott (CÐE, E. horsfieldii (J.B.L.T. Lescherault de la Tour) testa, embryos) J.J. Bennett (DÐF, testa, embryos) E. trifoliolata E.D. Merrill (not illustrated) Genista C. Linnaeus (30.22) ...... 1120 G. cephalantha É. Spach (not illustrated) Eutaxia R. Brown (24.26) ...... 1006 G. germanica C. Linnaeus (CÐE, testa, embryos) E. microphylla (R. Brown) J.M. Black (CÐE, testa, embryos) Genistidium I.M. Johnston (8.11) ...... 420 G. dumosum I.M. Johnson (AÐE, fruit and valve, Eversmannia A.A. von Bunge (18.01) ...... 882 seed, testa, embryos) E. hedysaroides A.A. von Bunge (not illustrated) E. subspinosa (F.E.L. von Fischer ex A.-P. Geoffroea N. von Jacquin (4.15) ...... 262 de Candolle) B.A. Fedtschenko (AÐE, G. decorticans (J. Gillies ex W.J. Hooker & fruits, seeds, testa, embryos) G.A.W. Arnott) A.E. Burkart (DÐE, testa) G. spinosa N. von Jacquin (F, embryos) Exostyles H.W. Schott (1.10) ...... 2 Gliricidia K.S. Kunth (8.06)...... 406 Eysenhardtia K.S. Kunth (6.02) ...... 276 G. brenningii (H.A.T. Harms) M. Lavin E. orthocarpa (A. Gray) S. Watson (not illustrated) (not illustrated) E. parvifolia T.S. Brandegee (not illustrated) G. maculata (K.S. Kunth) K.S. Kunth ex G.W. E. spinosa G. Engelmann (CÐE, testa, embryos) Walpers (not illustrated) G. sepium (N. von Jacquin) K.S. Kunth ex G.W. Faba P. Miller, see Vicia, p. 894 Walpers (A, CÐE, dehisced fruit and valves, testa, embryos) Fiebrigiella H.A.T. Harms (14.06) ...... 766 F. gracilis H.A.T. Harms (AÐF, fruit, fruit and Glottidium A.N. Desvaux (8.01A) ...... 392 article, seed, testa, embryos) G. vescarium A.N. Desvaux (AÐE, fruits (dehisced fruit and nondehisced endocarp), Fissicalyx G. Bentham (4.07) ...... 244 seeds, testa, embryos) F. fendleri G. Bentham (AÐE, fruits, seed, testa, embryos) Glycine C.L. von Willdenow (10.35) ...... 516 G. max (C. Linnaeus) E.D. Merrill (B, DÐF, seeds, Flemingia W. Roxburgh ex W.T. Aiton (10.77) ...... 620 testa, embryos) F. macrophylla (C.L. von Willdenow) E.D. Merrill (CÐE, testa, embryos) Glycyrrhiza C. Linnaeus (16.22) ...... 866 G. lepidota F.T. Pursh (CÐE, testa, embryos)

1194 Gompholobium J.E. Smith (24.01) ...... 960 H. mexicana J.N. Rose (not illustrated) G. latifolium J.E. Smith (E, embryos) H. parvifolia H.S. Irwin & M.T.K. Arroyo G. papuanum E.D. Merrill & L.M. Perry (not illustrated) (not illustrated) H. robusta H.S. Irwin & M.T.K. Arroyo G. scabra R. Brown (CÐD, testa) (not illustrated)

Gonocytisus É. Spach (30.20) ...... 1116 Hebestigma I. Urban (8.02) ...... 396 G. angulatus (C. Linnaeus) É. Spach (CÐE, H. cubense (K.S. Kunth) I. Urban (AÐE, valves, testa, embryos) seeds, testa, embryos)

Goodia R.A. Salisbury (23.05) ...... 948 Hedysarum C. Linnaeus (18.02) ...... 884 G. lotifolia G. Salisbury (not illustrated) H. coronarium C. Linnaeus (CÐE, testa, embryos) G. medicagnea F.H.J. von Mueller (CÐE, testa, embryos) Herpyza C. Wright (10.28) ...... 2, 500 H. grandiflora C. Wright (AÐB, fruit, seed) Grazielodendron H.C. de Lima (4.09) ...... 248 G. rio-docensis H.C. de Lima (AÐE, fruits, seed, Hesperolaburnum R.C.J.E. Maire (30.10) ...... 1094 testa, embryos) H. platycarpum (R.C.J.E. Maire) R.C.J.E. Maire (AÐE, valves showing endocarp and epicarp, Gueldenstaedtia F.E.L. von Fischer (16.19) ...... 858 seeds, testa, embryos) G. himalaica J.G. Baker (CÐE, testa, embryos) Hesperothamnus T.S. Brandegee (Millettieae) ...... 338 Halimodendron F.E.L. von Fischer ex A.-P. de Candolle H. littoralis (T.S. Brandegee) T.S. Brandegee (16.10) ...... 836 (BÐE, seed, testa, embryos) H. halodendron (P.S. von Pallas) A. Voss (AÐE, fruits, seeds, testa, embryos) Hippocrepis C. Linnaeus (13.13) ...... 742 H. ciliata C.L. von Willdenow (not illustrated) Hammatolobium E. Fenzl (13.17) ...... 752 H. emerus (C. Linnaeus) P. Lassen (B, fruits and H. kremerianum (E.St.-C. Cosson) C.H. Muller fruit segments) (E, embryos) H. unisiliquosa C. Linnaeus (DÐF, testa, embryos) H. lotoides E. Fenzl (AÐD, fruits with calyx and 1 article, seeds, testa) Hoita P.A. Rydberg (12.06) ...... 708 H. macrostachys (A.-P. de Candolle) P.A. Rydberg Haplormosia H.A.T. Harms (2.16) ...... 154 (CÐE, testa, embryos) H. monophylla (H.A.T. Harms) H.A.T. Harms H. strobilina (J.W. Hooker & G.A. Arnott) G.A.W. (A, CÐE, fruits, testa, embryos) Rydberg (not illustrated)

Hardenbergia G. Bentham (10.48) ...... 546 Holocalyx M. Micheli (1.12) ...... 110 H. violacea (G.V. Schneevoogt) W.T. Stearn H. balansae M. Micheli (AÐF, fruits, seeds, testa, (A, CÐE, fruits, testa, embryos) embryos, embryo magnified)

Harleyodendron R.S. Cowan (1.09) ...... 106 Hovea R. Brown ex W.T. Aiton (23.04) ...... 946 H. unifoliolatum R.S. Cowan (AÐD, fruit, seeds, H. heterophylla A. Cunningham ex J.D. Hooker testa) (not illustrated) H. linearis (J. Smith) R. Brown (not illustrated) Harpalyce A.-P. de Candolle (22.02) ...... 936 H. elliptica A.-P. de Candolle (CÐE, testa, H. sect. Harpalyce (not illustrated) embryos) H. sect. Brazilianae M.T.K. Arroyo (not illustrated) H. sect. Cubenses M.T.K. Arroyo (not illustrated) H. brasiliana G. Bentham (CÐE, testa, embryos) H. macrocarpa N.L. Britton & P. Wilson (not illustrated)

1195 Humularia P. Duvigneaud (14.15) ...... 786 I. glandulosa J.C. Wendland (not illustrated) H. chevalieri (E.A.J. De Wildeman) P.A. I. kirilowii C.J.M. Maximowicz & I.V. Palibin Duvigneaud (DÐF, testa, embryos) (DÐF, testa, embryos) H. drepanocephalus (E.G. Baker) P.A. I. trifoliata C. Linnaeus (not illustrated) Duvigneaud (B, fruit) I. glandulosa J.C. Wendland (not illustrated) I. linifolia (C. Linnaeus) A.J. Retzius Hybosema H.A.T. Harms (8.04) ...... 400 (not illustrated) H. ehrenbergii (F.R.R. Schlechter) H.A.T. Harms I. luzonensis. I. de Kort & G. Thijsse (AÐE, valves, seeds, testa, embryos) (not illustrated) I. parvifolia B. Heyne ex R. Wright & G.A.W. Hymenocarpos C.G. Savi (13.04)...... 724 Arnott (not illustrated) H. circinnatus (C. Linnaeus) C.G. Savi (AÐE, I. zollingeriana F.A.W. Miquel (not illustrated) fruits, seeds, testa, embryos) Inocarpus J.R. Forster & J.G. Forster (2.10) ...... 138 Hymenolobium G. Bentham (4.03) ...... 232 I. fagifer (S. Parkinson) F.R. Fosberg (BÐE, seeds, H. heringerianum C.T. Rizzini (CÐE, seeds, testa, testa, embryos) embryos) H. modestum A. Ducke (not illustrated) Isotropis G. Bentham (24.06) ...... 970 I. wheeleri G. Bentham (CÐE, testa, embryos) Hypocalyptus C.P. Thunberg (26.01) ...... 1028 H. coluteoides (J.B.A.P. de M. de Lamarck) R.M.T. Jacksonia R. Brown ex J.E. Smith (24.08)...... 972 Dahlgren (not illustrated) J. ramosissima G. Bentham (CÐE, testa, embryos) H. sophoroides (P.J. Bergius) H.E. Baillon (AÐE, fruits and valve, seeds, testa, embryos) Jansonia R. Kippist ex J. Lindley (24.17) ...... 2

Imbralyx Geesink, see Fordia, p. 334 Kennedia É.P. Ventenat (10.47) ...... 544 K. macrophylla (C.F.W. Meissner) G. Bentham Indigastrum H.F. Jaubert & É. Spach, see Indigofera, p. 430 (CÐE, testa, embryos) I. argyraeum (C.F. Ecklon & C.L.P. Zeyher) B.D. Schrire (not illustrated), see Indigofera, Kotschya S.F.L. Endlicher (14.11) ...... 778 p. 430 K. africana S.F.L. Endlicher var. bequaertii I. fastigiatum (E.H.F. Meyer) B.D. Schrire (E.A.J. De Wildeman) B. Verdcourt (DÐF, (not illustrated), see Indigofera, p. 430 testa, embryos) I. parvifolium (B. Heyne ex R. Wright & G.A.W. Arnott) B.D. Schrire (not illustrated), see Kummerowia A.K. Schindler (11.25) ...... 694 Indigofera, p. 430 K. stipulacea (C.J. Maximowicz) T. Makino (CÐE, testa, embryos) Indigofera C. Linnaeus (9.07) ...... 430 I. subgen. Indigastrum (H.F. Jaubert & É. Spach) Kunstleria D. Prain (Millettieae) ...... 340 J.B. Gillett, (not illustrated) I. subgen. Microcharis (G. Bentham) J.B. Gillett Lablab M. Adanson (10.61) ...... 576 (not illustrated) L. purpureus (C. Linnaeus) R. Sweet (AÐE, fruits, I. argyraea C.F. Ecklon & C.L.P. Zeyher seeds, testa, embryos) (not illustrated) I. australis C.L. von Willdenow (not illustrated) Laburnum P.C. Fabricius (30.09) ...... 1092 I. colutea (N.L. Burman) E.D. Merrill L. anagyroides F.C. Medikus (CÐE, testa, (not illustrated) embryos) I. cryptantha G. Bentham ex W.H. Harvey (not illustrated) I. disjuncta J.B. Gillett (not illustrated) I. fastigiata E.H.F. Meyer (not illustrated)

1196 Lackeya R.H. Fortunato, L.P. de Queiroz, & G.P. Lewis, von Meyer) P.H. Davis & U. Plitmann see Dioclea, p. 478 (not illustrated) L. multiflora (J. Torrey & A. Gray) R.H. L. nigricans subsp. ervoides (Grande) G. Fortunato, L.P. de Queiroz, & G.P. Lewis Ladizinsky (not illustrated) (not illustrated), see Dioclea, p. 478 L. orientalis (P.E. Boissier) H. Handel-Mazzetti (not illustrated) Lamprolobium G. Bentham (23.02) ...... 924 L. tomentosus G. Ladizinsky (not illustrated) L. fruticosum G. Bentham (AÐE, valve, seeds, testa, embryos) Leptoderris S.T. Dunn (Millettieae) ...... 342 L. aurantiaca S.T. Dunn (E, testa, embryos) Lathyrus C. Linnaeus (19.02) ...... 898 L. nobilis S.T. Dunn (CÐD, testa) L. sect. Orobus (C. Linnaeus) J.G. Baker (not illustrated) Leptodesmia G. Bentham (11.20) ...... 682 L. amphicarpos C. Linnaeus (not illustrated) L. congesta G. Bentham ex J.G. Baker (CÐE, L. cassius P.E. Boissier (not illustrated) testa, embryos) L. ciliolatus K. H. Rechinger (not illustrated) L. hirsutus C. Linnaeus (not illustrated) Leptosema G. Bentham (24.07) ...... 2 L. lentiformis U. Plitman (not illustrated) L. sativus C. Linnaeus (CÐE, testa, embryos) Lespedeza A. Michaux (11.24) ...... 690 L. setifolius C. Linnaeus (not illustrated) L. formosa (J.R.T. Vogel) B.A.E. Koehne (not illustrated) Latrobea C.D.F. Meissner (24.18) ...... 990 L. stipulacea C.J. Maximowicz (not illustrated) L. tenella G. Bentham (CÐE, testa, embryos) L. striata C.P. Thunberg (not illustrated) L. virginica (C. Linnaeus) N.L. Britton (CÐE, Lebeckia C.P. Thunberg (27.10)...... 1052 testa, embryos) L. capensis (C. Linnaeus) G.C. Druce (CÐE, testa, embryos) Lessertia A.-P. de Candolle (16.04) ...... 824 L. benguellensis J.G. Baker (CÐE, testa, embryos) Lecointea A. Ducke (1.08) ...... 102 L. amazonica A. Ducke (AÐF, fruits, seeds in situ Leucomphalos G. Bentham ex L.D. Planchon (2.23) .... 172 testa, embryos, embryo magnified) L. capparideus G. Bentham ex L.D. Planchon L. tango (P.C. Standley) G.P. Yakovlev (GÐH, (AÐF, fruits, seed, testa, embryos, embryo embryos, embryo magnified) magnified)

Lembotropis A.H.R. Grisebach (30.16), see Cytisus, Liparia C. Linnaeus (25.04) ...... 1016 p. 1188 L. splendens (N.L. Burman) J.J. Bos & H.C.D. de Wit (AÐE, valves, seeds, testa, embryos) Lennea J.F. Klotzsch (8.03) ...... 398 L. modesta (P.C. Standley & J.A. Steyermark) Lonchocarpus K.S. Kunth (Millettieae) ...... 344 P.C. Standley & J.A. Steyermark (AÐE, (and see Muellera, p. 352) valves, seeds, testa, embryos) L. capassa R.A. Rolfe (CÐE, testa, embryos) L. fluvialis (C.A.M. Lindman) R.H. Fortunato & Lens P. Miller (19.03) ...... 902 R. Palese (not illustrated), see Muellera, L. culinaris F.C. Medikus (not illustrated) p. 352 L. culinaris subsp. orientalis (P.E. Boissier) L. sericeus (J.L.M. Poiret) K.S. Kunth ex A.-P. B.S. Williams (not illustrated) de Candolle (not illustrated) L. ervoides (G. de Brignoli di Brunnhoff) L. Grande (CÐE, testa, embryos) Lotononis (A.-P. de Candolle) C.F. Ecklon & J.M. L. nigricans (F.A. Marchall von Bieberstein) Zeyher (27.09) ...... 1050 D.A. Godman (not illustrated) L. bainesii J.G. Baker (CÐE, testa, embryos) L. montbretii (F.E.L. von Fischer & C.A.A.

1197 Lotus C. Linnaeus (13.07) ...... 730 Macrotyloma (R. Wight & G.A.W. Arnott) L. sect. Pseudosimpeteria T.E. Kramina B. Verdcourt (10.65) ...... 584 D.D. Sokoloff (not illustrated) M. axillare (E.H.F. Meyer) B. Verdcourt (CÐE, L. corniculatus C. Linnaeus (DÐF, testa, embryos) testa, embryos) L. maritimus C. Linnaeus (B, fruit cross-section with four wings) Margaritolobium H.A.T. Harms (Millettieae) ...... 2 L. roudairei E. Bonnet (not illustrated), and see Acmispon, p. 734 Marina F.M. Liebmann (6.07) ...... 286 M. neglecta (B.L. Robinson) R.C. Barneby (CÐE, Luetzelburgia H.A.T. Harms (2.03) ...... 124 testa, embryos) L. praecox (H.A.T. Harms) H.A.T. Harms (BÐE, seed, testa, embryos) Mastersia G. Bentham (10.38) ...... 524 M. bakeri (S.H. Koorders) C.A. Backer (CÐE, Lupinus C. Linnaeus (30.08) ...... 1088 testa, embryos) L. tauris G. Bentham (CÐE, testa, embryos) Mecopus J.J. Bennett (11.15) ...... 670 Luzonia A.D.E. Elmer (10.23)...... 2 M. nidulans J.J. Bennett (AÐF, “bird cage,” fruits, seeds, testa, embryos) Lygos M. Adanson, see Retama, p. 1118 Medicago C. Linnaeus (21.05) ...... 928 Maackia F.J. Ruprecht & C.J. Maximowicz (2.40)...... 206 M. ciliaris (C. Linnaeus) C. Allioni (CÐE, testa, M. amurensis F.J. Ruprecht (CÐE, testa, embryos) embryos) M. hypogaea E. Small (not illustrated) Machaerium C.H. Persoon (4.06)...... 240 M. plicata (P.E. Boissier) G.I. Sirjaev M. lunatum (C. Linnaeus filius) W.A. Ducke (not illustrated) (not illustrated) M. popovii (E.I. Korneva) G.I. Sirjaev M. opacum J.R.T. Vogel (CÐE, testa, embryos) (not illustrated) M. quinta (J.C.B.F. Aublet) N.Y. Sandwith M. scutellata (C. Linnaeus) P. Miller (not illustrated) (not illustrated) M. robiniifolium (A.-P. de Candolle) J.R.T. Vogel (not illustrated) Meizotropis J.O. Voigt (10.06) ...... 448 M. buteiformis J.O. Voigt (BÐE, seed, testa, Macropsychanthus H.A.T. Harms & C.A.G. embryos) Lauterbach (10.22) ...... 490 M. dolichobotrys L.B. Holthuis (CÐD, testa) Melilotus P. Miller (21.03) ...... 920 M. lauterbachii H.A.T. Harms (not illustrated) M. altissimus J.L. Thuillier (not illustrated) M. mindanaensis E.D. Merrill (A, fruit) M. bicolor P.E. Boissier & B. Balansa (not illustrated) Macroptilium (G. Bentham) I. Urban (10.71) ...... 602 M. indicus (C. Linnaeus) C. Allioni (CÐE, testa, M. atropurpureum (A. P. de Candolle) I. Urban embryos) (not illustrated) M. gracile (E.F. Poeppig ex G. Bentham) I. Urban Melliniella H.A.T. Harms (11.19) ...... 680 (not illustrated) M. micrantha H.A.T. Harms (AÐE, fruits with M. heterophyllum (C.L. von Willdenow) and without calyx, seeds, testa, embryos) R. Maréchal & J.C. Baudet (not illustrated) M. lathyroides (C. Linnaeus) I. Urban (CÐE, Melolobium C.F. Ecklon & J.M. Zeyher (30.01) ...... 1072 testa, embryos) M. decumbens (E.H.F. Meyer) J. Burtt Davy M. panduratum (C.F.P. von Martius ex (CÐE, testa, embryos) G. Bentham) R. Maréchal & J.C. Baudet (not illustrated) Microcharis G. Bentham, see Indigofera, p. 430 M. pedatum (J.N. Rose) R. Maréchal & M. disjunta (J.B. Gillett) B.D. Schrire J.C. Baudet (not illustrated) (not illustrated), see Indigofera, p. 430

1198 Mildbraediodendron H.A.T. Harms (1.07) ...... 2, 100 Myrocarpus F.F. Allemäo e Cysneiro (2.05) ...... 128 M. excelsum H.A.T. Harms (AÐF), fruit, seed M. frondosus F.F. Allemäo e Cysneiro (AÐE, with endocarp functioning as testa, endocarp, fruits, seeds, testa, embryos) embryos, embryo magnified Myrospermum N. von Jacquin (2.06) ...... 130 Millettia R. Wight & G.A.W. Arnott (Millettieae)...... 348 M. frutescens N. von Jacquin (AÐE, fruits, seed, M. cinerea G. Bentham (E, embryos) testa, embryos) M. grandis (E.H.F. Meyer) H.C. Skeels (CÐD, M. sousanum A. Delgado Salinas & M.C. testa) Johnston (not illustrated) M. racemosa (W. Roxburgh) G. Bentham, see Endosamara, p. 332 Myroxylon C. Linnaeus f. (2.07) ...... 132 M. balsamum (C. Linnaeus) H.A.T. Harms var. Mirbelia J.E. Smith (24.11) ...... 978 pereirae (J.F. Royle) H.A.T. Harms (CÐE, testa, M. grandiflora W. Aiton (CÐE, testa, embryos) embryos)

Monopteryx R. Spruce ex G. Bentham (2.31) ...... 186 Mysanthus G.P. Lewis & A. Delgado Salinas M. uaucu R. Spruce ex G. Bentham (AÐD, fruit, (10.72A) ...... 610 seed, embryos) M. uleanus (H.A.T. Harms) G.P. Lewis & A.O. Delgado Salinas var. uleanus (AÐE, fruits, Montigena P.B. Heenan, see Swainsona, p. 818 seeds, testa, embryos) M. novae-zelandiae (J.D. Hooker) P.B. Heenan (not illustrated), see Swainsona, p. 818 Nemcia K. Domin (24.15) ...... 986 N. dilatata (G. Bentham) M.D. Crisp (CÐE, Mucuna M. Adanson (10.03)...... 440 testa, embryos) M. subgen. Stizolobium (P. Browne) D. Prain (not illustrated) Neocollettia W.B. Hemsley (11.26) ...... 2 M. pruriens (C. Linnaeus) A.-P. de Candolle (CÐE, testa, embryos) Neodielsia H.A.T. Harms (16.18) ...... 845 M. sloanei W. Fawcett & A.B. Rendle (not illustrated) Neodunnia R. Viguier (Millettieae)...... 2

Muellera C. Linnaeus f. (Millettieae) ...... 352 Neoharmsia R. Viguier (2.34) ...... 192 M. fluvialis (C.A.M. Lindman) A. Burkart N. baroni (E. Drake del Castillo) R. Viguier ex (not illustrated) G. Bentham (CÐE, testa, embryos) M. moniliformis C. Linnaeus f. (BÐE, seeds, testa, embryos) Neonotonia J.A. Lackey (10.40)...... 528 N. verdcourtii D. Isely (CÐE, testa, embryos) Muelleranthus J. Hutchinson (23.08) ...... 954 N. wrightii (G.A.W. Arnott) J.A. Lackey (AÐB, M. crenulatus A.T. Lee (not illustrated) fruits, seeds) M. stipularis (J.M. Black) A.T. Lee (AÐE, fruits, seeds, testa, embryos) Neorautanenia H. Schinz (10.60) ...... 574 M. trifoliatus (F.H. Mueller) J. Hutchinson ex N. amboensis H. Schinz (A, CÐE, fruit, testa, A.T. Lee (not illustrated) embryos)

Mundulea (A.-P. de Candolle) G. Bentham Neorudolphia N.L. Britton (10.10)...... 458 (Millettieae) ...... 354 N. volubilis (C.L. von Willdenow) N.L. Britton M. sericea (C.L. von Willdenow) A.J.B. (AÐE, fruits, seeds, testa, embryos) Chevalier (CÐE, testa, embryos) Nephrodesmus A.K. Schindler (11.04)...... 2, 642 Murtonia W.G. Craib, see Desmodium, p. 652 N. albus A.K. Schindler (A, fruits) N. sericeus (B.P.G. Hochreutiner) A.K. Schindler (not illustrated)

1199 Nesphostylis B. Verdcourt (10.58) ...... 570 Ormocarpum A.M.F.J. Palisot de Beauvois (14.01) ..... 754 N. holosericea (J.G. Baker) B. Verdcourt (AÐE, O. kirkii S. le M. Moore (not illustrated) fruit, seeds, testa, embryos) O. orientale (C.P.J. Sprengel) E.D. Merrill (CÐE, testa, embryos) Nissolia N. von Jacquin (14.08) ...... 770 N. leiogyne N.Y. Sandwith (E, embryos) Ormosia G. Jackson (2.15) ...... 150 N. schottii (J. Torrey) A. Gray (CÐD, testa) O. panamensis G. Bentham (CÐE, testa, embryos)

Nogra E.D. Merrill (10.33)...... 512 Ornithopus C. Linnaeus (13.15) ...... 748 N. dalzellii (E.G. Baker) E.D. Merrill (AÐE, O. micranthus (G. Bentham) J. Arechaveleta & fruits, seeds, testa, embryos) Bolpardo (not illustrated) N. filicaulis (Kurz) E.D. Merrill (not illustrated) O. sativus F.d’A. Brotero (CÐE, testa, embryos) N. grahamii E.D. Merrill (not illustrated) Ostryocarpus J.D. Hooker (Millettieae) ...... 356 Notospartium J.D. Hooker (17.02) ...... 872 O. riparius J.D. Hooker (AÐC, fruit) N. carmichaeliae J.D. Hooker (CÐE, testa, embryos) Otholobium C.H. Stirton (12.08) ...... 712 O. caffrum (C.F. Ecklon & C.L.P. Zeyher) Olneya A. Gray (8.09) ...... 416 C.H. Stirton (not illustrated) O. tesota A. Gray (AÐE, fruits and valve, seeds, O. glandulosa (C. Linnaeus) J.W. Grimes testa, embryos) (not illustrated) O. hirtum (C. Linnaeus) C.H. Stirton (CÐE, Onobrychis P. Miller (18.06)...... 890 testa, embryos) O. melanotricha P.E. Boissier (CÐE, testa, O. pubescens (J.L.M. Poiret) J.W. Grimes embryos) (not illustrated) O. viciifolia J.A. Scopali (not illustrated) O. sericeum (J.L.M. Poiret) C.H. Stirton (not illustrated) Ononis C. Linnaeus (21.01)...... 914 O. natrix C. Linnaeus (not illustrated) Otion, see Gompholobium, p. 960 O. pubescens C. Linnaeus (CÐE, testa, embryos) Otoptera A.-P. de Candolle (10.56) ...... 564 Ophrestia H.M.L. Forbes (10.11) ...... 460 O. burchellii A.-P. de Candolle (AÐE, fruit, seeds, O. breviracemosa B. Verdcourt (not illustrated) testa, embryos) O. radicosa (A. Richard) B. Verdcourt (CÐE, testa, embryos) Oxylobium H.C. Andrews (24.09) ...... 974 O. cordifolium (H.C. Andrews) (not illustrated) Orbexilum C.S. Rafinesque-Schmaltz (12.07) ...... 710 O. microphyllum G. Bentham (not illustrated), O. onobrychis (T. Nuttall) P.A. Ryberg see Gompholobium, p. 960 (not illustrated) O. pultenaea A.-P. de Candolle (not illustrated) O. pedunculatum (P. Miller) A.M. Vail (CÐE, O. robustum J. Thompson (CÐE, testa, embryos) testa, embryos) Oxyrhynchus T.S. Brandegee (10.68)...... 596 Oreophysa (A.A. von Bunge ex P.E. Boissier) J.F.N. O. trinervis (J.D. Smith) V.E. Rudd (CÐE, testa,. Bornmüeller (16.06) ...... 828 embryos) O. microphylla (H.F. Jaubert & É. Spach) K. Browicz (AÐE, fruit, seed, testa, embryos) Oxytropis A.-P. de Candolle (16.17) ...... 854 O. campestris (C. Linnaeus) A.-P. de Candolle Ormocarpopsis R. Viguier (14.02) ...... 758 (not illustrated) O. aspera R. Viguier (AÐE, fruit, seed, testa, O. pilosa (C. Linnaeus) A.-P. de Candolle (CÐE, embryos) testa, embryos) O. riparia D.I. Litvinov (not illustrated)

1200 Pachecoa P.C. Standley & J.A. Steyermark P. cyphocalyx (A. Gray) P.A. Rydberg (14.23) ...... 800 (not illustrated) P. prismatica (M. Sessé y Lacastra & J.M. Mociño) P. esculentum (F.T. Pursh) P.A. Rydberg P.C. Standley & B.G. Schubert (AÐB, articles (CÐE, testa, embryos) and fruits, seeds) P. venezuelensis A. Burkart (CÐE, testa, embryos) Peltiera D.J. Du Puy & J.-N. Labat (14.02A) ...... 2

Pachyrhizus A. Richard ex A.-P. de Candolle Periandra C.F.P. von Martius ex G. Bentham (10.46) ...... 542 (10.15) ...... 470 P. erosus (C. Linnaeus) I. Urban (CÐE, testa, P. heterophylla G. Bentham (CÐE, testa, embryos) embryos)

Panurea R. Spruce ex G. Bentham (2.30) ...... 184 Pericopsis G.H.K. Thwaites (2.17) ...... 156 P. longifolia R. Spruce ex G. Bentham (AÐD, fruit, P. elata (H.A.T. Harms) M.S. Knaap-van testa, embryos) Meeuwen (CÐE, testa, embryos)

Papilionopsis C.G.G.J. van Steenis, see Desmodium, p. 652 Petaladenium A. Ducke (2.25) ...... 2

Paracalyx S.I. Ali (10.82) ...... 632 Petalostemon A. Michaux, see Cullen, p. 696 P. scariosus (W. Roxburgh) S.I. Ali (CÐE, testa, embryos) Peteria A. Gray (8.10) ...... 418 P. thompsoniae S. Watson (CÐE, testa, embryos) Paraderris (F.A.W. Miquel) R. Geesink, see Derris, p. 324 Petteria C.B. Presl (30.13) ...... 1101 Paramachaerium A. Ducke (4.10) ...... 250 P. ramentacea (F.W. Sieber) C.B. Presl (AÐE, P. gruberi G.K. Brizicky (not illustrated) fruit and valve, seeds, testa, embryos) P. krukovii V.E. Rudd (not illustrated) P. ormosioides (A. Ducke) A. Ducke Phaca C. Linnaeus, see Oxytropis, p. 854 (not illustrated) P. schomburgkii (G. Bentham) A. Ducke (BÐE, Phaseolus C. Linnaeus (10.72) ...... 606 seeds, testa, embryos) P. acutifolius A. Gray (CÐE, testa, embryos) P. schunkei V.E. Rudd (not illustrated) P. coccineus C. Linnaeus (not illustrated) P. leptostachyus G. Bentham (not illustrated) Parochetus F. Buchanan-Hamilton ex D. Don P. lunatus C. Linnaeus (not illustrated) (21.02) ...... 918 P. vulgaris C. Linnaeus (not illustrated) P. communis F. Buchanan-Hamilton ex D. Don (AÐE, fruits with calyx, seeds, testa, embryos) Phylacium J.J. Bennett (11.22) ...... 686 P. bracteosum J.J. Bennett (not illustrated) Parryella J. Torrey & A. Gray (6.03) ...... 278 P. majus H. Collett & W.B. Hemsley (CÐE, P. filifolia J. Torrey & A. Gray (AÐE, fruits with testa, embryos) calyx, seeds, testa, embryos) Phyllodium A.N. Desvaux (11.07) ...... 648 Pearsonia R.A. Dummer (27.01) ...... 1030 P. pulchellum (C. Linnaeus) A.N. Desvaux P. cajanifolia (C.E.O. Kuntze) R.M. Polhill (CÐE, testa, embryos) (CÐE, testa, embryos) Phyllota (A.-P. de Candolle) G. Bentham (24.19) ...... 992 Pediomelum P.A. Rydberg (12.03)...... 702 P. luehmannii F. von Mueller (not illustrated), P. argophyllum (F.T. Pursh) J.W. Grimes see Gompholobium, p. 960 (not illustrated) P. pleurandroides F.H.J. von Mueller (AÐE, P. castoreum (S. Watson) P.A. Rydberg fruits with calyx, seeds, testa, embryos) (not illustrated)

1201 Phylloxylon H.E. Baillon (9.01) ...... 424 Platymiscium J.R.T. Vogel (4.08) ...... 246 P. decipiens H.E. Baillon (not illustrated) P. curuense N. Zamora & B.B. Klitgaard P. xylophylloides (J.G. Baker) D.J. Du Puy, (not illustrated) J.-N. Labat, & B.D. Schrire (AÐE, fruits, P. filipes G. Bentham (CÐE, testa, embryos) damaged seed, testa, embryos). P. pinnatum (N. von Jacquin) A. Dugand (not illustrated) Physanthyllis P.E. Boissier, see Tripodion, p. 722 Platypodium J.R.T. Vogel (4.14)...... 260 Physostigma J.H. Balfour (10.52) ...... 554 P. elegans J.R.T. Vogel (AÐE, fruits, seeds, testa, P. venenosum J.H. Balfour (DÐF, testa, embryos) embryos)

Pickeringia T. Nuttall ex J. Torrey & A. Gray Platysepalum F.M.J. Welwitch ex J.G. Baker (29.06) ...... 1070 (Millettieae) ...... 362 P. montana T. Nuttall (A, fruit) P. hirsutum (S.T. Dunn) F.N. Hepper (CÐE, testa, P. montana T. Nuttall var. tomentosa (L.R. embryos) Abrams) I.M. Johnston (CÐE, testa, embryos) Podalyria C.L. von Willdenow (25.06)...... 1022 Pictetia A.-P. de Candolle (14.05) ...... 764 P. sericea (H.C. Andrews) R. Brown (A, CÐE, P. spinifolia (A.N. Desvaux) I. Urban (BÐE, seeds, valves, testa, embryos) testa, embryos) Podocytisus P.E. Boissier & T.H.H. von Heldreich Piptanthus R. Sweet (29.02) ...... 1060 (30.11) ...... 1096 P. nepalensis (W.J. Hooker) D. Don (BÐE, seeds, P. caramanicus P.E. Boissier & T.H.H. testa, embryos) von Heldreich (AÐE, fruits, seeds, testa, P. tomentosa A.R. Franchet (not illustrated) embryos)

Piscidia C. Linnaeus (Millettieae) ...... 358 Podolobium R. Brown, see Gompholobium, p. 960, P. mollis J.N. Rose (CÐE, testa, embryos) and Mirbelia, p. 978

Pisum C. Linnaeus (19.04) ...... 906 Podolotus J.F. Royle (13.08), see Cytisopsis, p. 716 P. fulvum J. Sibthorp & J.E. Smith (not illustrated) Poecilanthe G. Bentham (Millettieae) ...... 364 P. sativum C. Linnaeus subsp. elatius (C. von P. amazonica (W.A. Ducke) W.A. Ducke (B, DÐF, Steven ex F.A.M. von Bieberstein) fruit and valve, testa, embryos) P.F.A. Ascherson & K.O.R.P.P. Graebner P. falcata (J. Velloso de Miranda) A. Ducke (not (CÐE, testa, embryos) illustrated) P. effusa (J.E. Huber) W.A. Ducke (not illustrated) Plagiocarpus G. Bentham (23.03)...... 944 P. grandifora G. Bentham (not illustrated) P. axillaris G. Bentham (AÐE, fruit with calyx P. hostmannii (G. Bentham) G.J.H. Amshoff and valves, seeds, testa, embryos) (A, valve) P. itapuana G.P. Lewis (not illustrated) Platycelyphium H.A.T. Harms (2.38) ...... 200 P. subcordata G. Bentham (not illustrated) P. voense (H.G.A. Engler) H. Wild (AÐE, fruit, seeds, testa, embryos) Poiretia É.P. Ventenat (14.20) ...... 794 P. latifolia J.R.T. Vogel (not illustrated) Platycyamus G. Bentham (Millettieae) ...... 360 P. tetraphylla (J.L.M. Poiret) A.E. Burkart P. regnellii G. Bentham (AÐE, fruits, seed, testa, (CÐE, testa, embryos) embryos) Poitea É.P. Ventenat (8.05) ...... 402 Platylobium J.E. Smith (23.07) ...... 952 P. glaegeoides É.P. Ventenat (BÐE, seeds, testa, P. formosum J.E. Smith (CÐE, testa, embryos) embryos)

1202 Polhillia C.H. Stirton (30.04) ...... 1082 Psorothamnus P.A. Rydberg (6.06) ...... 284 P. pallens C.H. Stirton (CÐE, testa, embryos) P. arborescens (J. Torrey) R.C. Barneby (CÐE, testa, embryos) Pongamia É.P. Ventenat (Millettieae)...... 366 P. kingii (S. Watson) R.C. Barneby (not illustrated) P. pinnata (C. Linnaeus) J.B. Pierre (AÐE, fruit, seeds, testa, embryos) Pterocarpus N. von Jacquin (4.17)...... 266 P. angolensis A.-P. de Candolle (not illustrated) Pongamiopsis R. Viguier (Millettieae)...... 368 P. erinaceus J.L.M. Poiret (CÐE, testa, embryos) P. pervilleana (H.E. Baillon) R. Viguier (CÐE, P. indicus C.L. von Willdenow (not illustrated) testa, embryos) P. officinalis N. von Jacquin (not illustrated)

Priestleya A.-P. de Candolle, see Liparia, p. 1016, and Pterodon J.R.T. Vogel (3.03) ...... 224 Xiphotheca, p. 1008 P. polygalaeflorus G. Bentham (CÐE, testa, embryos) Pseudarthria R. Wight & G.A.W. Arnott (11.11)...... 658 P. fagifolia J.G. Baker (not illustrated) Ptycholobium H.A.T. Harms (Millettieae) ...... 370 P. hookeri R. Wight & G.A.W. Arnott (CÐE, P. biflorum (E.H.F. Meyer) R.K. Brummitt testa, embryos) (A, fruits) P. viscida (C. Linnaeus) R. Wight & G.A.W. P. contortum (N.E. Brown) R.K. Brummitt Arnott (not illustrated) (CÐE, testa, embryos)

Pseudeminia B. Verdcourt (10.30) ...... 504 Ptychosema G. Bentham ex J. Lindley (23.09) ...... 956 P. comosa (J.G. Baker) B. Verdcourt (AÐE, fruits, P. anomalum G. Bentham (AÐE, fruit, seed, testa, seeds, testa, embryos) embryos) P. pusillum G. Bentham (not illustrated) Pseudoeriosema L. Hauman (10.12) ...... 462 P. andongense (F.M.J. Welwitsch ex J.G. Baker) Pueraria A.-P. de Candolle (10.32) ...... 508 L. Hauman (CÐE, testa, embryos) P. montana (J. de Loureiro) E.D. Merrill var. lobata (C.L. von Willdenow) L.J.G. van der Pseudolotus K.H. Rechinger (13.09), see Cytisopsis, p. 821 Maesen & S. Almeida (CÐE, testa, embryos)

Pseudovigna (H.A.T. Harms) B. Verdcourt (10.31) ..... 506 Pultenaea J.E. Smith (24.23) ...... 1000 P. argentea (C.L. von Willdenow) B. Verdcourt P. incurvata A. Cunningham (not illustrated) (AÐE, fruits, seeds, testa, embryos) P. neurocalyx P.K.N.S. Turczanivow (not illustrated) Psophocarpus N.J. de Necker ex A.-P. de Candolle P. obcordata (R. Brown) G. Bentham (CÐE, (10.51) ...... 552 testa, embryos) P. palustris A.N. Desvaux (CÐE, testa, embryos) Pycnospora R. Brown ex R. Wight & G.A.W. Arnott Psoralea C. Linnaeus (12.09) ...... 714 (11.12) ...... 660 P. aphylla C. Linnaeus (DÐE, testa) P. lutescens (J.L.M. Poiret) A.K. Schindler (AÐE, P. pinnata C. Linnaeus (AÐC, F, calyx with a fruits, seeds, testa, embryos) fruit inside and a cupulum, fruit without calyx, seeds, embryos) Pyranthus D.J. Du Puy & J.-N. Labat (Millettieae) ...... 372 P. alasoa D.J. Du Puy & J.-N. Labat (A, CÐF, Psoralidium P.A. Rydberg (12.04) ...... 704 fruits, seeds, testa, embryos) P. lanceolatum (F.T. Pursh) P.A. Rydberg (CÐE, P. ambatoana (H.E. Baillon) D.J. Du Puy & J.-N. testa, embryos) Labat (B, fruits) P. tenuifolium (F.T. Pursh) P.A. Rydberg (not illustrated)

1203 Rafnia C.P. Thunberg (27.05) ...... 1038 Rothia C.H. Persoon (27.02) ...... 1032 R. amplexicaulis (C. Linnaeus) C.P. Thunberg R. indica (C. Linnaeus) G.C. Druce (CÐE, testa, (CÐE, testa, embryos) embryos)

Ramirezella J.N. Rose (10.66A) ...... 592 Rupertia J.W. Grimes (12.05) ...... 706 R. strobilophora (B.L. Robinson) J.N. Rose (A, CÐE, R. hallii (P.A. Rydberg) J.W. Grimes (not fruits, testa, embryos) illustrated) R. physodes (D. Douglas ex W.J. Hooker) J.W. Ramorinoa C.L. Spegazzini (4.11 ...... 252 Grimes (AÐE, fruits, seed, testa, embryos) R. girolae C.L. Spegazzini (AÐE, fruits, seed, R. rigida (S.B. Parish) J.W. Grimes (not illustrated) testa, embryos) Sakoanala R. Viguier (2.35) ...... 194 Rehsonia L.R. Stritch, see Wisteria, p. 384 S. madagascariensis R. Viguier (CÐE, testa, embryos) Requienia A.-P. de Candolle (Millettieae) ...... 374 S. villosa R. Viguier (B, seeds) R. sphaerosperma A.-P. de Candolle (BÐE, seeds, testa, embryos) Salweenia E.G. Baker (2.42) ...... 210 S. wardii E.G. Baker (AÐE, fruits, seeds, testa, Retama C.S. Rafinesque-Schmaltz (30.21) ...... 1118 embryos) R. sphaerocarpa (C. Linnaeus) P.E. Boissier (CÐE, testa, embryos) Sarcodum J. de Loureiro (Millettieae)...... 376 S. binnendyckianum (W.S. Kurz) R. Geesink Rhodopis I. Urban (10.09) ...... 456 (DÐF, testa, embryos) R. lowdenii W.S. Judd (A, B, G, fruits, embryos) S. scandens J. de Loureiro (AÐC, fruit, seed) R. planisiliqua (C. Linnaeus) I. Urban (CÐF, fruits, seeds, testa) Sartoria P.E. Boissier & T.H.H. von Heldreich (18.05) ...... 2 Rhynchosia J. de Loureiro (10.80) ...... 626 R. baukea J.-N. Labat & D.J. Du Puy (not Schefflerodendron H.A.T. Harms (Millettieae) ...... 378 illustrated) S. usambarense H.A.T. Harms (BÐE, seed, testa, R. phaseoloides (O.P. Swartz) A.-P. de Candolle embryos) (CÐE, testa, embryos) Scorpiurus C. Linnaeus (13.14)...... 744 Rhynchotropis H.A.T. Harms (9.03) ...... 426 S. muricatus C. Linnaeus (CÐE, testa, embryos) R. poggei (P.H.W. Taubert) H.A.T. Harms S. muricatus var. muricatus (not illustrated) (AÐE, fruit, seeds, testa, embryos) S. muricatus var. subvillosus (C. Linnaeus) J.B.A.P.M. de Lamarck (not illustrated) Riedeliella H.A.T. Harms (2.08) ...... 134 S. subvillosus C. Linnaeus (not illustrated) R. magalhaesii (C.T. Rizzini) H.C. de Lima & S. sulcatus C. Linnaeus (not illustrated) A.M.S. de Fonseca Vaz (CÐE, testa, embryos) S. vermiculatus C. Linnaeus (not illustrated)

Robinia C. Linnaeus (8.07) ...... 410 Securigera A.P. de Candolle (13.12)...... 740 R. pseudoacacia C. Linnaeus (CÐE, testa, S. securidaca (C. Linnaeus) A. von Degen & I. embryos) Doerfler (CÐE, testa, embryos)

Robynsiophyton R. Wilczek (27.03) ...... 1034 Sellocharis P.H.W. Taubert (30.07) ...... 2 R. vanderystii R. Wilczek (AÐE, fruits, seeds, testa, embryos) Sesbania G.A. Scopoli (8.01) ...... 388 S. sect. Glottidium (A.N. Desvaux) M. Lavin (not illustrated), see Glottidium, p. 392 S. emerus (J.B.C.F. Aublet) I. Urban (CÐE, testa, embryos)

1204 Shuteria R. Wight & G.A.W. Arnott (10.41) ...... 530 Sphinctospermum J.N. Rose (8.12) ...... 422 S. involucrata (N. Wallich) R. Wight & G.A.W S. constrictum (S. Watson) J.N. Rose (AÐE, fruits, Arnott (CÐE, testa, embryos) seeds, testa, embryos)

Sinodolichos B. Verdcourt (10.34) ...... 514 Spirotropis E.L.R. Tulasne (2.32) ...... 2, 188 S. lagopus (S.T. Dunn) B. Verdcourt (AÐE, fruits, S. longifolia (A.-P. de Candolle) H.E. Baillon (A, seed, testa, embryos) fruits)

Smirnowia A.A. von Bunge (16.08) ...... 832 Spongiocarpella G.P. Yakovlev & N. Ulziykhutag S. turkestana A.A. von Bunge (AÐE, fruit, seeds, (16.13) ...... 2 testa, embryos) Stauracanthus J.H.F. Link (30.24) ...... 1126 Smithia W. Aiton (14.12) ...... 780 S. boininii (P.B. Webb) G.A. da S.F. Sampaio S. elliotti E.G. Baker var. elliotti (B, young fruit) (A, inner and outer valve surfaces) S. purpurea W.J. Hooker (DÐF, testa, embryos) S. genistoides (F.d’A. Brotero) G.A. da S.F. Sampaio (CÐE, testa, embryos) Soemmeringia C.F.P. von Martius (14.10) ...... 776 S. semperflorens C.F.P. von Martius (AÐF, partial Stirtonanthus B.-E. Van Wyk & A.L. Schutte fruit, fruit and fruits in marcescent corollas, (25.07) ...... 2 seeds, testa, embryos) Stizolobium P. Browne, see Mucuna, p. 440 Sophora C. Linnaeus (2.45)...... 214 S. tomentosa C. Linnaeus (CÐE, testa, embryos) Stracheya G. Bentham (18.04) ...... 888 S. tibetica G. Bentham (AÐE, fruits, seeds, testa, Spartidium A.N. Pomel (27.04) ...... 1036 embryos) S. saharae (E.S.-C. Cosson & E. Reboul) A.N. Pomel (AÐE, fruit, seed, testa, embryos) Streblorrhiza S.F.L. Endlicher (17.01) ...... 870 S. speciosa S.F.L. Endlicher (AÐE, fruit, seed, Spartium C. Linnaeus (30.19) ...... 1114 testa, embryos) S. junceum C. Linnaeus (AÐE, dehisced fruit and valve, seeds, testa, embryos) Strongylodon J.R.T. Vogel (10.02) ...... 438 S. loheri S.-F. Huang (not illustrated) Spathionema P.H.W. Taubert (10.55) ...... 562 S. macrobotrys A. Gray (CÐE, testa, embryos) S. kilimandscharicum P.H.W. Taubert (AÐE, fruits, seeds, testa, embryos) Strophostyles S. Elliott (10.70) ...... 600 S. helvula (C. Linnaeus) S. Elliott (CÐE, testa, Spatholobus J.C. Hasskarl (10.05) ...... 446 embryos) S. suberectus D.B. Dunn (CÐE, testa, embryos) Stylosanthes O.P. Swartz (14.25) ...... 804 Sphaerolobium J.E. Smith (24.02) ...... 964 S. guianensis (J.B.C.F. Aublet) O.P. Swartz S. grandiflorum G. Bentham (not illustrated) (DÐF, testa, embryos) S. vimineum J.E. Smith (CÐE, testa, embryos) Styphnolobium H.W. Schott (2.44)...... 212 Sphaerophysa A.-P. de Candolle (16.07) ...... 830 S. affine (J. Torrey & A. Gray) W.G. Walpers S. kotschyana P.E. Boissier (not illustrated) (CÐE, testa, embryos) S. salsula (P.S. von Pallas) A.-P. de Candolle (AÐE, fruits, seeds, testa, embryos) Sutherlandia R. Brown (16.03) ...... 822 S. frutescens (C. Linnaeus) R. Brown (BÐE, seeds, Sphenostylis E.H.F. Meyer (10.57) ...... 566 testa, embryos) S. stenocarpa (C.F. Hochstetter ex A. Richard) H.A.T. Harms (CÐE, testa, embryos)

1205 Swainsona R.A. Salisbury (16.02) ...... 818 Teramnus P. Browne (10.36)...... 520 S. novae-zelandiae J.D. Hooker (not illustrated) T. uncinatus (C. Linnaeus) O.P. Swartz (CÐE, S. maccullochiana F.H. Mueller (CÐE, testa, testa, embryos) embryos) Terua P.C. Standley & F.J. Hermann, see Lonchocarpus, Swartzia J.C.D. von Schreber (1.01) ...... 78 p. 344 S. fistuloides H.A.T. Harms (not illustrated), and see Bobgunnia, p. 84 Tetragonolobus G.A. Scopoli, see Lotus, p. 730 S. latifolia G. Bentham var. sylvestris R.S. Cowan (IÐL, embryos, embryos magnified) Teyleria C.A. Backer (10.39) ...... 526 S. madagascariensis A.N. Desvaux (not illustrated), T. koordersii (C.A. Backer) C.A. Backer (AÐE, and see Bobgunnia, p. 84 fruits, seeds, testa, embryos) S. oraria R.S. Cowan (GÐH, embryos, embry magnified) Thermopsis R. Brown (29.04) ...... 1064 S. panamensis G. Bentham (not illustrated) T. inflata J. Cambessedes (not illustrated) S. simplex (O.P. Swartz) C.P.J. Sprengel (CÐF, T. divaricarpa A. Nelson (not illustrated) testa, embryos, embryos magnified) T. gracilis T.J. Howell (not illustrated) T. lupinoides (C. Linnaeus) J.H.F. Link (CÐE, Sweetia C.P.J. Sprengel (2.02) ...... 122 testa, embryos) S. fruticosa C.P.J. Sprengel (AÐE, fruits, seeds, T. rhombifolia (T. Nuttall ex F.T. Pursh) J. testa, embryos) Richardson (not illustrated)

Tadehagi H. Ohashi (11.13) ...... 662 Tipuana (G. Bentham) G. Bentham (4.13) ...... 258 T. rogeri (A.K. Schindler) H. Ohashi T. tipu (G. Bentham) C.E.O. Kuntze (AÐE, fruits (not illustrated) seed, testa, embryos) T. triquetrum (C. Linnaeus) H. Ohashi (AÐE, fruits, seeds, testa, embryos) Trifidacanthus E.D. Merrill (11.05) ...... 644 T. triquetrum subsp. rodgeri (A.K. Schindler) T. unifoliolatus E.D. Merrill (AÐE, fruit and article, H. Ohashi (not illustrated) seed, testa, embryos)

Taralea J.B.C.F. Aublet (3.02) ...... 222 Trifolium C. Linnaeus (21.06) ...... 932 T. crassifolia (G. Bentham) W.A. Ducke (CÐE, T. sect. Lotoidea H.J.N. von Crantz (not illustrated) testa, embryos, external embryo) T. pannonicum N. von Jacquin (CÐE, testa, T. oppositifolia J.C.B.F. Aublet (not illustrated) embryos) T. repens C. Linnaeus (not illustrated), Taverniera A.-P. de Candolle (18.03) ...... 886 see Adesmia, p. 810 T. lappacea (P. Forsskål) A.-P. de Candolle T. subterraneum C. Linnaeus (not illustrated) (CÐE, testa, embryos) Trigonella C. Linnaeus (21.04) ...... 924 Teline F.K. Medikus, see Genista, p. 1120 T. foenum-graecum C. Linnaeus (CÐE, testa, embryos) Templetonia R. Brown (23.01) ...... 938 T. biloba (G. Bentham) R.M. Polhill (not illustrated) Tripodion F.C. Medikus (13.03)...... 722 T. incana J.H. Ross (not illustrated) T. teraphyllum (C. Linnaeus) J.P. Fourreau (AÐE, T. retusa (É.P. Ventenat) R. Brown (CÐE, testa, fruits and fruiting calyx, seeds, testa, embryos) embryos) Uleanthus H.A.T. Harms (2.29) ...... 182 Tephrosia C.H. Persoon (Millettieae) ...... 380 U. erythrinoides H.A.T. Harms (AÐG, fruit, seed, T. purpurea (C. Linnaeus) C.H. Persoon (CÐE, testa, embryos, magnified embryo) testa, embryos)

1206 Ulex C. Linnaeus (30.25)...... 1128 Vermifrux J.B. Gillett (13.10) ...... 736 U. parviflorus P.A. Pourret de Figeac (CÐE, testa, V. abyssinicum (A. Richard) J.B. Gillett (AÐE, embryos) fruits, seeds, testa, embryos), and see ...... Dorycnopsis, p. 726 Uraria A.N. Desvaux (11.16)...... 672 U. cordifolia C.F.W. Wallroth (not illustrated) Vicia C. Linnaeus (19.01) ...... 894 U. picta (N. von Jacquin) A.N. Desvaux ex A.-P. V. subgen. Vicia (not illustrated) de Candolle (CÐE, testa, embryos) V. sect. Hypechusa (F.G.C. Alefeld) P.F.A. Acherson & K.O.R.P.P. Graebner (not illustrated) Urariopsis S.S. Schindler, see Uraria, p. 672 V. ervilia (C. Linnaeus) C.L. von Willdenow U. cordifolia (C.F.W. Wallroth) A.K. Schindler (not illustrated) (not illustrated), see Uraria, p. 672 V. faba C. Linnaeus (CÐE, testa, embryos) V. menziesii C.P.J. Sprengel (not illustrated) Uribea A. Dugand & R.C. Romero-Castañeda V. montbretii F.E.L. von Fischer & C.A.A. (2.04) ...... 126 von Meyer (not illustrated), see Lens, p. 902 U. tamarinoides A. Dugand & R.C. Romero- V. sativa C. Linnaeus (F, embryos) Castañeda (AÐE, fruits, seeds, testa, embryos) V. sativa subsp. amphicarpa (C. Linnaeus) J.A. Battandier (not illustrated) Urodon P.K.N.S. Turczaninow (24.22) ...... 998 U. capitatus P.K.N.S. Turczaninow (AÐE, fruits Vigna C.G. Savi (10.66) ...... 588 with calyx, seeds, testa, embryos) V. pilosa (J.T. Klein ex C.L. von Willdenow) J.G Baker (not illustrated) Vandasia K. Domin (Fabaceae), see Vandasina, p. 548 V. unguiculata (C. Linnaeus) G.W. Walpers subsp. unguiculata (CÐE, testa, embryos) Vandasia J. Velenovsky (Gasteromycetes), see Vandasina, p. 548 Viminaria J.E. Smith (24.03) ...... 966 V. juncea (H.A. Schader) J.C. von Hoffmannsegg Vandasina S. Rauschert (10.49) ...... 548 (AÐE, fruits with calyx, seeds, testa, embryos) V. retusa (G. Bentham) S. Rauschert (AÐE, fruits, seeds, testa, embryos) Virgilia J.L.M. Poiret (25.08) ...... 1024 V. oroboides (B. Bergius) T.M. Salter (A, CÐE, Vatairea J.B.C.F. Aublet (4.02) ...... 230 valves, testa, embryos) V. guianensis J.B.C.F. Aublet (not illustrated) V. lundellii (P.C. Standley) P.C. Standley (CÐE, Wajira M. Thulin, see Vigna, p. 588 endocarp (functioning as testa), embryos) Weberbauerella O.E. Ulbrich (14.18) ...... 2 Vataireopsis W.A. Ducke (4.01) ...... 226 V. speciosa W.A. Ducke (AÐE, fruits, seed, testa, Wiborgia C.P. Thunberg (27.11) ...... 1054 embryos) W. humilis (C.P. Thunberg) R. Dalhgren (not illustrated) Vatovaea E. Chiovenda (10.53)...... 558 W. leptoptera R. Dahlgren (not illustrated) V. pseudolablab (H.A.T. Harms) J.B. Gillett (AÐE, W. monoptera E.H.F. Meyer (not illustrated) fruits, seeds, testa, embryos) W. mucronata (C. Linnaeus) A.P. Druce (CÐE, testa, embryos) Vaughania S. Moore (9.02) ...... 2 W. obcordata (B. Bergius) C.P. Thunberg (not illustrated) Vavilovia A.A. Fedorov (19.05) ...... 908 W. sericea C.P. Thunberg (not illustrated) V. formosa (C. von Steven) A.A. Federov (AÐE, fruits, seeds, testa, embryos) Willardia J.N. Rose, see Lonchocarpus, p. 344

1207 Wisteria T. Nuttall (Millettieae) ...... 384 W. frutescens (C. Linnaeus) J.L.M. Poiret (CÐE, testa, embryos)

Xanthocercis H.E. Baillon (2.14) ...... 148 X. rabiensis L.J.G. van der Maesen (not illustrated) X. zambesiaca (E.G. Baker) N. Dumaz-le-Grand (CÐE, testa, embryos)

Xeroderris G.E. Roberty (Millettieae)...... 386 X. stuhlmannii (P.H.W. Taubert) F. de A. Mendonça & E.P. Sousa (AÐE, fruit, seeds, testa, embryos)

Xiphotheca C.F. Ecklon & J.M. Zeyher (25.01) ...... 1008 X. fruticosa (C. Linnaeus) A.L. Schutte & B.-E. van Wyk (AÐE, fruits, seeds, testa, embryos)

Yucaratonia A.E. Burkart, see Gliricidia, p. 406

Zabrotes subfasciatus, a bruchid, see Phaseolus, p. 606

Zollernia M.A.P. zu Wied-Neuwied & C.G.D. Nees von Esenbeck (1.11) ...... 108 Z. ilicifolia (A.T. Brongniart) J.R.T. Vogel (B, E seed, embryos) Z. kanukuensis R.S. Cowan (CÐD, testa) Z. magnifica A.M. de Carvalho & R.C. Barneby (not illustrated) Z. tango P.C. Standley (not illustrated), see Lecointea, p. 102

Zornia J.F. Gmelin (14.21) ...... 796 Z. glochidiata H.G.L. Reichenbach ex A.-P. de Candolle (CÐE, testa, embryos)

1208