Psoraleeae (12.01Ð12.09) faboid split; punctiform; marginal according to radicle tip; recessed; within corona. Hilum corona color lighter Genus: 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 longitu- dinal 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, (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).

698 A

B E

C D

699 Genus: 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 pubes- cence 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 meso- carp 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).

700 A

B E

C D

701 Genus: 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.

702 A

B E

C D

703 Genus: 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).

704 A

B

E

C D

705 Genus: 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 modi- fied 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).

706 A

B E

C D

707 Genus: 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).

708 A

B E

C D

709 Genus: 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 .” 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).

710 A

B E

C D

711 Genus: 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).

712 A

B E

C D

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).

714 A

B

C F

D E

715