Entada

Taxonomy

Entada M. Adanson Nom. cons. Fam. 2: 318, 554. Jul-Aug 1763.

Subfamily: Mimosoideae.
Phylogenetic Number: 2.3.14.
Tribe: Mimoseae.
Group: Entada.
Species Studied - Species in Genus: 13 studied; ca. 30 in genus.

Description

Fruit: A legume; unilocular; 1–200 cm long; 2–15 cm wide; 0.3–3 cm thick; length less than twice as long as width, or 2–9 times longer than wide, or more than 9 times longer than wide; with deciduous androecial sheath; with deciduous corolla; with deciduous calyx; without orifice formed by curving of fruit or fruit segments; straight to 1-coiled; not plicate; twisted, or not twisted; symmetrical, or asymmetrical; oblong, or falcate; with both sutures parallelly curved; not inflated; compressed to terete; without beak; rounded at apex; aligned with longitudinal axis of fruit; short tapered at base; aligned with longitudinal axis of fruit; with the apex and base uniform in texture; chartaceous to ligneous (rarely fleshy when fresh); seed chambers externally visible; margin constricted; margin slightly constricted along both margins to constricted along both margins, or constricted on 1 margin and slightly constricted on the other margin (dorsal and ventral margins respectively); margin without sulcus; margin plain; wing(s) absent; stipitate; with the stipe up to 35 mm long; indehiscent. Replum visible, or invisible. Epicarp dull; monochrome; brown; with surface texture uniform; glabrous; eglandular; without spines; not smooth; with elevated features; veined; reticulately veined; not tuberculate; exfoliating; without cracks; without embedded tissue, much thicker than epicarp, running from base to apex. Mesocarp present, or absent; surface not veined; 1-layered; without balsamic vesicles; with fibers; without reniform canals; fibrous throughout; ligneous. Endocarp present; visible; dull; opaque; monochrome, or mottled; tan; with mottling more or less uniform (darker in most seed chambers for most winged endocarps); with brown overlay; smooth; without adhering pieces of testa; septate to nonseptate; sub coriaceous (when segments winged), or ligneous (when segments not winged); not exfoliating; remaining fused to mesocarp and epicarp, or separating from mesocarp (in largest fruits); remaining fused to epicarp, or separating from epicarp (in largest fruits); entire, or separating into 1-seeded winged segments (and separating from replum). Seed(s) 6–14; length transverse to fruit length to oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus 15–33 mm long; of 1 length only; filiform; hooked to plicate. Aril absent.

Seed: 5–80 mm long; 3–70 mm wide; 1–40 mm thick; not overgrown; not angular; symmetrical; circular to oblong, or irregular; terete to 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 present; without pieces of adhering epicarp; not adhering to endocarp; free from endocarp; glossy to dull; not modified by a bloom; colored; monochrome; dark brown (occasionally with adnate tan to ocher patches of endocarp tissue); glabrous; smooth to not smooth; with elevated features; rugose; osseous to coriaceous. Pleurogram present, or absent; 90–100 %. Pseudopleurogram absent. Fracture lines present, or absent. Rim absent. Wing(s) absent. Raphe not visible. Hilum present; visible; without faboid split; punctiform, or larger than punctiform; up to 15 mm long (on some of largest seeds); with curved outline to straight outline; elliptic; linear; apical at apex of radicle tip; flush to recessed; not within corona, halo, or rim. Lens not discernible (on large pleurogrammatic seeds), or discernible (on smaller nonpleurogrammatic seeds); 0.5–0.7 mm long; with margins straight; oblong; not in groove of raphe; mounded; similar color as testa; lighter than testa; tan; not within corona, halo, or rim. Endosperm absent. Cotyledons smooth; both outer faces flat; both the same thickness; both more or less of equal length; not folded; margin entire 180 degrees from base of radicle; similar at apex; partially concealing radicle, or not concealing radicle; notched at radicle; with lobes; with lobes touching (auriculate), or not touching; without basal groin formed by lobes; with the interface division terminating at base of radicle; without margins recessed; inner face flat; glabrous on inner face. Embryonic axis straight; parallel to length of seed; without a joint evident between the radicle and the cotyledons. Radicle differentiated from cotyledon; straight with embryonic axis; centered between cotyledons. Plumule moderately developed; glabrous.

Distribution

Pantropic and subpantropic (Most numerous in Africa).

New World and Old World; pantropical and pansubtropical (most common in Africa); Mexico, West Indies, Central America, and South America; Peru, Brazil, Ecuador, and the Guianas; Africa, Madagascar, India, Indochina, China, Japan, Indonesia and the Philippines, Australia, Fiji, and Hawaii.

Generic Notes

Fruit and seed characters in this genus are divisive and are used to subdivide the genus. Brenan (1966) in revising Entada for "Flora Zambesiaca" described sections and subsections for all Entada spp. of the world but assigned only African species to his two subgenera (Entada and Acanthentada Brenan, now the genus Pseudoentada N.L. Britton and J.N. Rose). Brenan opined that "there is not sufficient justification for recognizing generic segregates from Entada in Africa, at least for the time being." Lewis and Elias (1981) recognized Entada and Pseudoentada. Perhaps a critical revision of Entada will resolve the problem created by having species with such disparate fruit and seed characters in the same genus. The seeds of E. gigas and E. phaseoloides are wide ranging drift seeds (Gunn et al., 1976).

 Fruit and seed:  E. abyssinica  Steudel ex A. Richard - bottom left entire fruit (background) and partial fruit (foreground);  E. glandulosa  Pierre ex Gagnepain - bottom right fruit segments;  E. phaseoloides  (C. Linnaeus) Merrill - top center partial seed in situ.
Fruit and seed: E. abyssinica Steudel ex A. Richard - bottom left entire fruit (background) and partial fruit (foreground); E. glandulosa Pierre ex Gagnepain - bottom right fruit segments; E. phaseoloides (C. Linnaeus) Merrill - top center partial seed in situ.
 Fruit:  E. africana  Guillemin & Perrottet - bottom right fruit segment;  E. gigas  (C. Linnaeus) Fawcett & Rendle - top left partial seed in situ, bottom left fruit;  E. polystacha  (C. Linnaeus) de Candolle - top right partial fruit showing 1-seeded fruit segments, indurate replum, seed in situ within fruit segment.
Fruit: E. africana Guillemin & Perrottet - bottom right fruit segment; E. gigas (C. Linnaeus) Fawcett & Rendle - top left partial seed in situ, bottom left fruit; E. polystacha (C. Linnaeus) de Candolle - top right partial fruit showing 1-seeded fruit segments, indurate replum, seed in situ within fruit segment.
 Seed, cotyledon, embryo, and testa:  E. abyssinica  Steudel ex A. Richard - top center seed topography, bottom right testa SEM;  E. gigas  (C. Linnaeus) Fawcett & Rendle - bottom far left seed topography, bottom left center seed in longitudinal section showing cavity between cotyledon, top right testa SEM;  E. glandulosa  J.B.L. ex Gagnepain - top left center seed topography;  E. polystacha  (C. Linnaeus) de Candolle - top far left cotyledon concealing all but tip of radicle (L) and embryonic axis (R);  E. pursaetha  de Candolle - bottom center seed;  E.  spp. - center seeds.
Seed, cotyledon, embryo, and testa: E. abyssinica Steudel ex A. Richard - top center seed topography, bottom right testa SEM; E. gigas (C. Linnaeus) Fawcett & Rendle - bottom far left seed topography, bottom left center seed in longitudinal section showing cavity between cotyledon, top right testa SEM; E. glandulosa J.B.L. ex Gagnepain - top left center seed topography; E. polystacha (C. Linnaeus) de Candolle - top far left cotyledon concealing all but tip of radicle (L) and embryonic axis (R); E. pursaetha de Candolle - bottom center seed; E. spp. - center seeds.