Cordyla

Taxonomy

Cordyla J. de Loureiro Fl. Cochinch. 402, 411. Sep 1790.

Subfamily: Faboideae.
Phylogenetic Number: 3.1.06.
Tribe: Swartzieae.
Group: Aldina.
Species Studied - Species in Genus: 4 studied; 5 in genus.

Description

Fruit: A legume; unilocular; 4–10 cm long; 3–3.5 cm wide; 2.5–3.5 cm thick; length less than twice as long as width, or 2–9 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; not plicate; not twisted; asymmetrical, or symmetrical; (sub-) circular, or ovate, or oblong; with both sutures parallelly curved, or both sutures nearly straight, or 1 straight and 1 curved suture; widest near middle or D-shaped; not inflated; terete; without beak, or with beak; declined; with solid beak the same color and texture as fruit; rounded at apex, or tapered at apex; aligned with longitudinal axis of fruit, or oblique with longitudinal axis of fruit, or right-angled with longitudinal axis of fruit; short tapered at base; aligned with longitudinal axis of fruit, or oblique with longitudinal axis of fruit; with the apex and base uniform in texture; ligneous (when fresh pulpy within between the seeds (Capuron, 1968Capuron, 1968:
Capuron R. 1968. Contributions a l'étude de la flore forestière de Madagascar. Adansonia 8(ser. 2): 11&-16, 17&-37, 189&-222.)); seed chambers externally invisible, or visible; seed chambers with the raised seed chambers not torulose; margin not constricted; margin without sulcus; margin plain; wing(s) absent; stipitate, or substipitate; with the stipe 0.1–25 mm long; indehiscent. Replum invisible. Epicarp dull; monochrome; dark reddish to greenish brown; with surface texture uniform; glabrous; eglandular; without spines; not smooth; with elevated features; not veined; not tuberculate; ribbed; not exfoliating; without cracks; without embedded tissue, much thicker than epicarp, running from base to apex. Mesocarp present; thick; surface not veined; 1-layered; without balsamic vesicles; without fibers; without reniform canals; solid, or fleshy; ligneous (or subligneous). Endocarp present (but testa fused to endocarp); visible; opaque; without adhering pieces of testa; nonseptate; coriaceous; not exfoliating; separating with mesocarp from epicarp; without wings; entire. Seed(s) 1–6; length parallel with fruit length, or oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus of 1 length only; thick; straight. Aril absent.

Seed: 20–30 mm long; 15–22 mm wide; 9–16 mm thick; overgrown, 1 seed filling entire fruit cavity (Corner 1976); angular; asymmetrical; ovate, or reniform; compressed, or terete; 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 absent; partially adhering to endocarp. Endosperm absent. Cotyledons not smooth; somewhat wrinkled; 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; completely concealing radicle; entire over radicle; without lobes; with the interface division terminating at base of radicle; without margins recessed; reddish tan, or brown (reddish); 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; triangular; lobe tip straight; straight with embryonic axis; centered between cotyledons; much less than 1/2 length of cotyledons. Plumule well developed; glabrous.

Distribution

Tropical Africa.

Old World; tropical Africa.

Generic Notes

In Herendeen'S (1994) cladistic analyses Cordyla and Mildbraediodendron (3.1.7) formed a well defined clade supported by five characters. Within Swartzieae and Sophoreae (3.2) some of their seed structures are also unique: well developed, straight embryos at the center of the cotyledons which are deeply divided almost to their center. Twenty fruit and seed characters, as well as distribution, separate Cordyla from Dupuya: Cordyla with fruit with beak declined, short tapered at base; mesocarp 1-layered; endocarp coriaceous, separating with mesocarp from epicarp; aril absent; seed overgrown, 15–22 mm wide, angular, asymmetrical; testa absent; endosperm absent; cotyledons wrinkled, both outer faces convex, not folded, completely concealing radicle, entire over radicle, inner face flat; embryonic axis parallel to length of seed; radicle triangular; distribution tropical Africa, versus Dupuya with fruit with beak straight, rounded or tapered at base; mesocarp 2-layered; endocarp spongy, remaining fused to mesocarp and epicarp; aril present; seed not overgrown, 8–12 mm wide, not angular, symmetrical; testa present; endosperm present; cotyledons 1–6 grooves on each face, outer face of one cotyledon flat and of other cotyledon concave, with both folded, partially concealing radicle, notched at radicle, inner face with central ridge on 1 cotyledon and central groove on other; embryonic axis oblique to length of seed, radicle bulbose; distribution endemic to Madagascar. In our opinion, Dupuya is a good genus. The parallels between Swartzia-Bobgunnia (3.1.0 and 3.1.0) and Cordyla-Dupuya are striking. In both cases, the former have over grown seeds with all the characteristics of that condition, and the latter have typical faboid seeds.

Tribal Notes

Tribe Swartzieae

Swartzieae has been assigned to Caesalpinioideae (Cowan, 1968), Swartzioideae (de Candolle, 1825a, 1825b; Corner, 1976), and Faboideae (Bentham, 1865, Hutchinson, 1964, Cowan, 1981). In 1968, Cowan (1968) was unable "finally to resolve the sub-familial relationship of Swartzia," but in 1981, he (Cowan 1981) placed it in the Faboideae and stated, "... features appear to support the arrangement adopted here (Cowan 1981) with the Swartzieae representing a relatively less-advanced position in the Papilionoideae (Faboideae). This conclusion is now supported by wood anatomy ..., by nodulation proclivity ..., and by chemistry ...; chromosome numbers of n=8, 10 or 14 ..., as well as pollen morphology ..., do not negate this conclusion." In the most recent assessment of the Fabaceae, Polhill (1994a, 1994b) maintained Swartzieae as a basal tribe of Faboideae, "transitional to the Caesalpinioideae." He transferred four genera from Sophoreae (2) into the Swartzieae, Amburana (3.1.15), Ateleia (3.1.13), Cyathostegia (3.1.14), and Holocalyx (3.1.12), and arranged the genera in four groups corresponding to clades in Herendeen's (1994) cladistic analysis. Herendeen carried out cladistic analyses using morphological characters of all Swartzieae genera, 19 genera of Sophoreae, and three Caesalpinioideae genera. He concluded that Swartzieae is polyphyletic and that it should be disbanded and its genera transferred to Sophoreae. Preliminary rbcL data (Doyle et al. 1997) supported his conclusions. Ireland et al. (2000) also carried out molecular phylogenetic studies. They also concluded that Swartizeae is polyphyletic, and suggested that possibly tribe Swartzieae could be maintained with Swartzia (3.1.01), Bobgunnia (3.1.01A), Bocoa (3.1.02), Ateleia (3.1.13), Cyathostegia (3.1.14) and the current remaining Swartzieae genera transferred to other tribes. Our seed data neither support nor refute the overall outlines of Herendeen's cladograms; they are discussed below for a few genera. Ferguson and Skvarla (1991) reported on the pollen morphology of Aldina and Swartzia (1.01), and the nine other genera of Swartzieae are covered in Ferguson and Skvarla (1988). Their data are summarized in a computer-generated key in Vezey et al. (1991). The pollen data for the tribe should be compared with our seed-fruit morphological data.