Dupuya

Taxonomy

Dupuya J.H. Kirkbride, ined.

Subfamily: Faboideae.
Phylogenetic Number: 3.1.06A.
Tribe: Swartzieae.
Group: Aldina.
Species Studied - Species in Genus: 2 studied; 2 in genus.

Description

Fruit: A legume; unilocular; 3.5–10 cm long; 2–3.5 cm wide; 2–3.5 cm thick; length less than twice as long as width to 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; symmetrical; elliptic to oblong; not inflated; terete; with beak (5–8 mm long), or without beak; straight; with solid beak the same color and texture as fruit; tapered at apex, or rounded at apex; aligned with longitudinal axis of fruit; tapered at base, or rounded at base; aligned with longitudinal axis of fruit; with the apex and base uniform in texture; drupaceous (when fresh, Du Puy et al. [2002]), or ligneous (when dry); seed chambers externally invisible, or visible; seed chambers with the raised seed chambers not torulose; margin not constricted, or constricted (when 2 or more seeds in fruit); margin slightly constricted along both margins; margin without sulcus; margin plain; wing(s) absent; stipitate; with the stipe 15–25 mm long; indehiscent. Replum invisible. Epicarp dull, or glossy; monochrome; light to dark brown or reddish to purplish brown; with surface texture uniform; glabrous; eglandular; without spines; not smooth; with elevated features; not veined; not tuberculate; longitudinally ribbed (with 20–40 ribs); not exfoliating; without cracks; without embedded tissue, much thicker than epicarp, running from base to apex. Mesocarp present; thick; surface not veined; 2-layered; without balsamic vesicles; without fibers; without reniform canals; with solid layer over solid layer; ligneous. Endocarp present; visible; dull; opaque; monochrome; white (when fresh, Du Puy et al. [2002]), or brown (pale, when dry); spongy; without adhering pieces of testa; nonseptate; spongy (when dry disintegrating at the touch); not exfoliating; remaining fused to mesocarp and epicarp; without wings; entire. Seed(s) 1–3(–4); length parallel with fruit length; neither overlapping nor touching; in 1 series. Aril present; dry; when dry rim-aril; entire; covering less than 1/2 of seed; without tongue (or flap) on lips of 2-lipped rim-aril; dark brown.

Seed: 17–20 mm long; 8–12 mm wide; 7–9 mm thick; not overgrown; not angular; symmetrical; elliptic to oblong, or ovate (slightly); compressed; 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; with pieces of adhering epicarp; completely adhering to endocarp; free from endocarp; dull; not modified by a bloom; colored; streaked; with frequent streaks; reddish to dark brown, or red (Du Puy, 2002); with brown overlay; glabrous; not smooth; with recessed features; faintly grooved, or striate; coriaceous. Pleurogram absent. Pseudopleurogram absent. Fracture lines absent. Rim absent. Wing(s) absent. Raphe visible (11–15 mm long); from hilum through lens to base of seed and terminating; not bifurcating; lighter than testa; reddish brown; recessed. Hilum present; visible; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform; 1.5–2 mm long; with curved outline; elliptic; recessed; within halo; halo lighter than testa. Lens not discernible. Endosperm present; thick; not pluglike and not resembling tip of radicle; covering at least 1/2 of embryo, but not entire embryo; adnate to testa. Cotyledons not smooth; 1–3 grooves on each face to 4–6 grooves on each face (longitudinally); outer face of one cotyledon flat and other cotyledon concave (partially folded cotyledon with face flat in center and fully folded cotyledon convex in center due to folding); both the same thickness; both more or less of equal length; with both folded; sufficiently folded for inner face to touch itself and not sufficiently folded for inner face to touch itself (1 cotyledon partially folded and the other fully folded with edges touching in center); portions of inner folded face equal and unequal (fully folded cotyledon with edges touching and equal and other partially folded without edgse touching and equal); margin entire 180 degrees from base of radicle; similar at apex; partially concealing radicle; notched at radicle; with lobes; with lobes touching (auriculate); with basal groin formed by lobes; with the interface division terminating at base of radicle; without margins recessed; white; inner face with central ridge on 1 and central groove on other (fully folded cotyledon with longitudinal central ridge and the partially folded one with longitudinal central groove); glabrous on inner face. Embryonic axis straight; oblique to length of seed; without a joint evident between the radicle and the cotyledons. Radicle differentiated from cotyledon; bulbose; lobe tip straight; straight with embryonic axis; centered between cotyledons; less than 1/2 length of cotyledons. Plumule moderately developed; glabrous.

Distribution

Endemic to Madagascar.

Old World; endemic to Madagascar.

Generic Notes

Capuron (1968) described and illustrated the fruit of C. haraka R. Capuron. Du Puy et al. (2002) treated the Madagascarian Cordyla and observed, "the Malagasy species are distinctive as the inner stamens are sterile." Their descriptions of its fruits and seeds are excellent, and clearly demonstrate that the Madagascarian Cordyla are also different from the remainder of Cordyla because of their fruit and seed structures. Twenty fruit and seed characters, as well as distribution, separate Dupuya from Cordyla: 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, versus 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. 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.

 Fruit and seed:  D. haraka  (H.E. Baillon) J.H. Kirkbride, ined. - second fruit from the left;  D. madagascariensis  (R. Viguier) J.H. Kirkbride, ined. - first and third fruits from the left, the solitary seed, and a magnification of its hilum.
Fruit and seed: D. haraka (H.E. Baillon) J.H. Kirkbride, ined. - second fruit from the left; D. madagascariensis (R. Viguier) J.H. Kirkbride, ined. - first and third fruits from the left, the solitary seed, and a magnification of its hilum.
 Seed, cotyledon, embryo, and testa:  D. madagascariensis  (R. Viguier) J.H. Kirkbride, ined. - embryo, cotyledons, and testas.
Seed, cotyledon, embryo, and testa: D. madagascariensis (R. Viguier) J.H. Kirkbride, ined. - embryo, cotyledons, and testas.