Copaifera

Taxonomy

Copaifera C. Linnaeus Nom. cons. Sp. Pl. ed. 2. 557. Sep 1762.

Subfamily: Caesalpinioideae.
Phylogenetic Number: 1.4.44.
Tribe: Detarieae.
Group: Detarium.
Species Studied - Species in Genus: 22 studied; 25–30 in genus.

Description

Fruit: A legume; unilocular; 1.5–7 cm long; 1.2–4.5 cm wide; 0.8–2.5 cm thick; length less than twice as long as width; 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, or asymmetrical; nearly circular; with 1 straight and 1 curved suture; widest at base; not inflated; compressed to terete; without beak; tapered at apex; aligned with longitudinal axis of fruit to oblique with longitudinal axis of fruit; rounded at base; right angled with longitudinal axis of fruit; with the apex and base uniform in texture; ligneous; seed chambers externally visible; margin not constricted; margin without sulcus; margin plain; wing(s) absent; substipitate; with all layers dehiscing; splitting along suture(s). Dehiscence of valves along 1 suture, or both sutures (ventrally, or an initial separation along ventral suture followed by complete dehiscence along dorsal suture); passive (valves falling separatley). Replum invisible. Epicarp dull to glossy; monochrome; reddish brown to red; with surface texture uniform; glabrous; eglandular; without spines; not smooth; with elevated features; veined, or not veined; transversely veined relative to fruit length; not tuberculate; rugose, or glandular dotted ("resinous globules"); not exfoliating; without cracks; without embedded tissue, much thicker than epicarp, running from base to apex. Mesocarp present; surface not veined; 1-layered; without balsamic vesicles; with fibers, or without fibers; without reniform canals; fleshy when fresh, fibrous throughout and solid (or entirely solid); ligneous (when dry). Endocarp present; visible; dull; opaque; monochrome; reddish brown to white; without adhering pieces of testa; nonseptate; not exfoliating; remaining fused to mesocarp and epicarp; entire. Seed(s) 1(–2); length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus 0.1–5 mm long; of 1 length only; filiform to thick; straight. Aril often present, or absent (orange yellow to red or blackish red); fleshy; when fleshy arched shape to caplike; entire; covering thick to thin and usually 1/2 to nearly all of seed.

Seed: 13–28 mm long; 9–17 mm wide; 1–32 mm thick; not overgrown; not angular; symmetrical, or asymmetrical; oblong, or irregular; terete to 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; without pieces of adhering epicarp; not adhering to endocarp; free from endocarp; glossy (or nearly so and usually partially concealed by thin aril); not modified by a bloom; colored; monochrome; black to brown; glabrous; smooth, or not smooth; with elevated features; rugose; osseous. Pleurogram absent. Pseudopleurogram absent. Fracture lines present; pattern type not stated. Rim absent. Wing(s) absent. Raphe visible (inconspicuous from hilum through lens and across apex to base of seed). Hilum present; fully concealed; concealed by funicular remnant; without faboid split; larger than punctiform; 0.1–2 mm long; with curved outline, or angular outline; elliptic; triangular; nearly marginal according to radicle tip; flush; not within corona, halo, or rim. Lens discernible, or not discernible; 0.1–0.3 mm long; with margins curved; circular; not in groove of raphe; mounded; dissimilar color from testa; red; 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; similar at apex; completely concealing radicle; split over radicle; with lobes; with lobes overlapping, or touching (auriculate); with basal groin formed by lobes, or 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; oblique to length of seed; without a joint evident between the radicle and the cotyledons. Radicle differentiated from cotyledon; centered between cotyledons. Plumule rudimentary; glabrous.

Distribution

Tropical America and Africa and African southern deciduous woodlands.

New World and Old World; West Indies, Central America, and South America (tropical); Argentina, Peru, Brazil, Ecuador, and the Guianas; Africa and Indonesia and the Philippines (including African deciduous woodlands).
 

Generic Notes

If fruits are damaged, Resin globules from the mesocarp may be present on the surface of the epicarp. A coumarin odor was detected during drilling seeds of C. officinalis. Several species are the source of copaiba, an oleoresin compound with pharmaceutical uses. J. Langenheim and Silva are studying the genus. Cowan and Polhill (1981a) noted that this genus may include Pseudosindora (4.44). Hou (1994, Blumea 38:313–330) synonmized Pseudosindora under Copaifera.

Tribal Notes

Tribe Detarieae

Bruneau et al. (2000) carried out extensive phylogenetic analyses of tribes Amherstieae and Detarieae. They concluded that they form a single monophyletic group. Therefore, they supported Polhill's (1995a, 1995b) decision to unite the two tribes.

 Fruit:  C. baumiana  H.A.T. Harms - bottom left seed topography;  C. glycycarpa  W.A. Ducke - top left fruit;  C. langsdorfii  R.L. Desfontaines - center seed topography;  C. luetzelburgii  H.A.T. Harms - bottom center fruit, top center seed in situ;  C. mildbraedii  H.A.T. Harms - right seed in situ.
Fruit: C. baumiana H.A.T. Harms - bottom left seed topography; C. glycycarpa W.A. Ducke - top left fruit; C. langsdorfii R.L. Desfontaines - center seed topography; C. luetzelburgii H.A.T. Harms - bottom center fruit, top center seed in situ; C. mildbraedii H.A.T. Harms - right seed in situ.
 Seed, cotyledon, embryo, and testa:  C. officinalis  C.L. von Willdenow - far left cotyledon split and concealing radicle (above) and embryonic axis (below), top left center seed topography, testa SEMs;  C.  spp. - bottom left center seeds.
Seed, cotyledon, embryo, and testa: C. officinalis C.L. von Willdenow - far left cotyledon split and concealing radicle (above) and embryonic axis (below), top left center seed topography, testa SEMs; C. spp. - bottom left center seeds.