Calophaca

Taxonomy

Calophaca F.E.L. von Fischer ex A.P. de Candolle Prodr. 2: 270. Nov (med.) 1825.

Subfamily: Faboideae.
Phylogenetic Number: 3.16.12.
Tribe: Galegeae.
Subtribe: Astragalinae.
Species Studied - Species in Genus: 3 studied; 5 in genus.

Description

Fruit: A legume; unilocular; 1–3 cm long; 0.3–0.5 cm wide; 0.4–0.5 cm thick; 2–9 times longer than wide; with deciduous androecial sheath; with deciduous corolla; with persistent calyx; with calyx shorter than fruit; without orifice formed by curving of fruit or fruit segments; straight; not plicate; not twisted; asymmetrical; linear; with both sutures nearly straight; inflated (when mature); terete (when mature); with beak; straight; with solid beak the same color and texture as fruit; short tapered at apex; aligned with longitudinal axis of fruit, or oblique with longitudinal axis of fruit; tapered at base; aligned with longitudinal axis of fruit; with the apex and base uniform in texture; coriaceous; seed chambers externally invisible; margin not constricted; margin without sulcus; margin plain, or embellished; margin with fringe (plain and glandular hairs); wing(s) absent; nonstipitate; with all layers dehiscing; splitting along suture(s). Dehiscence of valves along both sutures; apical and down; active; with valves twisting. Replum invisible. Epicarp dull; monochrome; dark reddish brown; with surface texture uniform; pubescent and indurate; with hairs erect; with 1 type of pubescence, or 2 types of pubescence; puberulent, or villous (restricted to sutures or throughout); with pubescence reddish brown, or gray; with long and short gray plain-tipped hairs; with pubescence uniformly distributed; with simple hairs and glandular hairs, or glandular hairs; pliable; with hair bases plain; glandular; with glandular hairs (reddish-brown), or dots (reddish-brown); without spines; not smooth; with elevated features; veined; reticulately veined; not tuberculate; 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 vitriol layer over solid layer; coriaceous. Endocarp present; visible; dull; opaque; monochrome; tan; cobwebby and spongy (but thinly filled); without adhering pieces of testa; subseptate, or nonseptate; with septa thin (tissue paper-like), flexible; with septa eglandular; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; without wings; entire. Seed(s) 1–8; length oblique to fruit length; neither overlapping nor touching; in 1 series. Funiculus 1–2 mm long; of 1 length only; somewhat filiform; straight, or curved. Aril absent.

Seed: 4–8 mm long; 0.4–5 mm wide; 0.1–3.3 mm thick; not overgrown; not angular; asymmetrical; (sub-) reniform, or oblong; compressed; with surface smooth; with visible radicle and cotyledon lobes, or without visible radicle and cotyledon lobes; without external groove between radicle and cotyledon lobes; with deep 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; dull; not modified by a bloom; colored; monochrome; reddish brown; glabrous; smooth, or not smooth; with elevated features; warty, or wrinkled; coriaceous. Pleurogram absent. Pseudopleurogram absent. Fracture lines absent. Rim absent. Wing(s) absent. Raphe not visible. Hilum present; partially concealed, or fully concealed; concealed by funicular remnant, or radicle lobe, or wing; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; larger than punctiform, or punctiform; 0.1–0.7 mm long; with curved outline; elliptic; marginal according to radicle tip, or between cotyledon and radicle lobe; flush; not within corona, halo, or rim. Lens discernible, or not discernible; 1 mm long; with margins straight; linear; not in groove of raphe; confluent with hilum; flush; dissimilar color from testa; darker than testa; black; not within corona, halo, or rim. Endosperm present; thin; not pluglike and not resembling tip of radicle; covering entire embryo; adnate to testa, or embryo. 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; not concealing radicle; entire over radicle; without lobes; with the interface division terminating at base of radicle; without margins recessed; reddish to dark reddish brown; inner face flat; glabrous on inner face. Embryonic axis deflexed; oblique to length of seed; without a joint evident between the radicle and the cotyledons. Radicle differentiated from cotyledon; bulbose; lobe tip curved, or hooked; deflexed and parallel to cotyledon length; centered between cotyledons; less than 1/2 length of cotyledons, or 1/2 to nearly length of cotyledons. Plumule rudimentary; glabrous.

Distribution

Central Asia (4 spp.) and Russia (C. wolgarica).

Old World; Europe to Mediterranean to Russia to China.

Tribal Notes

Tribe Galegeae

Traditionally this tribe has been called Galegeae. Reveal (1997) reported that the name Astragaleae was published before the name Galegeae. Following the International Code of Botanical Nomenclature (Greuter et al., 1994), the oldest name for a taxon must be used, so Reveal suggested that this tribe should be called Astragaleae. In 1999 Reveal (1999) reversed himself, so that this tribe remains the Galegeae. Welsh (1960) reported on the Galegeae of north-central United States. Sanderson and Liston (1995) carried out cladistic analyses of Galegeae genera using molecular data. They concluded that Galegeae is paraphyletic having given rise to tribes Cicereae (20), Hedysareae (18), Trifolieae (21), and Fabeae (19), and therefore requiring a re-evaluation of the circumscription of Galegeae. Heenan (1995, 1998c), utilizing unpublished nuclear ribosomal DNA ITS data, concluded that "Carmichaelia (17.05) is nested within (the) 'Astragalean clade' of Galegeae" and is the sister group of Clianthus. He therefore supported the proposal of Sanderson and Wojciehowski (1996) that Carmichaelieae should not be recognized at tribal level, but rather included in Galegeae.

 Fruit and seed:  C.  spp. - fruits (closed and dehisced) and seeds.
Fruit and seed: C. spp. - fruits (closed and dehisced) and seeds.
 Cotyledon, embryo, and testa:  C. wolgarica  (C. Linnaeus f.) F.E.L. von Fischer ex A.P. de Candolle - embryo, cotyledons, and testa SEMs.
Cotyledon, embryo, and testa: C. wolgarica (C. Linnaeus f.) F.E.L. von Fischer ex A.P. de Candolle - embryo, cotyledons, and testa SEMs.