Caragana

Taxonomy

Caragana P.C. Fabricius Enum. ed. 2. 421. Sep-Dec 1763.

Subfamily: Faboideae.
Phylogenetic Number: 3.16.11.
Tribe: Galegeae.
Subtribe: Astragalinae.
Species Studied - Species in Genus: 21 studied; ca. 80 in genus.

Description

Fruit: A legume; unilocular; 2–6 cm long; 0.25–0.6 cm wide; 0.15–0.4 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; not inflated, or inflated; compressed; without beak, or with beak; declined; with solid beak the same color and texture as fruit; short tapered at apex; oblique with longitudinal axis of fruit; short tapered at base; oblique 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; 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; glabrous; glandular, or eglandular; with glandular dots; without spines; not smooth; with elevated features; veined; reticulately veined; not tuberculate; somewhat wrinkled; not exfoliating; without cracks; without embedded tissue, much thicker than epicarp, running from base to apex. Mesocarp present; thin; surface not veined; 1-layered; without balsamic vesicles; without fibers; without reniform canals; solid; coriaceous. Endocarp present; visible; dull; opaque; monochrome; reddish tan; smooth; without adhering pieces of testa; nonseptate; chartaceous; not exfoliating; remaining fused to mesocarp and epicarp; without wings; entire. Seed(s) 2–5; length parallel with fruit length; neither overlapping nor touching; in 1 series. Funiculus 0.5–1 mm long; of 1 length only; thick; straight. Aril present; dry; when dry rim-aril; entire; white.

Seed: 2.5–10 mm long; 2–4.5 mm wide; 2–4.5 mm thick; not overgrown; not angular, or angular (barely); asymmetrical; oblong, or linear, or circular, or reniform; terete, or compressed; with surface smooth; with visible radicle and cotyledon lobes; with external groove between 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; dull, or glossy; not modified by a bloom; colored; monochrome, or mottled and streaked; with frequent mottles; with frequent streaks; dark reddish to organgish brown, or orange; with brown overlay (dark reddish); glabrous; smooth, or not smooth; with recessed features; pitted with small separate pits; chartaceous, or coriaceous. Pleurogram absent. Pseudopleurogram absent. Fracture lines absent. Rim absent. Wing(s) absent. Raphe visible, or not visible; from hilum to lens (at base of seed), or lens to base of seed and terminating; not bifurcating; color of testa (barely darker); flush. Hilum present; partially concealed; concealed by funicular remnant; with faboid split; with the lips of the faboid split the same color as the rest of the hilum; punctiform, or larger than punctiform; 0.3–0.5 mm long; with curved outline; circular; marginal according to radicle tip, or between cotyledon and radicle lobe; flush; not within corona, halo, or rim. Lens discernible, or not discernible; 0.2–1 mm long; with margins straight, or curved; wedge-shaped; circular, or elliptic; not in groove of raphe; adjacent to hilum, or confluent with hilum; 0.1–3 mm from hilum; mounded, or recessed; similar color as testa; barely darker than testa; reddish brown; not within corona, halo, or rim. Endosperm present; thin; not pluglike and not resembling tip of radicle; covering entire embryo; adnate to testa. Cotyledons smooth; both outer faces convex; both the same thickness; both more or less of equal length, or 1 longer than other; not folded (though occasionally 1 cotyledon larger than other); 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 tan; 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; linear; lobe tip straight; deflexed and parallel to cotyledon length; centered between cotyledons; less than 1/2 length of cotyledons. Plumule rudimentary, or moderately developed; glabrous.

Distribution

Eastern Europe, Asia, and naturalized in North America.

Old World; eastern Europe to Mediterranean to Russia to Southwest Asia to India to China to Korea to Mongolia.

Generic Notes

Yakovlev and Sviazeva (1984, 1985a, 1987) have treated some of the Russian species, and Gorbunova (1984) established a sectional and subsectional sequence for the genus.

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. - valves, dehisced fruits, and seeds.
Fruit and seed: C. spp. - valves, dehisced fruits, and seeds.
 Cotyledon, embryo, and testa:  C. densa  V.L. Komarov - embryo, cotyledons, and testa SEMs.
Cotyledon, embryo, and testa: C. densa V.L. Komarov - embryo, cotyledons, and testa SEMs.