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South African Journal of Botany 81 (2012) 124–127 www.elsevier.com/locate/sajb

Short communication A review of section Cyanospermum (, ) in South Africa ⁎ A.N. Moteetee a, , J.S. Boatwright b, T.P. Jaca a

a University of Johannesburg, Department of Botany and Biotechnology, P. O. Box 524, Auckland Park 2006, Johannesburg, South Africa b Department of Biodiversity and Conservation Biology, University of the Western Cape, Private Bag X17, Bellville 7535, Cape Town, South Africa

Received 10 May 2012; received in revised form 15 June 2012; accepted 22 June 2012 Available online 20 July 2012

Abstract

We review Rhynchosia section Cyanospermum in South Africa. It is characterised by a robust habit, racemose inflorescence of numerous flowers with a large, tomentose calyx with broad lobes, and dark blue seeds. The section is represented in the Flora of southern Africa Region by two species, R. hirta and R. resinosa. However, the latter species lacks the diagnostic dark blue seeds of the section and has not been included here. Rhynchosia hirta appears to be allied to the Madagascan R. leandrii from which it can be distinguished by its larger flowers and seeds. © 2012 SAAB. Published by Elsevier B.V. All rights reserved.

Keywords: Fabaceae; Phaseoleae; Rhynchosia; South Africa;

1. Introduction allied to Eriosema (DC.) G.Don. within the Cajaninae (Doyle and Doyle, 1993). These two genera, together with Paracalyx Ali and The genus Rhynchosia Lour. comprises about 230 species Flemingia Ait.f., differ from the rest of the subtribe in having fewer distributed mainly in Africa and Madagascar but extending to than 3 ovules and inconspicuous arils on the seeds (Lackey, 1981). warm temperate and tropical Asia, northern Australia and tropical Rhynchosia and Eriosema share pinnately compound leaves and and subtropical America (Schrire, 2005). It derives its name from axillary racemes but can be distinguished from each other by the a Greek word “rhynch” meaning “beak” in reference to the shape point of attachment of the funicle to the hilum in the seed: attached of the keel in R. volubilis Lour. (the type species). According to in the middle in Rhynchosia and at the end of the hilum in Harvey (1862),however,“the carina is scarcely rostrate” in Eriosema (Lackey, 1981). Rhynchosia. The current sectional classification of Rhynchosia (Baker, Rhynchosia is a member of subtribe Cajaninae Benth. of 1923) divides the South African species into five sections: tribe Phaseoleae (Fabaceae). Cajaninae is monophyletic (Kajita Cyanospermum (Wight and Arnott) Benth. (one spp.), et al., 2001) and diagnosed by the presence of vesicular glands Arcyphyllum Torrey and Gray (two spp.), Chrysoscias E.Mey. (on stems, calyces, ovaries, fruits and on the lower surface of (four spp.), Polytropia Presl. (two spp.), and Eurhynchosia Baker f. the leaflets), and bulbous-based hairs (which cover the whole (59 ssp.), which also includes Copisma E.Mey. and Orthodanum plant), and also by the lack of bracteoles, with the exception of E.Mey. The aptly named section Cyanospermum is characterised Adenodolichos Harms (Lackey, 1977; Moteetee and Van Wyk, by its dark blue seeds, as well as robust stems, racemose 2006).Basedonmoleculardata,Rhynchosia appears to be closely inflorescences with numerous flowers, and tomentose calyces with broad lobes. In his treatment of Rhynchosia, Baker ⁎ Corresponding author. Tel.: +27 11 5592977; fax: +27 11 5592411. (1871) placed five species in this section, all distributed in E-mail address: [email protected] (A.N. Moteetee). tropical Africa. Two of these, R. hirta (Andr.) Meikle & Verdc

0254-6299/$ -see front matter © 2012 SAAB. Published by Elsevier B.V. All rights reserved. doi:10.1016/j.sajb.2012.06.007 A.N. Moteetee et al. / South African Journal of Botany 81 (2012) 124–127 125 and R. resinosa (Hochst. ex A. Rich.) Baker also extent to north- Inflorescences 7–many-flowered axillary racemes, occasionally eastern and northern parts of South Africa respectively. The latter branched, 10–40 cm long, reddish brown pubescent. Flowers 15– species was curiously not included in Baker's(1923)revision of 28 mm long; bracts broadly elliptic or broadly ovate, (10)12– South African species of Rhynchosia. The species lacks the 18 × 6–10 mm, reddish brown pubescent, caducous; bracteoles diagnostic dark blue seeds of the section, having seeds that absent. Calyx bilabiate, entirely velvety canescent or sometimes are black or “dark reddish-brown mottled with black” (Gillett brownish pubescent at base; lobes of upper lip connate almost to et al., 1971) and has not been included here in anticipation of apex, those of the lower lip unequal, oblong-elliptic, carinal lobe molecular studies of the genus. Another species from Gabon, longer than lateral lobes, 14–28 mm long, lateral lobes 8–20 mm R. gabonensis Jongkind (Jongkind, 1994) also belongs to this long. Corolla persistent, mostly shorter than calyx, varying from section. The nomenclature of R. hirta was clarified by Verdcourt greenish-cream or yellow striped with red, through orange or pink and Meikle (1967). We provide a taxonomic treatment of the with red or purple venation to brick red; standard suborbicular, taxon in South Africa, including a description, illustrations and a 15–22 × 12–18 mm, slightly emarginate, glabrous, with two distribution map. Although this species was treated for Flora of callosities at base, claw 2–4 mm long; wings narrower and Tropical East Africa (Gillett et al., 1971), the treatment did not shorter than keel, 10–12×3–4 mm, spurred at base, without include correct typification which is provided here. sculpturing, claw 3–4 mm long; keel larger than wings, 12–14 × 5–6 mm, pocketed, claw 5–6 mm long. Stamens 2. Taxonomic treatment diadelphous (nine filaments fused, vexillary stamen free to the base), anthers monomorphic, dorsifixed. Ovary narrowly Rhynchosia hirta (Andr.) Meikle & Verdc. in Taxon 16: 462 oblong, subsessile, pubescent, 2-ovuled; style curved up- (1967); Verdc. in Fl. Trop. E. Afr.: 720 (1971); Drummond in wards, glabrous. Fruit narrowly oblong, 15–20×5–6 mm, Kirkia 8: 226 (1972); Gonçalves in Garcia de Orta, Sér. Bot. 5: densely velvety canescent or rusty-brown pubescent, plicate, 106 (1982); Lock, Leg. Afr. Check-list: 430 (1989). Dolichos 2-seeded. Seeds depressed-globular, 5–7×3–4 mm, smooth hirtus Andr., Bot. Repos. 7: t. 446 (1807). Type: grown in England and glossy, dark blue when fresh, becoming black when dry, from seed said to have been sent from the Cape of Good Hope, remainingattachedtopod(Fig. 1). (illustrationinAndr.,Bot.Repos.7:t.446.(1807),lectotype,here designated). 2.1. Flowering time: August–March Cylista villosa Ait., Hort. Kew., ed. 1, 3 : 36 (1789); Ait. f., Hort. Kew., ed 2, 4: 300 (1812), non Rhynchosia villosa Druce Distribution and habitat: R. hirta is widespread from northeast (1917). South Africa northwards into East and Central Africa and Rhynchosia cyanosperma Benth. ex Baker in Fl.Trop. Afr. westwards into northern Nigeria), and further east to the 2: 218 (1871). E.G. Baker in J. Linn. Soc., Bot. 40: 58 (1911); Mascarene Islands, India and Sri Lanka (Verdcourt, 2001). In Eyles in Trans. Roy. Soc. South Africa 5: 382 (1916); Baker f., South Africa it occurs in Limpopo and Mpumalanga Provinces Legum. Trop. Africa: 469 (1929). Type: Malawi, ‘Zambesi land, (Fig. 2). It grows in sandy soil in grasslands, rocky outcrops, open Manganja Hills’, Meller s.n. (K, lectotype, designated by Gillett woodlands and forest margins. et al., 1971). Diagnostic characters: R. hirta has a similar growth form to the Rhynchosia tomentosa (Roxb.) Baill. in Bull. Mens. Soc. Madagascan species R. leandrii Du Puy and Labat. When Linn. Paris 1: 387 (1883), non Hook. & Arn. (1835) nec Kurz. describing the latter species, Du Puy and Labat (1996) did (1874) Cyanospermum tomentosum (Roxb.) Wight & Arn., not specify its sectional placement, but it conforms to sect. Prodr. Fl. Ind. Orient.: 260 (1834). Cylista tomentosa Roxb., Pl. Cyanospermum in its robustness, reddish brown pubescence, Coromandel 3: 16, t. 221 (1811). Type: grown in Calcutta from and dark blue seeds and may belong here. The flowers of R. hirta seed sent from Mysore, (illustration in Pl. Coromandel 3: t. 221 aremuchlargerthanthoseofR. leandrii (e.g. standard petal (1811), lectotype, here designated). 22×18 mm vs 6×6 mm), as well as the seeds (up to 7 mm long Rhynchosia albiflora (Sims) Alston in Trimen, Handb. Fl. compared to up to 4 mm) (Du Puy et al., 2002). Ceylon 6: 85 (1931); Robyns, Fl. Sperm. Parc Nat. Alb. 1: 348 (1948); Brenan, Check-list For. Trees Shrubs Tang. Terr.: 439 3. Additional specimens examined (1949); Hauman in Fl. Congo Belge 6: 190 (1954); Meikle in Fl. W. Trop. Afr., ed. 2, 1: 554 (1958). Cylista albiflora Sims in South Africa. LIMPOPO: 2329 (Polokwane): Louis Trichardt Curtis' Bot. Mag. 43: t. 1859 (1815). Type: grown in England (−BB), E. Koker 28 (PRE). 2330 (Tzaneen): Soutpansberg from seeds received from Mauritius, illustration in Curtis' Bot. District, Elim Hospital, Vutshila Crafts (− AA), C. Liengme Mag. 43: t. 1859 (1815). 1455 (PRE); Tshakoma, (− AB), A.A. Obermeyer 30341 Vigorous twining or climbing herb, 2–8mlong.Stems densely (PRE); Westfalia Landgoed (− CA), N. Grobbelaar 410 puberulous with short, brownish hairs when young, glabrescent. (PRE); Fanie Bothadam Nature Reserve (−CC), M.F. Borcherds Leaves trifoliolate, leaflets broadly elliptic to ovate or almost (PRE); Letaba (− CD), A.T. Renny s.n. PRE 5613 (PRE). cordate, 55–160× 40–110 mm, sparsely pubescent to glabrescent MPUMALANGA: 2429 (Zebediela): Blouberg district, above, densely pubescent with reddish brown hairs beneath; Potgietersrust (− AA); D. Grosskopf 9008 (NBG). 2430 stipules elliptic-lanceolate, (7) 10–20 × 3–4 mm; stipels linear- (Pilgrim'sRest):Lekgalameetse Nature Reserve, behind lanceolate, 5–7×0.5–1.0 mm; petiole 35–100 (140) mm long. Balloon house no. 10 (− AB), M. Stalmans 1372 (PRE); 126 A.N. Moteetee et al. / South African Journal of Botany 81 (2012) 124–127

Fig. 1. Vegetative and reproductive morphology of Rhynchosia hirta: (A) flowering branch; (B) calyx opened with upper lobes to left; (C) standard petal; (D) wing petal; (E) keel petal; (F) stamens; (G) pistil; (H) lateral view of pod; (I) lateral view of seed attached to pod. Vouchers: A from G.B. Deall 42 (PRE); B–IfromA.T. Renny s.n. PRE 5613 (PRE). Scale bars: A, 30 mm; B, F, 20 mm; A, C–I, 1 mm.

Fig. 2. Geographical distribution of R. hirta in South Africa. A.N. Moteetee et al. / South African Journal of Botany 81 (2012) 124–127 127

Blyderiviers Poort Nature Reserve, Belvedere (−AD), J. Hankey, Papilionoideae–Phaseoleae) from Madagascar and the Comoro Islands. S.L. Turner, A.E. Aubrey 1033 (PRE). 2431 (Acornhoek): 2 miles Bulletin du Muséum National d'Histoire Naturelle. Section B, Adansonia: − Botanique Phytochemie Sér. 4 (18), 85–94. [3.2 km] N of Bushbuck Ridge ( CC), L.E. Codd 1662 (PRE). Du Puy, D.J., Labat, J.-N., Rabevohitra, R., Villiers, J.F., Bosser, J., Moat, J. 2530 (Lydenburg):Sabie(−BB), G.B. Deall 42 (PRE); (Eds.), 2002. The Leguminosae of Madagascar. Royal Botanic Gardens, Kew. Mataffinkop District (−BD), J. Onderstall 1064 (PRE). 2531 Gillett, J.B., Polhill, R.M., Verdcourt, B., 1971. Flora of Tropical East Africa, (Komatipoort): White River Municipal Town lands (− AC), Leguminosae: Subfamily Papilionoideae (Part 4). Crown Agents. J. Onderstall 1355 (PRE); Barberton (− CC), W.E. Holt 292 Harvey, W.H., 1862. Leguminosae. In: Harvey, W.H., Sonder, O.W. (Eds.), Flora Capensis, vol.2, pp. 242–247. (PRE). Jongkind, C.C.H., 1994. Novivates gabonenses (18): a new species of Rhynchosia (Leguminosae: Papilionoideae). Bulletin du Jardin botanique national de – Acknowledgements Belgique 63, 219 221. Kajita, T., Ohashi, H., Tateishi, Y., Bailey, C.D., Doyle, J.J., 2001. rbcL and legume phylogeny, with particular reference to Phaseoleae, Millettieae and allies. The curator of PRE is gratefully acknowledged for assistance Systematic Botany 26, 515–536. with herbarium material. The University of Johannesburg and the Lackey, J.A., 1977. A revised classification of the tribe Phaseoleae (Leguminosae: National Research Foundation Thuthuka programme are thanked Papilionoideae), and its relation to canavanine distribution. Botanical Journal of – for financial support. the Linnean Society 74, 163 178. Lackey, J.A., 1981. Tribe 10. Phaseoleae. In: Polhill, R.M., Raven, P.H. (Eds.), Advances in Legume Systematics, 1. Royal Botanic Gardens, Kew, References pp. 301–327. Moteetee, A.N., Van Wyk, B.-E., 2006. A revision of the genus Bolusafra (tribe Baker, E.G., 1923. Revision of South African species of Rhynchosia. Bothalia Phaseoleae, Fabaceae). South African Journal of Botany 72, 604–608. 1, 113–138. Schrire, B.D., 2005. Phaseoleae. In: Lewis, G., Schrire, B.D., Mackinder, B., Lock, Baker, J.G., 1871. Leguminosae. In: Flora of Tropical East Africa, 218. M. (Eds.), Legumes of the World. Royal Botanic Gardens, Kew, pp. 393–430. Doyle, J.J., Doyle, J.L., 1993. Chloroplast DNA phylogeny of the Papilionoid Verdcourt, B., 2001. Rhynchosia. In: Pope, G.V., Polhill, R.M. (Eds.), Flora legume tribe Phaseoleae. Systematic Botany 18, 309–327. Zambesiaca, vol. 3 (5). Royal Botanic Gardens, Kew, pp. 168–210. Du Puy, D.J., Labat, J.-N., 1996. New taxa and nomenclatural changes in Verdcourt, B., Meikle, R.D., 1967. The identity of Dolichos hirtus Andrews. Rhynchosia Lour. and Eriosema (DC.) Reichb. (Leguminosae– Taxon 16, 462–463.

Edited by JC Manning