South African Journal of Botany 72 (2006) 147 – 149 www.elsevier.com/locate/sajb

Studies in in southern 39: articulatus L. and Cyperus corymbosus Rottb.

K.D. Gordon-Gray a,*, C.J. Ward b, T.J. Edwards a

a School of Botany and Zoology, University of KwaZulu-Natal Pietermaritzburg, Private Bag X01, Scottsville 3209, South Africa b P.O. Box 30501, Mayville 4058, South Africa

Received 23 July 2004; accepted 14 June 2005

Abstract

Cyperus articulatus L. and Cyperus corymbosus Rottb. were studied in southern Africa and were found to be synonymous. The latter bears laminae, has longer and frequently has less septate culms than the former but these differences are phenotypic. Phenotypes formerly referred to C. corymbosus are generally rare and scattered, occupying the drier fringing zones of populations. A map of the distributional range of C. articulatus in southern Africa is provided, along with an illustration of the Fcorymbosus_ phenotype. D 2005 SAAB. Published by Elsevier B.V. All rights reserved.

Keywords: Cyperus articulatus; Cyperus corymbosus; Southern Africa

Cyperus articulatus L. and Cyperus corymbosus Rottb., two C. articulatus often occurs in almost pure stands in tropical tropical, perennial, rhizomatous, helophytic hydrophytes had and warm temperate localities that provide permanent water. It been considered specifically distinct until Tucker (1994), after is distinguished by its robust, leafless culms up to 1600Â8mm an extensive survey of Mexican, South American and Old that are septate-noded and solid-pithed. In life the nodes are not World specimens, found he could not substantiate accepted always clearly defined externally but may be detected by gentle character differences. Consequently he treated C. corymbosus finger pressure or longitudinal sectioning. On drying, with as a synonym of C. articulatus. Study in southern Africa has shrinkage of pith, the septa stand out markedly. Bracts of the supported Tucker’s treatment. We propose that the former is a brown, branched, terminal inflorescences are short (5–9 mm), laminate form with longer bracts and frequently less septate- stiff and sharp and never exceed primary ray length. The noded culms. Commonly sympatric, the two entities are subterranean, curved stoloniferous are scale-covered distributed widely; C. corymbosus is rare and scattered within when young and terminate, at intervals, in erect, emergent or adjacent to more extensive stands of C. articulatus. culms. Corm development at the joints is not usually Cyperus articulatus L. Sp. pl. 1: 44 (1753), Kunth: 53 (1837), evident apart from occasional slight enlargements and we Steudel: 30 (1855), Boeck.: 274 (1869/70), C.B.Clarke: 356 found no reference to corm production in the literature. In areas (1901/02), Ku¨kenthal: 77 (1935), Ayers: 1–104 (1946), with seasonal climate, fluctuating water levels and greater R.W.Haines and Lye: 183 (1983), Tucker: 95 (1994). Lectotype: competition, the facies is modified. Culms remain tall, but Sloane, Voy. Jamaica Nat. His. 1: 121, t, 81, Figure 1 (1707). are slender with fewer septa. Occasionally short, inconspicuous Cyperus corymbosus Rottb. Descr. pl. rar. 19 (1772); blades terminate the exposed lower leaf sheaths. Bracts are C.B.Clarke: 554 (1894), C.B.Clarke: 181 (1897/98), longer (20–36 mm), becoming leaf-like, but seldom exceed the C.B.Clarke: 357 (1901/02), Ku¨kenthal.: 80 (1935), R.W.Haines inflorescence (Fig. 1). and Lye: 183 (1983). Type: Suriname, Rolander s.n. (holo., C. corymbosus exhibits a close morphological relationship Herb. Rottb. C.). with this less robust expression of C. articulatus. Accepted differences between the two species have been based upon, for * Corresponding author. C. corymbosus, culms 1- or 2-bladed, bracts leaf-like, culm E-mail address: [email protected] (K.D. Gordon-Gray). septa sparse or absent (Clarke, 1901/02; Ku¨kenthal, 1935;

0254-6299/$ - see front matter D 2005 SAAB. Published by Elsevier B.V. All rights reserved. doi:10.1016/j.sajb.2005.06.004 148 K.D. Gordon-Gray et al. / South African Journal of Botany 72 (2006) 147–149

extracted from earlier literature without determining any significant anatomical divergence. However, this is a limited assessment requiring confirmation. Synonymy is further supported by distribution. C. articu- latus, except for Malaysia, is common pantropically; C. corymbosus is reported from numerous localities within, or fringing the range of C. articulatus but is rare. The only exception to this pattern known to us is in Sri Lanka (Ceylon) where Koyama (1985) reported the apparent replacement of C. articulatus by C. corymbosus over a century. He commented after describing C. articulatus—FIt is rather interesting that this species has never been collected since Trimen’s time_ (c. 1888). Such a change could be the result of gradual modification of habitat, perhaps in conjunction with herbivory or other consistent mutilation. In southern Africa C. articulatus is well-known and widely represented in fresh, or slightly saline, mostly moving water of estuaries, river banks, and wet grassland from sea level to about 1065m altitude (Fig. 2). It is reported from northern Namibia including Caprivi, Botswana, Swaziland and from the Limpopo, Mpumalanga and KwaZulu- Natal (Archer, 2003; Retief and Herman, 1997) and the Eastern Cape provinces of South Africa. C. corymbosus is likewise widespread extra territorially, but poorly represented in herbaria and referred to as Frare_ or Funcommon_. Clarke (1897/98) in his treatment of Cyperus for southern Africa, Fig. 1. Cyperus articulatus (corymbosus form): whole plant showing one leaf omitted C. articulatus, but did include C. corymbosus, citing blade drying off. Culm articulations are not well defined (Ward 14828). only Dre`ge 4406 from FNatal, near the mouth of the Umzimkulu River_ [now Mzimkhulu (Port Shepstone)] and Ayers, 1946). Tucker (1994) was unable to substantiate these adding FKunth, Enum. ii: 53, partly, i.e. the Dre`ge examples character differences in a study of Fsome 400 Old World and with septations of stems obsolete_: [Note: a fragment of the South American collections labeled as one or other of the inflorescence of Dre`ge 4406 is preserved in PRE]. The species species_. He therefore treated C. corymbosus and C. articulatus has been recollected from Dre`ge’s locality, nearly two centuries as synonymous. later (CJ Ward 14828). Metcalfe (1971) examined culms and rhizomes of C. Ward (1980) recorded both species from Isipingo, noting the articulatus from Tanzania and cited features of C. corymbosus elongated rhizomes, tuberous [sic] only in C. corymbosus. The

Fig. 2. Known distribution of Cyperus articulatus in southern Africa. K.D. Gordon-Gray et al. / South African Journal of Botany 72 (2006) 147–149 149 presence of well defined underground storage organs in this – 3030 (Port Shepstone): Mzimkhulu River mouth (-CB), species (initially termed tubers, but now determined as corms, Dre`ge 4406** (K, PRE), CJ Ward 14828** (NH, NU, PRE, since they are stems which become hard and woody and are UDW). tunicated by frayed leaf sheaths) was a significant distinction – 3128 (Umtata): Elliotdale, The Haven (-BB), JL Gordon- from C. articulatus. The presence of corms in an allied species Gray 1007** (NU); Kei River Estuary (-CB), CJ Ward prompted the suggestion (Gordon-Gray, 1995) that C. cor- 14750 (GRA, NU, UDW). ymbosus might be a hybrid in South Africa. Subsequent study has dispelled this hypothesis. Our experience is that corm production is induced in Acknowledgements habitats influenced by seasonal change, fluctuating water levels and increased competition, and is almost always We express thanks for help received from the South accompanied by sparse leaf blades and short leaf-like bracts. African Biodiversity Institute, Pretoria, and the Herbarium, Such occur most commonly during colonization and in School of Botany and Zoology, University of KwaZulu-Natal, areas fringing favourable habitats. Further, we now have Pietermaritzburg. evidence of corms in what, on characters of emergent culms, must be named C. articulatus—from both wild collections References and cultivated plants (CJ Ward 15045; 15111; 15169). Clearly in southern Africa, as in other areas of distribution, C. Archer, C., 2003. Cyperaceae. In: Germishuizen, G., Meyer, N.L. (Eds.), Plants corymbosus is synonymous with C. articulatus (representing of Southern Africa: An Annotated Check List, Strelitzia, vol. 14. National Botanical Institute, Pretoria. an environmental or juvenile expression of the latter). We Ayers, B. Bro, 1946. The genus Cyperus in Mexico. Catholic Universities of point to obvious difficulties that underlie the lack of corms on America, Biological Studies 1, 1–104. specimens of C. articulatus. These include the tropical watery Boeckeler, O., 1869/70. Die Cyperaceae des Ko¨ninglichen Herbariums zu habitat and the early decay of interconnecting stolons that Berlin. Linnaea 36, 271–768. leave the longer lasting culms isolated. Who has investigated Clarke, C.B., 1894. Cyperaceae. In: Durand, T., Schinz, H.Conspectus Florae Africae ou E´ nume´ration des Plantes d’Afrique, vol. 5. Friedlaender, Berlin adequately what exists in substrates where unfriendly biota and Klincksieck, Paris, pp. 526–692. (from bilharzia to crocodiles) lurk? Most herbarium speci- Clarke, C.B., 1897/98. Cyperaceae. Flora Capensis, vol. 7. Reeve and Co, mens comprise a growing stolon severed at a point behind London, pp. 149–310. one or two flowering culms—young portions where corm Clarke, C.B., 1901/02. Cyperaceae. Flora of Tropical Africa, vol. 8. Reeve and enlargement has not yet commenced. Co, London. Gordon-Gray, K.D., 1995. Cyperaceae in Natal: Strelitzia 2. National Botanical Key to superscripts *articulatus (cormose) form; **corym- Institute, Pretoria. bosa form Haines, R.W., Lye, K.A., 1983. The Sedges and Rushes of East Africa. East Selected specimens African Natural History Society, Nairobi. Koyama, T.M., 1985. Cyperaceae. In: Dassanyaka, M.D., Fosberg, F.R. (Eds.), A R – 1113 Angola (Cuanza Sul Prov.): Novo Redondo, Negunza revised Handbook to the Flora of Ceylon, vol. 5, pp. 125–405 Rotterdam. Ku¨kenthal, G. (1935/36) Cyperaceae.– Scirpoideae – Cypereae. Das Pflan- River, CJ and JD Ward 76 (NU, UDW). zenreich 4, 20 heft 101: 163–671. Leipzig. – 1713 Namibia (Swartbooisdrif): Cunene River, Epupa Falls Kunth, C.S., 1837. Cyperographica synoptica. Enumeration Plantarum 2. J (-AA), CJ Ward 12047 (NU, UDW). Cotta, Stuttgart and Tu¨bingen. –1934 Mozambique (Beira): Gorongosa National Park, Metcalfe, C.R., 1971. Anatomy of the : V. Cyperaceae. Pungwe River, CJ Ward 7797 (NU, UDW). Clarendon Press, Oxford. Retief, E., Herman, P.P.J., 1997. Cyperaceae. Plants of the Northern Provinces – 2829 (Harrismith): Weenen County, Brakfontein, near Frere of South Africa: Keys and Diagnostic Characters, Strelitzia, vol. 6. National (-DD), Acocks 10827 (PRE). Botanical Institute, Pretoria. – 2831 (Nkandla): Mtunzini, Sihayi Lagoon (-DD), CJ and Steudel, E.G. Von, 1855. Synopsis Plantarum Glumacearum, Pars 2 Cyper- MC Ward 57 (NU, UDW); upper Mfule (-CB), CJ Ward aceae. Stuttgart. 15044, 15045* (NH, NU, UDW, PRE). Tucker, G.C., 1994. Revision of the Mexican Species of Cyperus (Cyperaceae). Systematic Botany Monographs, vol. 43. American Society of Plant – 2832 (Mtubatuba): Hluhluwe Game Reserve, Hluhluwe Taxonomists, Ann Arbor, Michigan. River (-AA), CJ Ward 15111* (NH, NU, PRE, UDW); Ward, C.J., 1980. The Plant Ecology of the Isipingo Beach Area, Natal, South Richards Bay, Thulazihleka (-CC), CJ Ward 15169* (NH, Africa. Memoirs of the Botanical Survey of South Africa, vol. 45. NU, PRE, UDW). – 2930 (Pietermaritzburg): Merrivale Shooting Range, sedge marsh (-CA), Kotze 86, 147, 148** (NU); Isipingo Beach (-DD), CJ Ward 390, 664** (NU).