Phytotaxa 166 (1): 033–048 www.mapress.com / phytotaxa / Copyright © 2014 Magnolia Press ISSN 1179-3155 (print edition) Article PHYTOTAXA ISSN 1179-3163 (online edition) http://dx.doi.org/10.11646/phytotaxa.166.1.2 Taxonomic changes in C4 Cyperus (Cypereae, Cyperoideae, Cyperaceae): combining the sedge genera Ascolepis, Kyllinga and Pycreus into Cyperus s.l. ISABEL L ARRIDON 1 , K ENNETH BAUTERS 1 , MARC REYNDERS 1 , 2 , W I M HUYGH 1 & PAUL GOETGHEBEUR1 1 Ghent University, Department of Biology, Research Group Spermatophytes, K.L. Ledeganckstraat 35, BE-9000 Gent, Belgium; isabel.larridon@ugent.be, kenneth.bauters@ugent.be 2 Botanic Garden Meise, Nieuwelaan 38, BE-1860 Meise, Belgium. Abstract The sedge genera Alinula, Ascolepis, Kyllinga, Lipocarpha, Pycreus, Queenslandiella, Remirea, Sphaerocyperus and Volkiella (Cyperaceae) were recognised at generic level because they possess specialised inflorescence and/or flower characters. However, recent molecular phylogenetic analyses show that these genera are all nested in a paraphyletic Cyperus s.s. and therefore should be viewed as part of a broadly circumscribed genus Cyperus. For all species of Alinula and for the single species of Queenslandiella, Remirea and Sphaerocyperus, Cyperus names were already published by other authors. For the species of Lipocarpha and Volkiella, Cyperus names and a new sectional classification are published in a separate paper including a detailed molecular phylogenetic hypothesis for these taxa. Based on a study of herbarium specimens and literature, in this paper, twenty species of Ascolepis, seventeen species of Kyllinga, and six species of Pycreus, which do not yet have a validly published and legitimate name in Cyperus, are formally included into Cyperus as new combinations or new names. Notes on the synonymy of an African Pycreus species are also included. Introduction The tribe Cypereae is the second most species rich tribe of the Cyperaceae family. Its largest genus Cyperus (Linnaeus 1753: 44) forms the most diverse sedge genus in the tropics. It also represents one of the most ecologically important angiosperm genera in tropical wetlands. As a result of recent molecular phylogenetic studies (e.g. Muasya et al. 2002, 2009a, Simpson et al. 2007, Larridon et al. 2011a, 2013, Bauters et al. 2014), the need became clear to create a new classification of this tribe reflecting its natural relationships, and to re-evaluate the morphological characters used for generic delimitation in this tribe (Muasya et al. 2009b; Vrijdaghs et al. 2011, Reynders et al. 2012, Larridon et al. 2011b, 2013, Bauters et al. 2014). Following recent molecular phylogenetic analyses (e.g. Simpson et al. 2007, Muasya et al. 2009a, Larridon et al. 2011a, 2013), Cyperus s.l. or the Cyperus clade is a monophyletic clade including a paraphyletic genus Cyperus s.s. encompassing at least 12 segregate genera (following the classification of Goetghebeur 1998 and Govaerts et al. 2007). Besides the paraphyletic nature of Cyperus s.s., there are several other arguments for a new classification of the taxa in the Cyperus clade, and more specifically to reduce the segregate genera into an expanded genus Cyperus. These arguments include: (1) the multiple origins of most of the characters used to circumscribe the segregate genera, such as, switches from spiral to distichous glume arrangement and the other way around, deciduous spikelets, pistil dimerisation and condensation of the inflorescence; (2) the fact that when using the current classification, taxa are often nested in other taxa of the same rank, for example, the genus Volkiella (Merxmüller & Czech 1953: 318) is nested in the genus Lipocarpha (Brown 1818: 459), which is nested in the group Cyperus species using the C4 photosynthetic pathway formerly known as the genus Mariscus (Vahl 1805: 372), and the monophyletic clade of Cyperus species using C4 photosynthesis is in turn nested into the group of Cyperus s.l. species using C3 photosynthesis; (3) the presence of many species with intermediate morphologies; Accepted by Muthama Muasya: 2 Apr. 2014; published: 17 Apr. 2014 33 and (4) the fact that the relationships between the different C4 segregate genera and the C4 Cyperus s.s. species and taxa cannot be entirely resolved with the current techniques because of a fast radiation of species at the base of the clade encompassing all C4 Cyperus s.l. species (Larridon et al. 2013). Larridon et al. (2011a, b) proposed two subgenera under Cyperus based on the type of photosynthesis used which is linked with the anatomy type: (1) Cyperus subgenus Anosporum (Pax 1887: 107) (C 3 photosynthesis and eucyperoid anatomy, paraphyletic but forming a clearly circumscribed natural group) and Cyperus subgenus Cyperus (C4 photosynthesis and chlorocyperoid anatomy, monophyletic). A new sectional classification for Cyperus subgenus Anosporum could already be presented based on a well-resolved phylogeny for the Cyperus s.l. taxa using C3 photosynthesis (Larridon et al. 2011a, b, Fig. 1). The segregate genera Courtoisina (Soják 1979 publ. 1980: 193), Kyllingiella (Haines & Lye 1978: 176) and Oxycaryum (Palla 1908: 169) were formally integrated into new or existing monophyletic sections of Cyperus along with some intermediate species of Cyperus (Larridon et al. 2011b, Larridon & Goetghebeur 2013, Fig. 1). Larridon et al. (2013a) suggested merging the nine C4 segregate genera into Cyperus subgenus Cyperus based on molecular phylogenetic analyses (Fig. 1, see also Bauters et al. 2014). These segregate genera are: Ascolepis (Steudel 1855: 105), Alinula (Raynal 1977: 43), Kyllinga (Rottbøll 1773: 12), Lipocarpha, Pycreus (Beauvois 1816: 48), Queenslandiella (Domin 1915: 415), Remirea (Aublet 1775: 44), Sphaerocyperus (Lye 1972a: 214) and Volkiella. Following this suggestion, Bauters et al. (2014) proposes a new sectional classification under Cyperus for the species formerly placed in the segregate genera Lipocarpha and Volkiella. Also, W. Huygh et al. (unpubl. data) recommend including Kyllinga into Cyperus subgenus Cyperus as a monophyletic section. FIGURE 1. Simplified cladogram of the Cyperus clade based on the results of Larridon et al. (2011a, 2013), adapted from Reynders (2013). Colours: purple = Ficinia clade. Green = C3 Cyperus grade. Red = C4 Cyperus clade. 34 • Phytotaxa 166 (1) © 2014 Magnolia Press LARRIDON ET AL. At this stage it is not possible to present a detailed new subdivisional classification for all groups of Cyperus subgenus Cyperus. However, since the C4 segregate genera are clearly nested in Cyperus s.l. (Muasya et al. 2002, Larridon et al. 2011a, 2013), the species currently included in these segregate genera should be formally included into Cyperus. In this paper, the species that do yet have a Cyperus name, are included in Cyperus either with a new combination or a new name. For the four Alinula species and for the single species of Queenslandiella, Remirea and Sphaerocyperus, Cyperus names were already published by other authors (Vahl 1805, Steudel 1854, Kükenthal 1935–36, 1937, 1943, Kern 1958a, b, Lye 1983b, 1994). As Kyllinga and Pycreus have always been considered as closely related to Cyperus, most Kyllinga and Pycreus species already have (often homotypic) synonyms under Cyperus. Therefore, for Kyllinga and Pycreus, only a few (mostly fairly recent) names still need to be transferred to Cyperus.For Ascolepis, none of the species have a synonym under Cyperus yet, since authors have always considered it as a genus separate from Cyperus. It is important to note that since the type collections are difficult to access, little attention is given here to the Asian species of Pycreus pending an urgent need of confirmation of the taxonomic status of these species. Among them are the Indian taxa described by Govindarajalu (e.g. 1991), who applied an extremely narrow morphological species concept. For example, Prasad (2009) synonymised five of these species with P. malabaricus. Also in the P. pumilus complex, similar reevaluations are needed for the Indian species. Collections of the African species are more accessible, e.g. through JSTOR Global Plants (2013), the Kew Herbarium Catalogue (2013) and the catalogue of the Musée National d’Histoire Naturelle in Paris (2013). Therefore, it was possible to evaluate their taxonomic status here. New combinations or new names are listed for the African taxa we consider to be valid species. Several other species needed to be synonymised. Material and methods We examined a large number of herbarium specimens from B, BM, BR, CEBU, GENT, K, L, LISC, MO, NY, P, SRGH, TAN, UPS, US, WAG and YA (abbreviations according to Holmgren et al., 1900; underlined herbaria have been visited during this study). Our understanding of the taxa was further supplemented with our own observations in the field, and from collections in the Ghent University Botanical Garden. Additional information on species and (type) specimens was obtained from literature (incl. protologues) and the online databases JSTOR Global Plants (2013), Tropicos.org (2013) and Govaerts et al. (2013). An overview of all published generic and infrageneric names is available for the Cyperus clade (Huygh et al. 2010a, Larridon et al. 2011c, Reynders et al. 2011) and served as the nomenclatural base for our efforts to construct a modern classification of the giant genus Cyperus. Images of spikelets presented the figure were taken with a Nikon SMZ800 stereoscopic microscope, equipped with a Nikon digital camera DXM1200 (Nikon, Tokyo, Japan) and edited with Adobe Photoshop CS3 (Adobe Systems Inc., San Jose, USA). Taxonomic treatment ‘Ascolepis’ Cyperus sect. Ascolepis (Nees ex. Steud.) Bauters, stat. & comb. nov. Basionym: Ascolepis Nees ex Steud., Syn. Pl. Glumac. 2: 105 (Steudel 1855). Type: Ascolepis eriocauloides (Steudel 1842: 597) Nees (1855: 105). Synonyms:—Platylepis Kunth (1837: 269), nom. illeg. Lectotype (designated by Huygh et al. 2010a):—Platylepis capensis Kunth (1837: 269) [≡ Ascolepis capensis (Kunth) Ridley (1884: 164)]. Pterachne Schrader ex Nees (1842: 62), nom. nud. in syn. Pterogyne Schrader ex Nees (1842: 62), nom. nud. in syn. Antrolepis Welwitsch (1859: 578), nom. provis. Note:—This section includes, as defined here, 20 species. According to Govaerts et al. (2013), Ascolepis includes 22 species. Since Ascolepis lineariglumis Lye (1983a: 561) and Ascolepis elata Welwitsch (1869: 79) are both included in the complex of Ascolepis protea Welwitsch (1969: 75) s.l., we chose not to combine them into Cyperus as separate species. Also, Ascolepis menonguensis Meneses (1956: 259), a species only known from its COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L. Phytotaxa 166 (1) © 2014 Magnolia Press • 35 type-collection, is probably an aberrant form of Ascolepis protea. Furthermore, we chose not to combine the various infraspecific names under Ascolepis protea into Cyperus pending confirmation of the taxonomic status of these taxa. Cyperus ampullaceus (J.Raynal) Bauters, comb. nov. Basionym: Ascolepis ampullacea J.Raynal, Adansonia, n.s., 13: 159 (Raynal 1973). Type:—ZAMBIA. Northern Province: Mporokoso District, Five miles north of Muzombwe, west side of Mweru-wa-Ntipa, 3500 ft, 16 April 1961, J.B. Phipps & L.D.E.F. Vesey-FitzGerald 3233 (holotype NY!, isotypes K!, P!, SRGH!). Description:—Goetghebeur (1980: 296). Cyperus ascocapensis Bauters, nom. nov. Basionym: Platylepis capensis Kunth, Enum. Pl. 2: 269 (Kunth 1837) ≡ Ascolepis capensis (Kunth) Ridley (1884: 164), non Cyperus capensis (Steudel 1829: 153) Endlicher (1842: 94). Type:—SOUTH AFRICA, Omsamcaba, 12 May 1832, J.F. Drège 4389 (isotype P!). Synonyms:—Platylepis dioica Steudel (1855: 131), non Cyperus dioicus Johnston (1924: 988). Type:—SOUTH AFRICA. Katberg: 4500–5000 ft, 13 November 1832, J.F. Drège 3953 (holotype P! isotypes P!) Ascolepis capensis var. lacera C.B.Clarke in Durand & Schinz (1894: 651), nom. nud. Ascolepis capensis var. pleiostachya Kükenthal (1921: 9). Type: not located. Description:—Goetghebeur (1980: 291). Cyperus ascodensus Goetgh., nom. nov. Basionym: Ascolepis densa Goetgh., Adansonia, n.s., 19: 285 (Goetghebeur 1980), non Cyperus densus Link (1820: 83). Type:—ZAMBIA. Mapanza: Choma, 20 March 1958, E.A. Robinson 2814 (holotype SRGH!, isotypes K!, M!, P, PRE!). Description:—Goetghebeur (1980: 285). Cyperus ascofibrillosus Goetgh., nom. nov. Basionym: Ascolepis fibrillosa Goetgh., Bull. Jard. Bot. Natl. Belg. 47: 439 (Goetghebeur 1977), non Cyperus fibrillosus Kükenthal (1921: 1). Type:—CONGO. 19 February 1954, R. Devred 1517 (holotype BR!, isotypes BRVU, K!, P, SRGH!). Description:—Goetghebeur (1980: 289). Cyperus ascohemisphaericus Goetgh., nom. nov. Basionym: Ascolepis hemisphaerica Peter ex Goetgh., Adansonia, n.s., 19: 283 (Goetghebeur 1980), non Cyperus hemisphaericus Boeckeler (1859: 436). Lectotype (designated by Goetghebeur 1980: 285):—BURUNDI. Bei Niakassu östlich, 4 March 1926, A. Peter 38250 (holotype B!, isotypes B!, K!, P, SRGH!). Description:—Goetghebeur (1980: 283). Cyperus asconeglectus Goetgh., nom. nov. Basionym: Ascolepis neglecta Goetgh., Bull. Jard. Bot. Natl. Belg. 47: 441 (Goetghebeur 1977), non Cyperus neglectus Parlatore (1846: 131). Type:—CONGO. Kilambwe (Kamina), 23 March 1955, S. Risopoulos 389 (holotype BR!). Description:—Goetghebeur (1980: 287). Cyperus ascopinguis Goetgh., nom. nov. Basionym: Ascolepis pinguis C.B.Clarke, Ann. Mus. Congo Belge, Bot., II, 1(2): 69 (Clarke 1900), non Cyperus pinguis (Clarke) Mattfeld & Kükenthal (Kükenthal 1936: 583). Lectotype (designated by Goetghebeur 1980: 295):—CONGO. [Kalemie], June 1894, G. Descamp s.n. (holotype BR!). Description:—Goetghebeur (1980: 295). Cyperus ascopusillus Goetgh., nom. nov. Basionym: Ascolepis pusilla Ridl., Trans. Linn. Soc. London, Bot. 2: 164 (Ridley 1884), non Cyperus pusillus Vahl (1805: 303). Type:—ANGOLA. Circa Lopollo et Monino, May 1860, F.M.J. Welwitsch 1678 (holotype BM!, isotype BR!). Synonyms:—Lipocarpha multibracteata Clarke (1902: 472), non Cyperus multibracteatus Boeckeler (1875: 107). Type:— ANGOLA. District Huilla, May 1860, F.M.J. Welwitsch 6773 (holotype BM!, isotype BR!). Ascolepis pusilla var. cylindrica Hooper (1983: 608). Type:—TANZANIA. Iringa, just north of township, 1650 m, 15 July 1956, E. Milne-Redhead & P. Taylor 11202 (holotype K!). Ascolepis pusilla var. echinata Hooper (1983: 607). Type:—TANZANIA, T4, Sumbawanga to Mpanda road, 8 km north of Sumbawanga, Fiengalezia villaga, 10 June 1980, S.S. Hooper, C.C. Townsend & L. Mwasumbi 1927 (holotype K!). 36 • Phytotaxa 166 (1) © 2014 Magnolia Press LARRIDON ET AL. Ascolepis pusilla var. microcuspis Lye (1983a: 564). Type:—ZAMBIA, Mazabuka, Siamambo, Choma, 4300ft, 20 March 1958, E.A. Robinson 2815 (holotype K!, isotype PRE!). Description:—Goetghebeur (1980: 297). Cyperus ascospinulosus Goetgh., nom. nov. Basionym: Ascolepis spinulosa Goetgh., Bull. Jard. Bot. Natl. Belg. 47: 438 (Goetghebeur 1977), non Cyperus spinulosus Roxburgh (1820: 207). Type:—CONGO. Haut Katanga: environs de Lubudi, 1937, D. Cabu s.n. (holotype BR!). Description:—Goetghebeur (1980: 287). Cyperus ascotrigonus Goetgh., nom. nov. Basionym: Ascolepis trigona Goetgh. (Goetghebeur 1980: 286), non Cyperus trigonus Boeckeler (1888: 11). Type:— ZAMBIA. 10 km East of Kasama, 12 January 1961, E.A. Robinson 4253 (holotype SRGH!, isotypes BR!, K, M!, P, PRE!). Description:—Goetghebeur (1980: 286). Cyperus brasiliensis (Kunth) Bauters, comb. nov. Basionym: Platylepis brasiliensis Kunth, Enum. Pl. 2: 269 (Kunth 1837) ≡ Ascolepis brasiliensis (Kunth) Benth. ex C.B.Clarke (Clarke 1894: 651). Type:—BRAZIL. Sellow s.n., (holotype K!, isotype P!, HAL!). Synonyms:—Platylepis gujanensis Nees (1842: 63) ≡ Kyllinga decora Steudel (1855: 317), nom. illeg. Type:—GUYANA. Schomburgk 109 (holotype: B, isotypes BM, K!, U!, W). Platylepis leucocephala Nees (1842: 63). ≡ Ascolepis leucocephala (Nees) Eiten (1963: 221). Lectotype (designated here):— BRAZIL. H.K. Beyrich s.n. (P). Platylepis xanthocephala Nees (1842: 62). Type:—BRAZIL. February 1837, G. Gardner 715 (holotype: K!, isotypes BM!, K!, P!). Description:—Goetghebeur (1980: 293). Cyperus dipsacoides (Schumach.) Bauters, comb. nov. Basionym: Kyllinga dipsacoides Schumach., Beskr. Guin. Pl.: 41 (Schumacher 1827) ≡ Ascolepis dipsacoides (Schumach.) Raynal (1968: 99). Type:—GUINEA. P. Thonning s.n. (holotype C!, isotypes C!). Synonyms:—Ascolepis setigera Hutchinson (1936: 472), nom. inval. Description:—Goetghebeur (1980: 299). Cyperus eriocauloides (Steud.) Bauters, comb. nov. Basionym: Kyllinga eriocauloides Steud., Flora 25: 597 (Steudel 1842) ≡ Isolepis ascolepis Richard (1850–51: 501) ≡ Ascolepis eriocauloides Neex ex Steud. (Steudel 1855: 105). Type:—ETHIOPIA. Prope Gafta, 15 September 1838, W. Schimper 1195 (holotype P!, isotypes B!, BM!, BR!, GH!, HOH!, K!, L, MO!, P!, S!, TUB!, WAG!). Description:—Goetghebeur (1980: 283). Cyperus erythrocephalus (S.S.Hooper) Bauters, comb. nov. Basionym: Ascolepis erythrocephala S.S.Hooper, Kew Bull. 37: 605 (Hooper 1983). Type:—TANZANIA. About 6.5 km west of Songea, 990 m, 28 April 1956, E. Milne-Redhead & P. Taylor 9940 (holotype K!, isotypes BM!, NY!, P!). Description:—Hoenselaar et al. (2010: 269). Cyperus majestuosus (P.A.Duvign. & G.Léonard) Bauters, comb. nov. Basionym: Ascolepis majestuosus P.A.Duvign. & G.Léonard, Bull. Soc. Roy. Bot. Belgique 90: 188 (Duvigneaud 1958). Type:—CONGO. Duvigneaud & Timperman 2361 A 1 (holotype BRLU!). Description:—Goetghebeur (1980: 299). Cyperus metallorum (P.A.Duvign. & G.Léonard) Bauters, comb. nov. Basionym: Ascolepis metallorum P.A.Duvign. & G.Léonard, Bull. Soc. Roy. Bot. Belgique 90: 268 (Duvigneaud 1958). Type:—CONGO, Duvigneaud 3061 (holotype BRLU). Description:—Goetghebeur (1980: 281). Cyperus pseudopeteri (Goetgh.) Bauters, comb. nov. Basionym: Ascolepis pseudopeteri Goetgh., Adansonia, n.s., 19: 286 (Goetghebeur 1980). Type:—ZAMBIA. District Mwinilunga, Zambezi Rapids, 3 miles North of Kalene Hill, 4300 ft, 24 December 1969, B.K. Simon & G. Williamson 1991 (holotype SRGH, isotypes K!, P). Description:—Goetghebeur (1980: 286). COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L. Phytotaxa 166 (1) © 2014 Magnolia Press • 37 Cyperus proteus (Welw.) Bauters, comb. nov. Basionym: Ascolepis protea Welw., Trans. Linn. Soc. London 27: 75 (Welwitsch 1869). Type:—ANGOLA. Pungo Adongo: prope Funda-Quilombo, December 1856, F.M.J. Welwitsch 1667 (holotype BM!, isotype K!). Synonyms:—Ascolepis elata Welwitsch (1869: 79). Type:—ANGOLA. F.M.J. Welwitsch 1670 (holotype BM, isotype K). Ascolepis menonguensis Meneses (1956: 259). Type:—ANGOLA. Gossweiler 3189 (holotype LISC, isotype K). Ascolepis lineariglumis var. lineariglumis Lye (1983a: 561). Type:—ZAMBIA. 13 km NW of Kabwe-Bonanza, 14°22’S, 28°23’E, 1200 m, 8 April 1972, J. Kornas 1559 (holotype KRA, isotype K). Ascolepis lineariglumis var. pulcherrima Lye (1983a: 562). Type:—ZAMBIA. Abercorn district, Saisi River Marsh, 1500 m, 27 February 1957, H.M. Richards 8353 (holotype K). Description:—Goetghebeur (1980: 275). Cyperus siamensis (C.B.Clarke) Bauters, comb. nov. Basionym: Scirpus squarrosus var. siamensis C.B.Clarke, Beih. Bot. Centralbl. 27 (2): 460 (Clarke 1910) ≡ Scirpus chinensis var. siamensis (C.B.Clarke) Raymond (1957: 124) ≡ Scirpus siamensis (C.B.Clarke) Kern (1958a: 219) ≡ Ascolepis dipsacoides subsp. siamensis (C.B.Clarke) Raynal (1968: 99). Type:—THAILAND. [Wang Chao], 100m, 14 October 1904, C.C. Hosseus 101 (holotype K, isotype B!, P, M!). Synonym:—Ascolepis gracilis Turrill (1915: t. 3020), non Cyperus gracilis Brown (1810: 213). Type:—THAILAND. Kerr 2261 (holotype: K, isotype: BM) Description:—Goetghebeur (1980: 299). Cyperus vatkeanus (Boeckeler) Goetgh., comb. nov. Basionym: Ascolepis vatkeana Boeckeler, Allg. Bot. Z. Syst. 2 (4): 55 (Boeckeler 1896). Type:—NAMIBIA, Owamboland, Dr. C. Höpfner 82 (isotype Z). Synonyms:—Ascolepis speciosa Welwitsch (1869: 78), non Cyperus speciosus Vahl (1805: 364). Type:—ANGOLA. Prope Lopollo, March 1860, F.M.J. Welwitsch 1674 (holotype BM!). Ascolepis oliveri Vatke & Höpfner ex C.B.Clarke in Durand & Schinz (1894: 652), nom. nud. Description:—Goetghebeur (1980: 289). ‘Kyllinga’ Cyperus afro-occidentalis (Lye) Huygh, comb. nov. Basionym: Kyllinga afro-occidentalis Lye, Nordic J. Bot. 1: 744 (Lye 1981 publ. 1982). Type:—CÔTE D’IVOIRE. Zanzan, along road from Bouna to Ferkessédougou, 9°35’24”N, 4°18’23”W, 4 August 1967, Geerling & Bokdam 448 (holotype WAG, isotypes BR656056!, K00416263!, P00584993!). Description:—Lye (1981 publ. 1982: 744). Cyperus auratus (Nees) Huygh, comb. nov. Basionym: Kyllinga aurata Nees, Linnaea 10: 139 (Nees 1836) ≡ Kyllinga erecta var. aurata (Nees) Kükenthal (1913: 91) ≡ Cyperus erectus var. auratus (Nees) Kükenthal (1936: 589) ≡ Kyllinga colorata var. aurata (Nees) Lye (1972b: 218). Lectotype (designated here):—SOUTH AFRICA. Eastern Cape: Zwartskop River, 152–305 m, March 1846, Ecklon & Zeyher s.n. (lectotype P00573301!). Synonyms:—Kyllinga consanguinea Kunth (1837: 135). Lectotype (designated here):—SOUTH AFRICA. Glenfilling, below 305 m, 1 January 1832, Drège s.n. (lectotype P00573210!, isolectotype P00573209!). Kyllinga intricata Chermezon (1919: 211) ≡ Kyllinga erecta var. intricata C.B. Clarke (1894: 529) nom. nud. ≡ Cyperus erectus var. intricatus (Chermezon) Kükenthal (1936: 590) Kyllinga brevifolia subsp. intricata (Chermezon) Lye (1981 publ. 1982: 747) ≡ Cyperus brevifolius subsp. intricatus (Chermezon) Lye (Haines & Lye 1983, App. 3: 2). Type:— MADAGASCAR. Itasy, Imerina, December 1880, Hildebrandt 3788 (holo-: K000416337!). Description:—Nees (1836: 139). Cyperus beninensis (Samain, Reynders & Goetgh.) Huygh, comb. nov. Basionym: Kyllinga beninensis Samain, Reynders & Goetgh., Novon 16: 516 (Samain et al. 2006). Type:—BENIN. Borgou: Tchaourou, Oauri-Maro, 333 m, 27 August 1999, Sinsin 3038 (holotype: WAG0087242!). Description:—Samain et al. (2006: 516). Cyperus brunneoalatus (Cherm.) Huygh, comb. nov. Basionym: Kyllinga brunneoalata Cherm., Bull. Soc. Bot. France 83: 492 (Chermezon 1936). Lectotype (designated here):— 38 • Phytotaxa 166 (1) © 2014 Magnolia Press LARRIDON ET AL. GABON. Woleu-Ntem: Abam, 662 m, 1°37’59”N, 11°22’0”E, 13 May 1933, Le Testu 9128 (lectotype P00573142!, isolectotypes BR000000871945!, K000416271!, P00573140!, P00573141!). Description:—Chermezon (1936: 492). Cyperus cardosoi (Meneses) Huygh, comb. nov. Basionym: Kyllinga cardosoi Meneses, Garcia de Orta 4: 240 (Meneses 1956). Type:—ANGOLA. Bié: Silva Porto (Kuito), 1500 m, s.d., Cardoso s.n. (not located). Description:—Meneses (1956: 240). Cyperus carinalaevis (Lye & Mesterházy) Huygh, comb. nov. Basionym: Kyllinga carinalaevis Lye & Mesterházy Nordic J. Bot. 30: 385 (Lye & Mesterházy 2012). Type:—BENIN. Atakora: Boukoumbé, 232 m, 1°06′00″N, 10°09′56″W, 7 December 2009, A. Mesterházy MABEN 56 (holotype: K, isotypes: BP, BENIN). Description:—Lye & Mesterházy (2012: 385). Cyperus cataphyllatus (Huygh & Schouppe) Huygh, comb. nov. Basionym: Kyllinga cataphyllata Huygh & Schouppe, Blumea 55: 291 (Huygh et al. 2010b). Type:—BURUNDI. Bujumbura Rural: S. of Bugurama, direction Ijenda to Kigezi, 2080 m, 27 January 1984, Van der Veken 1984-123 (holotype GENT151693!, isotype GENT!). Description:—Huygh et al. (2010b: 291). Cyperus chrysanthoides (Mtot.) Huygh, comb. nov. Basionym: Kyllinga chrysanthoides Mtot., Bot. Jahrb. Syst. 111: 292 (Mtotomwema 1990a). Type:—BURUNDI. Bururi: Munini, 1900 m, 4°0’0”S, 29°45’0”E, 13 January 1977, Reekmans 6736 (holotype K, isotypes GENT!, P). Description:—Mtotomwema (1990a: 292). Cyperus ghanechinatus Huygh, nom. nov. Basionym: Kyllinga echinata S.S.Hooper, Kew Bull. 26: 580 (Hooper 1972), non Cyperus echinatus (Linnaeus 1753: 50) Alph.Wood (Wood 1861: 734). Type:—GHANA. Northern: between Damongo and Kpirri Lake, 194 m, 9°4’4”N, 1°48’36”W, 31 March 1956, Adams 3961 (holotype K000416264!, isotypes GC, P00584871!). Description:—Hooper (1972: 580). Cyperus mbitheanus (Muasya) Huygh, comb. nov. Basionym: Kyllinga mbitheana Muasya, J. E. Afr. Nat. Hist. 60: 66 (Muasya et al. 2010). Type:—KENYA. Eastern: Machakos District, Kindaruma, Muasya & Musili 2658 (holotype EA, isotypes BOL!, K). Description:—Muasya et al. (2010: 66). Cyperus mindorensis (Steud.) Huygh, comb. nov. Basionym: Kyllinga mindorensis Steud., Syn. Pl. Glumac. 2: 67 (Steudel 1854). Type:—PHILIPPINES. Luzon: Calabarzon, Batangas, 1841, Cuming 1558 (holotype BM000959042!, isotypes K000290929!, P00578843!). Synonyms: — Scirpus cephalotes Jacquin (1771: 42), nom. illeg., non S. cephalotes Linnaeus (1762: 76) ≡ Kyllinga cephalotes (Jacq.) Druce (1917: 630), non Cyperus cephalotes Vahl (1805: 311). Type:—Not located. Thryocephalon nemorale J.R.Forster & G.Forster (1775: 129, pl. 65) ≡ Kyllinga nemoralis (J.R.Forster & G.Forster) Dandy ex Hutchinson & Dalziel (1936: 487), non Cyperus nemoralis Chermezon (1921: 553). Type:—FRENCH POLYNESIA. Tahiti: J.R. Forster & G. Forster s.n. (lectotype BM000990844!). Kyllinga monocephala Stokes (1812: 120) nom. illeg., non K. monocephala Rottbøll (1773: 13). Type:—JAMAICA, Broughton s.n. (not located). Remarks: K. monocephala Rottb. is a superfluous name. It is a homotypic synonym of Rhynchospora colorata (Linneaus) Pfeiffer (1935: 89) and has Schoenus coloratus Linnaeus (1753: 43) as basionym. Cyperus kyllingia Endlicher (1842: 94) is the homotypic synonym of K. monocephala Rottb. and consequently this name is also a homotypic synonym of R. colorata. Kyllinga monocephala var. subtriceps Kunth (1837: 130) ≡ Cyperus kyllingia f. subtriceps (Kunth) Kükenthal (1936: 608). Type: not located. Kyllinga gracilis Zollinger (1854: 63) nom. illeg., non K. gracilis Kunth (1837: 134). Type:—Not located. Cyperus leucocephalus Hasskarl (1848: 87) nom. illeg., non C. leucocephalus Retzius (1788: 11). Type:—Not located. Kyllinga monocephala var. humilis Boeckeler (1868: 429) ≡ Cyperus kyllingia f. humilis (Boeckeler) Kükenthal (1936: 608). Syntypes:—GUAM. Haenke s.n. (not located); s.l., Herb. Link s.n. (not located). COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L. Phytotaxa 166 (1) © 2014 Magnolia Press • 39 Kyllinga monocephala var. tenuis Boeckeler (1868: 429) ≡ Cyperus kyllingia f. tenuis (Boeckeler) Kükenthal (1936: 608). Syntypes:—MARIANA ISLANDS. Gaudichaud s.n. (not located); INDONESIA. Timor: Gaudichaud s.n. (not located). Kyllinga monocephala var. latifolia Boeckeler (1868: 429) ≡ Kyllinga seychellarum A. Jussieu ex C.B.Clarke (Clarke 1894: 531) ≡ Kyllinga planiculmis Boivin ex C.B.Clarke (Clarke 1894: 531) ≡ Cyperus kyllingia var. latifolius (Boeckeler) Kükenthal (1936: 608 p.p.). Type:—MAYOTTE. Boivin 3066 (not located). Kyllinga planiculmis var. mucronata Chermezon (1925: 613). Type:—MADAGASCAR. Antsinanana, Mahanoro, confluent de l’Onive et du Mangoro, 600 m, February 1925, Perrier de la Bâthie 17010 (holotype TAN!). Cyperus curvispiculosus Koyama (1955: 215) ≡ Kyllinga curvispiculosa (T.Koyama) P.H.Hô (Hô 1993: 699). Type:— VIETNAM. Lam Dong, Annam, Dran, 1034 m, 11°48’51”N, 108°34’34”E, 18 June 1921, Hayata 727 (holotype P00065565!) Description:—Hoenselaar et al. (2010: 335). Cyperus ngothe (Mtot.) Huygh, comb. nov. Basionym: Kyllinga ngothe Mtot., Bot. Jahrb. Syst. 111: 294 (Mtotomwema 1990a). Type:—KENYA. Central: Kitui district, Endau, 19 November 1979, Gatheri, Mungai & Kibui 79/89 (holotype EA!). Description:—Mtotomwema (1990a: 294). Cyperus nigriceps Huygh, nom. nov. Basionym: Kyllinga erecta var. lurida Kük., Notizbl. Bot. Gart. Berlin-Dahlem 9: 300 (Kükenthal, 1925) ≡ Cyperus erectus var. luridus (Kük.) Kükenthal (1936: 590) ≡ Kyllinga aurata var. lurida (Kük.) Napper (1971: 21) ≡ Kyllinga colorata var. lurida (Kük.) Lye (1972b: 218) ≡ Kyllinga brevifolia subsp. lurida (Kük.) Lye (1981 publ. 1982: 747) ≡ Cyperus brevifolius subsp. luridus (Kük.) Lye in Haines & Lye (1983: App. 3: 2) ≡ Kyllinga brevifolia var. lurida (Kük.) Beentje in Hoenselaar et al. (2010: 315), non Cyperus luridus as ‘Cyperus lurida’ Govindarajalu (1975: 189). Lectotype (designated here):—KENYA. Central: Mount Aberdare, Forest Station, 2300 m, 0°30’0”S, 36°40’0”E, 26 December 1921, R.E. & T.C.E. Fries 367 (lectotype K000416311!, isolectotype B10 0278625!). Description:—Hoenselaar et al. (2010: 315). Cyperus njombensis Huygh, nom. nov. Basionym: Kyllinga uniflora Mtot., Nordic J. Bot. 9: 640 (Mtotomwema 1990b), non Cyperus uniflorus Thunberg (1825: 8). Type:—TANZANIA. Njombe District, Njombe-Songea road, Mhoro 3934 (holotype K!, isotype: DAR). Description:—Hoenselaar et al. (2010: 342). Cyperus rukwanus Huygh, nom. nov. Basionym: Kyllinga alba-purpurea Lye, Nordic J. Bot. 1: 743 (Lye 1981 publ. 1982) ≡ Cyperus alba-purpureus (Lye) Lye (Haines & Lye 1983, App. 3: 2) non Cyperus albopurpureus Chermezon (1920: 344). Type:—UNITED REPUBLIC OF TANZANIA. Rukwa: Nsangu Forest area, 2000 m, 2 January 1962, Robinson 4863 (holotype K000416318!, isotype SRGH0106170-0!). Description:—Lye (1981 publ. 1982: 743). Cyperus ruwenzoriensis (C.B.Clarke) Huygh, comb. nov. Basionym: Kyllinga ruwenzoriensis C.B.Clarke in D.Oliver & auct. suc. (eds.), Fl. Trop. Afr. 8: 283 (Clarke 1901) ≡ Cyperus oblongus var. ruwenzoriensis (C.B.Clarke) Kükenthal (1936: 580) ≡ Kyllinga nervosa var. ruwenzoriensis (C.B.Clarke) Lye (1972b: 218). Type:—UGANDA. Western: Ruwenzori, Kivata, 2438 m, 0°22’41”N, 29°54’18”E, s.d., Scott-Elliot 7554 (holotype K000416308!, isotypes B10 0278623!, BM000922547!). Synonyms:—Kyllinga elatior Kunth (1837: 135), non Cyperus elatior Boeckeler (1870) ≡ Kyllinga polyphylla var. elatior (Kunth) Kükenthal (1925: 300) ≡ Cyperus aromaticus var. elatior (Kunth) Kükenthal (1936: 582). Type:—SOUTH AFRICA. Eastern Cape: Between Umtata Prov. and St Johns River Pondoland, 31°37’45”S, 29°12’4”E, 305 m, 7 February 1832, Drège 4384 (holotype B10 0166793!, isotypes K000416349!, K000416350!, K000416351!, P00573190!, P00573191!, P00573192!). Kyllinga pinguis Clarke (1906: 131) ≡ Cyperus pinguis (C.B.Clarke) Mattf. & Kük. (Kükenthal 1936: 583). Syntypes:— UNITED REPUBLIC OF TANZANIA. Tanga: Kwai, 1500 m, November 1899, Stern 235 (syntype B10 0278305!); UGANDA. Entebbe, Brown 26 (not located); KENYA. Nairobi, Linton 7 (not located). Description:—Hoenselaar et al. (2010: 345). Cyperus serratangulus (Peter & Kük.) Huygh, comb. nov. Basionym: Cyperus cartilagineus var. serratangulus Peter & Kük. in H.G.A.Engler (ed.), Pflanzenr., IV, 20 (101): 609 (Kükenthal 1936). Typre:—UNITED REPUBLIC OF TANZANIA. Singida: Uyansi, Chaya near Lake Chaya, 1240 m, 4 January 1926, A. Peter 45780 (holotype B10 01655685!, isotypes B10 01655686!). Description:—Kükenthal (1936: 609). 40 • Phytotaxa 166 (1) © 2014 Magnolia Press LARRIDON ET AL. ‘Pycreus’: new names and combinations in Cyperus Cyperus africanus (S.S.Hooper) Reynders, comb. nov. Basionym: Pycreus divulsus subsp. africanus S.S.Hooper, Kew Bull. 26: 579 (Hooper 1972) ≡ Pycreus africanus (S.S.Hooper) Reynders (Reynders & Goetghebeur 2010: 227). Type:—CAMEROON. Gaudua, foothills of Gotel Mountains, 17 July 1969, J.B. Hall 1381 (holotype K!, isotype P!). Description:—Reynders & Goetghebeur (2010: 227). Cyperus acaulescens Reynders, nom. nov. Basionym: Pycreus acaulis Nelmes, Kew Bull. 10: 91 (1955), non Cyperus acaulis Steudel (1842: 599). Type:—MALAWI. Nyika Plateau, Kaulime Pond, 27 June 1952, G. Jackson 870 (holotype K!, isotype BR!). Description:—Nelmes (1955: 91). Cyperus neocooperi Reynders, nom. nov. Basionym: Pycreus cooperi C.B.Clarke in W.H.Harvey & auct. suc. (eds.), Fl. Cap. 7: 160 (Clarke 1897) ≡ Cyperus cooperi (C.B.Clarke) Kükenthal (1934: 68) nom. illeg., non. Cyperus cooperi (C.B.Clarke) K.Schum. (1900: 328). Type:— SOUTH AFRICA, Orange Free State, 17 January 1861, T. Cooper 912 (holotype K!). Description:—Clarke (1897: 160). Note:—Cyperus neocooperi is very similar to C. aethiops, which has smaller glumes and a more northerly, afromontane distribution. Possibly, both could even be considered as conspecific. Cyperus aethiops belongs to Cyperus section Pycreus (C. sect. Polystachyi Kük., nom. illeg.), and C. neocooperi also fits in this group based on its elongated elliptic nutlets. Kükenthal (1936) placed C. neocooperi species in the very heterogenous C. sect. Lancei based on its rather large and dark glumes. However, blackish glumes are correlated with Cyperus species growing at higher elevation and originated multiple times within the genus, also, glume size does not seem to be reliable for infrageneric classification (Reynders 2013). Cyperus okavangensis (Podlech) Reynders, comb. nov. Basionym: Pycreus okavangensis Podlech, Mitt. Bot. Staatssamml. München 3: 522 (Podlech 1960). Type:—NAMIBIA, [Rundu], 11 May 1939, O.H. Volk 1966 (holotype M!, isotype PRE!). Description:—Podlech (1960: 522). Cyperus poikilostachys (Nelmes) Reynders, comb. nov. Basionym: Pycreus poikilostachys Nelmes, Kew Bull. 6: 320 (Nelmes 1951 publ. 1952). Type:—ZAMBIA. Mwinilunga District, 0.5 mile South of Matonchi Farm, 24 January 1938, E. Milne-Redhead 4311 (holotype K!, isotypes BR!, K!, PRE!). Description:—Nelmes (1951 publ. 1952: 320). Cyperus poikilostachys var. heterochrous (Nelmes) Reynders, comb. nov. Basionym: Pycreus heterochrous Nelmes, Kew Bull. 6: 321 (1951 publ. 1952). Type:—ZAMBIA, Mwinilunga District, Matonchi Farm, 0.5 mile South of farm, 24 January 1938, E. Milne-Redhead 4309 (holotype K, isotypes BR!, P!, isotype PRE!). Description:—Nelmes (1951 publ. 1952: 321) Note:—Several specimens collected by E. Milne-Redhead in the Zambezian region were simultaneously described as new species by Nelmes, among which P. poikilostachys and P. heterochrous. The latter two species only differ in their glume colour: Pycreus poikilostachys has dark brown glumes and P. heterochrous has pale reddish brown glumes. As both are sympatric with an absence of individuals showing intermediate glume colours, Nelmes (1938) considered both colour variants as different species. Glume colour alone is now considered to be an unreliable character for species delimitation in Cyperus (Reynders 2013). In addition, as is known from classic Mendelian genetics, it is possible that different colour variants can be present in the same populations without the presence of intermediates (dominant-recessive inheritance instead of intermediate inheritance). Different colour variants of a single species are usually treated on the variety rank or below. COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L. Phytotaxa 166 (1) © 2014 Magnolia Press • 41 Cyperus poikilostachys var. poikilostachys (autonym, automatically established here). Cyperus scaettae (Cherm.) Reynders Basionym: Pycreus scaettae Cherm., Rev. Zool. Bot. Afr. 24: 295 (Chermezon 1933 publ. 1934). Lectotype (designated here):—DEMOCRATIC REPUBLIC OF CONGO, Mubeza, 1930, H. Scaetta 58M (hololectotype BR!, isolectotype K!). Description:—Hoenselaar et al. (2010: 296). Note:—Cyperus scaettae belongs to a group of species from Zambezian Africa showing thick accumulations of fibres (remains of leaf sheaths) surrounding the bases of the culms and most often inflorescences are reduced, with only a few spikelets set in a single spike. With the description of different taxa showing these characteristics (e.g. Pycreus fibrillosus, P. scaettae, Fig. 2), relatively less attention has been paid to spikelets, glume and nutlet morphology than to the striking culm bases and glume colour. In addition, communication and consultation of type material seemed to have been rather limited in the short period these taxa have been described, combined and synonymised by different authors (Chermezon 1932, 1933, 1933 publ. 1934, Kükenthal 1921, 1935–36). Subsequently, species separations and especially synonymisations were not very clear from the beginning and several misinterpretations arose in later publications (e.g. Haines & Lye 1983). Comparision of spikelet characteristics shows Cyperus scaettae clearly differs from C. fibrillosus (Fig. 2). FIGURE 2. Comparison of the spikelets (a & c) and rachillas (b & d) of two fibrous “Pycreus” species clearly showing the difference between Cyperus fibrillosus (a & b) and C. scaettae (c & d). Cyperus fibrillosus has a flexuous rachilla that is hardly visible between the glumes. Lower glumes mostly have 1–2 additional nerves on their wings. Cyperus scaettae has larger spikelets with strongly imbricate glumes and a straight rachilla that is visible between the glumes. 42 • Phytotaxa 166 (1) © 2014 Magnolia Press LARRIDON ET AL. Cyperus scaettae var. vanderystii (Cherm.) Reynders, comb. nov. Basionym: Pycreus vanderystii Cherm., Rev. Zool. Bot. Africaines 24: 296 (Chermezon 1933 publ. 1934) ≡ Cyperus fibrillosus var. vanderystii (Cherm.) Kük. (Kükenthal 1936: 348). Lectotype (designated here):—CONGO, Vanderyst 16469 (hololectotype BR!). Description:—Chermezon (1934: 296). Note:—Chermezon (1933 publ. 1934) already remarked that Pycreus vanderystii is closely related to P. scaettae when describing both taxa in the same publication. They have the same habit, spikelet and glume shapes, and their rachillas are straight. Pycreus vanderystii only differs in its overall larger dimensions of the plant and of the glumes, and its yellowish glume colour. Although most plants can easily be placed in one of both taxa based on the average size of the glumes, there is a slight overlap in the glume size range. Since it concerns sympatric taxa, recognising P. vanderystii at variety level seems most appropriate. As discussed under Cyperus scaettae, C. fibrillosus is clearly distinct because of its flexuous rachilla and different glumes. Therefore, P. vanderystii should not be included into the latter species as was done by Kükenthal (1936). Cyperus scaettae var. scaettae (autonym automatically established here). Notes on synonymy of an African ‘Pycreus’ species Pycreus sanguineosquamatus has not yet been combined into Cyperus. However, in our opinion, this name represents a species that needs to be placed in synonymy of an older species, i.e. Cyperus fontinalis. Therefore, a new combination into Cyperus would be superfluous. Pycreus sanguineosquamatus Van der Veken (1955: 145). Type:—DEMOCRATIC REPUBLIC OF CONGO, May 1939, H. Bredo 2750 (holotype BR!, isotypes BR!, C!, NY!, P!, PRE!). = Cyperus fontinalis (Cherm.) Kükenthal (1935–36: 341) Basionym: Pycreus fontinalis Cherm., Bull. Soc. Bot. France 67: 327 (Chermezon 1920 publ. 1921). Type:—MADAGASCAR, March 1920, H. Perrier de la Bâthie 13056 (holotype P!, isotypes K!, P!). Description:—Kükenthal (1935–36: 341). Note:—Pycreus fontinalis is a rare species only known from its type locality near the hot water springs of Antsirabe, Madagascar. Pycreus sanguineosquamatus is described on material from salt marshes of volcanic origin in the region of Shaba (Democratic Republic of Congo). With the description of the latter species, Van der Veken (1955) already noted the similarity of this taxon with P. fontinalis. However, P. fontinalis was circumscribed by Chermezon (1920 publ. 1921) and (1935–36) as having a tiny rhizome. As Van der Veken (1955) interpreted the material from Congo as therophytic, the two species were assumed to be separated by their growth form. However, when comparing the type material of both taxa, this distinction of growth forms is, in our opinion, no more than a matter of interpretation as no striking differences could be observed between the plant bases of the Congolean and Malagasy material. In addition, glumes and nutlets (the taxonomically most important characteristics in this group of Cyperus) are very similar. The type locality of Cyperus fontinalis in Antsirabe seems to have been destroyed due to exploitation of the hot water from the springs by the local community and the species may even be locally extinct (field observations in Madagascar, April 2010). The salt marches in southern Congo are similarly being exploited. The salt marshes of this region are of volcanic origin and provide a habitat to a large number of endemic plant species that are adapted to the mineral rich soils (Symoens 1953) such as several short lived (endemic) Cyperus and Bulbostylis species. Considering the strongly local geographic distribution of the species (two known localities) with a possible extinction from Antsirabe, small number of plants and continuing human exploitation at both localities, we propose to give Cyperus fontinalis the status of Critically Endangered according to the IUCN categories and criteria (IUCN 2012). COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L. 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