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Taxonomic changes in C4 Cyperus (Cypereae, Cyperoideae, Cyperaceae): Combining the sedge genera Ascolepis, Kyllinga and Pycreus into Cyperus s.l.

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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.
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Accepted by Muthama Muasya: 2 Apr. 2014; published: 17 Apr. 2014 33
PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Copyright © 2014 Magnolia Press
Phytotaxa 166 (1): 033–048
www.mapress.com/phytotaxa/Article
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 LARRIDON1, KENNETH BAUTERS1, MARC REYNDERS1,2, WIM HUYGH1 & PAUL
GOETGHEBEUR1
1Ghent University, Department of Biology, Research Group Spermatophytes, K.L. Ledeganckstraat 35, BE-9000 Gent, Belgium;
isabel.larridon@ugent.be, kenneth.bauters@ugent.be
2Botanic 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;
LARRIDON ET AL.34 Phytotaxa 166 (1) © 2014 Magnolia Press
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) (C3 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.
Phytotaxa 166 (1) © 2014 Magnolia Press 35
COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L.
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
LARRIDON ET AL.36 Phytotaxa 166 (1) © 2014 Magnolia Press
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!).
Phytotaxa 166 (1) © 2014 Magnolia Press 37
COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L.
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 (185051: 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).
LARRIDON ET AL.38 Phytotaxa 166 (1) © 2014 Magnolia Press
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):—
Phytotaxa 166 (1) © 2014 Magnolia Press 39
COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L.
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).
LARRIDON ET AL.40 Phytotaxa 166 (1) © 2014 Magnolia Press
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).
Phytotaxa 166 (1) © 2014 Magnolia Press 41
COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L.
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.
LARRIDON ET AL.42 Phytotaxa 166 (1) © 2014 Magnolia Press
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. s c a e t t a e , 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 “Pycreusspecies 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.
Phytotaxa 166 (1) © 2014 Magnolia Press 43
COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L.
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).
LARRIDON ET AL.44 Phytotaxa 166 (1) © 2014 Magnolia Press
References
Aublet, J.B.C.F. (1775) Histoire des Plantes de la Guiane Françoise 1. Pierre-François Didot jeune, Paris.
http://dx.doi.org/10.5962/bhl.title.48831
Bauters, K., Larridon, I., Reynders, M., Huygh, W., Asselman, P., Vrijdaghs, A., Muasya, A.M. & Goetghebeur, P. (2014). A
new classification for Lipocarpha and Volkiella as infrageneric taxa of Cyperus (Cypereae, Cyperoideae, Cyperaceae):
insights from species tree reconstruction supplemented with morphological and floral developmental data. Phytotaxa
166(1): 1–32.
http://dx.doi.org/10.11646/phytotaxa.166.1.1
Beauvois, A.M.F.J., Palisot de (1816) Flore d'Oware et de Bénin, en Afrique 2 (15). Imprimerie de Fain, Paris.
Boeckeler, J.O. (1859) Beschreibungen neuer Cyperaceen, nebst berichtigenden Bemerkungen über einige ältere Cyperus-
Arten. Flora 42: 433–448.
Boeckeler, J.O. (1868) Die Cyperaceen des königlichen Herbariums zu Berlin 1. Die Cypereen, Scirpeen und Hypolytreen.
Linnaea 35 (5): 397–512.
Boeckeler, J.O. (1870) Die Cyperaceen des königlichen Herbariums zu Berlin 1. Die Cypereen, Scirpeen und Hypolytreen.
Linnaea 36 (3): 271–384.
Boeckeler, J.O. (1875) Ein Beitrag zur Kenntnis der Cyperaceen-Flora Neuholland's und einiger polynesischer Inseln. Flora 58
(7): 107112.
Boeckeler, J.O. (1888) Beiträge zur Kenntniss der Cyperaceen 1. Cyperaceae novae. Breitschädel & Vogt, Varel a.d. Jade.
Boeckeler, J.O. (1896) Diagnosen neuer Cyperaceen. Allgemeine Botanische Zeitschrift für Systematik, Floristik,
Pflanzengeographie etc. 1896: 53–56.
Brown R. (1810) Prodromus florae Novae Hollandiae et insulae Van-Diemen. Typis R. Taylor et socii, London.
http://dx.doi.org/10.5962/bhl.title.397
Brown, R. (1818) Observations, systematical and geographical, on professor Christian Smith's collection of plants from the
vicinity of the River Congo. In: Tuckey, J.H. (ed.), Narrative of an Expedition to Explore the River Zaire, usually called
the Congo in South Africa. Appendix 5: 420–485.
Chermezon, H. (1919) Kyllingia (Cypéracées) nouveaux de Madagascar. Bulletin du Muséum d'Histoire Naturelle (Paris) 25
(3): 208–212.
Chermezon, H. (1920) Cyperus nouveaux de Madagascar. Bulletin de la Société botanique de France 66: 338–353.
Chermezon, H. (1920 publ. 1921) Diagnoses de Pycreus et Cyperus nouveaux de Madagascar. Bulletin de la Société botanique
de France 67: 326–330.
Chermezon, H. (1921) Sur quelques Cypéracées nouvelles du Centre de Madagascar. Bulletin du Muséum d'Histoire Naturelle
(Paris) 27 (7): 552–554.
Chermezon, H. (1925) Diagnoses de Cypéracées nouvelles de Madagascar 2. Bulletin de la Société botanique de France 72:
613–618.
Chermezon, H. (1932) Sur quelques Cypéracées du Congo belge. Revue de Zoologie et de Botanique Africaines 22 (1): 63–71.
Chermezon, H. (1933) Cypéracées nouvelles du Gabon 2. Bulletin de la Société botanique de France 80: 506509.
Chermezon H. (1933 publ. 1934) Cypéracées nouvelles du Congo belge. Revue de Zoologie et de Botanique Africaines24(3) :
294–299.
Chermezon, H. (1936) Cypéracées nouvelles du Gabon 3. Bulletin de la Société botanique de France 83: 492496.
Clarke, C.B. (1894) Cyperaceae. In: Durand, T. & Schinz, H. (eds.), Conspectus Florae Africae 5. Jardin botanique de l’ Etat,
Brussels, pp. 526692.
Clarke, C.B. (1897) Cyperaceae. In: Thiselton-Dyer, W.T. (ed.), Flora capensis 7: 1–192. L. Reeve & Co., Ltd., Kent.
Clarke, C.B. (1900) Cyperaceae. In: De Wildeman E. & Durand T. (eds.), Contributions à la flore du Congo. 1. Annales du
Museé du Congo (Belge). Botanique. Séries 2, 1 (2): 65–71.
Clarke, C.B. (1901) Cyperaceae 1. In: Thiselton-Dyer WT. (ed.), Flora of Tropical Africa 8: 266384. Lovell Reeve & Co.,
London.
Clarke, C.B. (1902). Cyperaceae 2. In: Thiselton-Dyer WT. (ed.), Flora of Tropical Africa 8: 385–524. Lovell Reeve & Co.,
London.
Clarke, C.B. (1906) Beiträge zur Flora von Afrika 29. Cyperaceae africanae. Botanische Jahrbücher für Systematik,
Pflanzengeschichte und Pflanzengeographie 38 (2): 131–136.
Clarke, C.B. (1910) Cyperaceae. In: Hosseus, C.C. (ed.), Beiträge zur Flora Siams. Beihefte zum Botanischen Centralblatt 27
(2): 455–507.
Domin, K. (1915) Beiträge zur Flora und Pflanzengeographie Australiens. Bibliotheca Botanica 85: 414–498.
ttp://dx.doi.org/10.5962/bhl.title.58219
Duvigneaud, P.A. (1958) La végétation du Katanga et de ses sols métallifères. Bulletin de la Société Royale de Botanique de
Belgique 90: 127–286.
Druce, G.C. (1917) Nomenclatural notes: chiefly African and Australian. The Botanical Exchange Club and Society of The
British Isles 1916. Second Supplement: 602–655.
Eiten, L.T. (1963) Cyperaceae. In: Eiten, G. (ed.), Habitat flora of Fazenda Campininha, Sao Paulo, Brazil. Simposio sôbre o
Cerrado, Univ. Sao Paulo, pp. 207–208, 220–223.
Phytotaxa 166 (1) © 2014 Magnolia Press 45
COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L.
Endlicher, S.F.L. (1842) Catalogus Horti Academici Vindobonensis 1. Vienna, pp. 495.
Forster, J.R. & Forster J.G.A. (1775) Characteres Generum Plantarum:quas in itinere ad insulas maris Australis, collegerunt,
descripserunt, delinearunt, annis 17721775. London.
http://dx.doi.org/10.5962/bhl.title.4448
Goetghebeur, P. (1977) Studies in Cyperaceae. 1. Taxonomic notes on Ascolepis and Marisculus, a new genus of the tribe
Cypereae. Bulletin van de Nationale Plantentuin van België 47 (3–4): 435–447.
http://dx.doi.org/10.2307/3667910
Goetghebeur, P. (1980) Studies in Cyperaceae 2. Contribution towards a revision of the mainly African genus Ascolepis Nees
ex Steudel. Adansonia, ser. 2 (19): 269–305.
Goetghebeur, P. (1998) Cyperaceae. In: Kubitzki, F., Huber, H., Rudall, P.J., Stevens, P.S. & Stützel, T. (eds.), The families and
genera of vascular plants 4. Springer-Verlag, Berlin, pp. 141–190.
Govaerts, R., Simpson, D.A., Goetghebeur, P., Wilson, K.L., Egorova, T. & Bruhl, J. (2007) World Checklist of Cyperaceae.
Sedges. Kew Publishing, Royal Botanic Gardens, Kew, London, U.K., 780 pp.
Govaerts, R., Simpson, D.A., Goetghebeur, P., Wilson, K.L., Egorova, T. & Bruhl, J. (2013) World checklist of Selected Plant
Families. Cyperaceae. Kew: The Board of Trustees of the Royal Botanic Gardens, Kew. Available from: http://
www.kew.org/wcsp/monocots/ (accessed 7 July 2013).
Govindarajalu, E. (1975) Studies in Cyperaceae 13. Novelties in Cyperus L. subgen. Pycreus (Beauv.) C.B. Clarke.
Proceedings of the Indian Academy of Sciences. Plant Sciences 81B (5): 187–196.
Govindarajalu, E. (1991) Cyperaceae Indiae australis precursores: new species and combinations in Pycreus Beauv.
Proceedings of the Indian Academy of Sciences. Plant Sciences 100 (6): 423–433.
Haines, R.W. & Lye, K.A. (1978) Studies in African Cyperaceae 17. Kyllingiella R.W.Haines & Lye, gen. nov. Botaniska
Notiser 131: 175177.
Haines, R.W. & Lye, K.A. (1983) The sedges and rushes of East Africa. East African National History Society, Nairobi.
Hasskarl, J.K. (1848) Plantae Javanicae Rariores Adjectis Nonnullis Exoticis, in Javae Hortis Cultis Descriptae. Berlin.
http://dx.doi.org/10.5962/bhl.title.45111
Hoenselaar, K., Verdcourt, B. & Beentje, H.J. (2010) Cyperaceae. In: Beentje, H.J. (ed.), Flora of Tropical East Africa. Kew
Publishing, Royal Botanic Gardens, Kew, London, U.K., pp. 1–466.
Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (1990) Index Herbariorum. Part 1: The Herbaria of the World. New York
Botanical Garden, Bronx, NY.
Hooper, S.S. (1972) New taxa, names and combinations in Cyperaceae for the "Flora of West Tropical Africa". Kew Bulletin 26
(3): 577–583.
http://dx.doi.org/10.2307/4120322
Hooper, S.S. (1983) A new species of Ascolepis (Cyperaceae) from tropical Africa with notes on variation in A. pusilla Ridl.
Kew Bulletin 37: 605–608.
http://dx.doi.org/10.2307/4109731
Hutchinson, J. & Dalziel, J.M. (1936) Cyperaceae. Flora of West Tropical Africa. Edition 1, Volume 2, Part 2: 464–495.
Hô, P.H. (1993) Cayco Vietnam: An Illustrated Flora of Vietnam Volume 3. Mekong Printing, Santa Ana, California, 1276 pp.
Huygh, W., Larridon, I., Reynders, M., Muasya, A.M., Govaerts, R., Simpson, D.A. & Goetghebeur, P. (2010a) Nomenclature
and typification of names of genera and subdivisions of genera in Cypereae (Cyperaceae): 1. Names of genera in the
Cyperus clade. Tax o n 59 (6): 1883–1890.
Huygh, W., Schouppe, D., Larridon, I., Simpson, D.A. & Goetghebeur, P. (2010b) Kyllinga cataphyllata (Cyperaceae), a new
species from the highlands of West and Central Africa. Blumea 55 (3): 291–293.
http://dx.doi.org/10.3767/000651910x550936
IUCN (2012) IUCN Red List Categories and Criteria: Version 3.1. Second edition. Gland, Switzerland and Cambridge, UK:
IUCN. iv + 32pp.
Jacquin, N.J. (1770–71). Hortus Botanicus Vindobonensis 1. Leopoldi Joannis Kaliwoda, Vienna, 100 pp.
http://dx.doi.org/10.5962/bhl.title.531
Johnston, I.M. (1924) Expedition of the California Academy of Sciences to the Gulf of California. Botany. Proceedings of the
California Academy of Sciences 4 (12): 951–1218.
JSTOR Global Plants (2013). Available from: http://plants.jstor.org/ (accessed: 12 July 2013)
Kern, J.H. (1958a) Florae Malesianae praecursores 17. Notes on Malaysian and some SE Asian Cyperaceae 5. Blumea 9 (1):
215–236.
Kern, J.H. (1958b) Florae Malesianae precursores 21. Notes on Malayan and some SE Asian Cyperaceae 7. Acta Botanica
Neerlandica 7: 786800.
Kew Herbarium Catalogue (2013) Royal Botanic Gardens, Kew, Richmond, U.K. Available from: http://apps.kew.org/herbcat/
navigator.do/ (accessed: 7 August 2013).
Koyama, T. (1955) An enumeration of Hayata's Indo-Chinese collection of Cyperaceae: Carex and Cyperus. Naturaliste
Canadien 82 (1011): 194216.
Kükenthal, G. (1913) Cyperaceae novae. Repertorium Specierum Novarum Regni Vegetabilis 12: 9195.
http://dx.doi.org/10.1002/fedr.4870120404
Kükenthal, G. (1921) Cyperaceae. In: Fries, R.E. (ed.), Wissenschaftliche Ergebnisse der Schwedischen Rhodesia-Kongo-
LARRIDON ET AL.46 Phytotaxa 166 (1) © 2014 Magnolia Press
Expedition 19111912, I, Ergänzungsheft: 1–10, Tafel 1–3.
Kükenthal, G. (1925) Cyperaceae. In: Fries, R.E. & Fries, T.G.E. (eds.), Beiträge zur Kenntnis der Flora des Kenia, Mt Aberbare
und Mt Elgon 7. Notizblatt des Botanischen Gartens und Museums zu Berlin-Dahlem 9: 299–331.
http://dx.doi.org/10.2307/3994384
Kükenthal, G. (1934) Cyperaceae. In: Norlindh, T. & Weimarck, H. (eds.), Beiträge zur Kenntnis der Flora von Süd-Rhodesia 2.
Botaniska Notiser 1934: 6483.
Kükenthal, G. (193536) Cyperaceae – Scirpoideae – Cypereae. In: Engler A. (ed.), Das Pflanzenreich 4 (20) [Heft 101].
Engelmann, Berlin, pp. 1671.
Kükenthal, G. (1937) Cyperaceen. In: Peter, A. (ed.), Flora von Deutsch-Ostafrika. Repertorium Specierum Novarum Regni
Vegetabilis. Beihefte 40 (1): 381540
Kükenthal, G. (1943) Zur Kenntnis der Gattung Rhynchospora. Boissiera 7: 100104.
Kunth, C.S. (1837) Enumeratio Plantarum 2. Cyperographia synoptica. Sumtibus J.G. Cottae, Stuttgart & Tübingen.
Larridon, I., Reynders, M., Huygh, W., Bauters, K., Van de Putte, K., Muasya, A.M., Boeckx, P., Simpson, D.A., Vrijdaghs, A. &
Goetghebeur, P. (2011a) Affinities in C3 Cyperus lineages (Cyperaceae) revealed using molecular phylogenetic data and
carbon isotope analysis. Botanical Journal of the Linnean Society 167: 1946.
http://dx.doi.org/10.1111/j.1095-8339.2011.01160.x
Larridon, I., Reynders, M., Huygh, W., Bauters, K., Vrijdaghs, A., Leroux, O., Muasya, A.M., Simpson, D.A. & Goetghebeur, P.
(2011b) Taxonomic changes in C3 Cyperus (Cyperaceae) supported by molecular phylogenetic data, morphology,
embryology, ontogeny and anatomy. Plant Ecology and Evolution 144 (3): 327356.
http://dx.doi.org/10.5091/plecevo.2011.653
Larridon, I., Reynders, M., Huygh, W., Muasya, A.M., Govaerts, R., Simpson, D.A. & Goetghebeur, P. (2011c) Nomenclature
and typification of names of genera and subdivisions of genera in Cypereae (Cyperaceae): 2. Names of subdivisions of
Cyperus. Tax o n 60 (3): 868884.
Larridon, I., Bauters, K., Reynders, M., Huygh, W., Muasya, A.M., Simpson, D.A. & Goetghebeur, P. (2013) Towards a new
classification of the giant paraphyletic genus Cyperus (Cyperaceae): phylogenetic relationships and generic delimitation in
C4 Cyperus. Botanical Journal of the Linnean Society 172 (1): 106126.
http://dx.doi.org/10.1111/boj.12020
Larridon, I. & Goetghebeur, P. (2013). Cyperus steudneri, a new combination in Cyperus (Cyperaceae). Plant Ecology and
Evolution 146 (1): 138139.
http://dx.doi.org/10.5091/plecevo.2013.827
Link, J.H.F. (1820) Jahrbücher der Gewächskunde 1 (3), G.C. Nauck’s Buchhandlung, Berlin & Leipzig, 184 pp.
Linnaeus, C. (1753). Species Plantarum 1. Impensis Laurentii Salvii, Stockholm, 560 pp.
Linnaeus, C. (1762) Species Plantarum. Ed. 2 (1). Laurentii Salvii, Stockholm, 784 pp.
Lye, K.A. (1972a) Studies in African Cyperaceae 5. Sphaerocyperus Lye, gen. nov. Botaniska Notiser 125 (3): 212–216.
Lye, K.A. (1972b) Studies in African Cyperaceae 6. New species and combinations in Kyllinga. Botaniska Notiser 125 (3):
217–219.
Lye, K.A. (1981 publ. 1982) Studies in African Cyperaceae 21. New taxa and combinations in Kyllinga Rottb. Nordic Journal
of Botany 1 (6): 741–747.
http://dx.doi.org/10.1111/j.1756-1051.1981.tb01161.x
Lye, K.A. (1983a) Studies in African Cyperaceae 24. New taxa and combinations in Ascolepis and Isolepis. Nordic Journal of
Botany 2 (6): 561–566.
http://dx.doi.org/10.1111/j.1756-1051.1983.tb01050.x
Lye, K.A. (1983b). Studies in African Cyperaceae 25. New taxa and combinations in Cyperus L. Nordic Journal of Botany 3:
213–232.
http://dx.doi.org/10.1111/j.1756-1051.1983.tb01069.x
Lye, K.A. (1994) New combinations in African Cyperaceae. Lidia 3 (4): 131–132.
Lye K.A. & Mesterházy A. (2012) Studies in African Cyperaceae 35. Kyllinga carinalaevis sp. nov. from west tropical Africa.
Nordic Journal of Botany 30: 385–388.
http://dx.doi.org/10.1111/j.1756-1051.2012.01289.x
Meneses, O.J. Azancot de (1956) Ciperàceas de Angola existentes no herbàrio do Jardim e Museu Agricola do Ultramar
(LISJC). Garcia de Orta 4: 239–264.
Merxmüller, H. & Czech, G. (1953) Eine neue Gattung der Cyperaceae. Mitteilungen aus der Botanischen Staatssammlung
München I (8): 317–323.
Mtotomwema, K. (1990a) Two new yellow spiked Kyllinga species (Cyperaceae) from tropical Africa. Botanische Jahrbücher
für Systematik, Pflanzengeschichte und Pflanzengeographie 111 (3): 289–296.
Mtotomwema, K. (1990b) New species of Kyllinga (Cyperaceae) from tropical Africa. Nordic Journal Botany 9 (6): 637–641.
http://dx.doi.org/10.1111/j.1756-1051.1990.tb00554.x
Muasya, A.M., Simpson, D.A. & Chase, M.W. (2002) Phylogenetic relationships in Cyperus L. s.l. (Cyperaceae) inferred from
plastid DNA sequence data. Botanical Journal of the Linnean Society 138 (2): 145–153.
http://dx.doi.org/10.1046/j.1095-8339.2002.138002145.x
Muasya, A.M., Simpson, D.A., Verboom, G.A., Goetghebeur, P., Naczi, R.F.C., Chase, M.W. & Smets, E. (2009a) Phylogeny of
Phytotaxa 166 (1) © 2014 Magnolia Press 47
COMBINING SCOLEPIS, KYLLINGA AND PYCREUS INTO CYPERUS S.L.
Cyperaceae based on DNA sequence data: Current progress and future prospects. Botanical Review 75 (1): 2–21.
http://dx.doi.org/10.1007/s12229-008-9019-3
Muasya, A.M., Vrijdaghs, A., Simpson, D.A., Chase, M.W., Goetghebeur, P. & Smets, E. (2009b) What is a genus in Cypereae:
Phylogeny, character homology assessment and generic circumscription in Cypereae. Botanical Review 75 (1): 52–66.
http://dx.doi.org/10.1007/s12229-008-9018-4
Muasya, A.M., Musili, P.M. & Vrijdaghs, A. (2010) Kyllinga mbitheana (Cyperaceae)—Description, floral ontogeny and
pollen micromorphology of a new species from Kenya. Journal of East African Natural History 99 (1): 65–75.
http://dx.doi.org/10.2982/028.099.0105
Musée National d’Histoire Naturelle (2013). Herbarium catalogue, Musée National d’Histoire Naturelle, Paris, France. Available
from: http://coldb.mnhn.fr/ (accessed: 26 July 2013).
Napper, D. (1971) Cyperaceae of East Africa 5. Journal of East African Natural History 28 (124): 1–24.
Nees von Esenbeck, C.G. (1836) Cyperaceae Capenses Secundum Novissimas Ecklonii Collectiones. Linnaea 10: 129–207.
Nees von Esenbeck, C.G. (1842) Florae Brasiliensis Cyperographia. In: von Martius, C.F.P. (ed.), Flora Brasiliensis 2 (1). R.
Oldenbourg, München & Leipzig, pp. 1–226.
Nelmes, E. (1955) Notes on Cyperaceae. 36. New annual species of Pycreus from tropical Africa 2. Kew Bulletin 10 (1): 91–92.
http://dx.doi.org/10.2307/4113918
Palla, E. (1908) Cyperaceae. In: von Wettstein, R. & Schiffner, V. (eds) Ergebnisse der botanischen Expedition der kaiserlichen
Akademie der Wissenschaften nach Südbrasilien 1901. 1. Pteridophyta und Anthophyta 1. Denkschriften der
Mathematisch-Naturwissenschaftliche Klasse der Kaiserlichen Akademie in Wien 79: 168200.
Parlatore, F. (1846) Flora Palermitana ossia descrizione delle piante che crescono spontanee nella valle di Palermo. Famiglia
2A. Cyperaceae. Giornale Botanico Italiano 2 (1): 117-173.
Pax, F. (1887) Cyperaceae. In: Engler, A. & Prantl, K. (eds.) Naturlichen Pflanzenfamilien 1, 2, 2: 98126. W. Engelmann,
Leipzig.
Pfeiffer, H. (1935) Conspectus Cyperacearum in America meridionale nascentium 6. Bemerkungen zum Gattungswert von
Dichromena Michx. und Pleurostachys Brongn. Repertorium Specierum Novarum Regni Vegetabilis 38: 8899.
http://dx.doi.org/10.1002/fedr.19210171312
Podlech, D. 1960. Über einige Cyperaceen Südafrikas. Mitteilungen der Botanischen Staatssammlung, München 3: 521–530.
Prasad, V.P. 2009. Five new synonyms of Pycreus malabaricus (Cyperaceae). Rheedea 19 (12): 1924.
Raymond, M. (1957) Some new or critical Scirpus from Indo-China. Naturaliste Canadien 84: 111–150.
Raynal, J. (1968) Notes cypérologiques 11. Sur quelques Scirpus et Ascolepis de l'Ancien Monde. Adansonia, sér. 2, 8 (1): 85–
104.
Raynal, J. (1973) Notes cypérologiques 19. Contribution à la classification de la sous-famille des Cyperoideae. Adansonia, sér.
2, 13 (2): 145–171.
Raynal, J. (1977) Notes cypérologiques 31. Mélanges nomenclaturaux (Cyperoideae) Adansonia, sér. 2, 17: 4347.
Retzius, A.J. (1788) Observationes botanicae 5. Siegfried Lebrecht Crusium, Leipzig.
http://dx.doi.org/10.5962/bhl.title.11760
Reynders, M. (2013) The challenging taxonomy and evolution of C4 Cyperus (Cyperaceae). A focus on Pycreus and its laterally
compressed pistils. Ph.D. thesis, Ghent University, Belgium: 382 p.
Reynders, M. & Goetghebeur, P. (2010). Reestablishment of Pycreus sect. Tuberculati Cherm. Blumea 55 (3): 226–230.
http://dx.doi.org/10.3767/000651910x539993
Reynders, M., Huygh, W., Larridon, I., Muasya, A.M., Govaerts, R., Simpson, D.A. & Goetghebeur, P. (2011) Nomenclature
and typification of names of genera and subdivisions of genera in the Cypereae (Cyperaceae): 3. Names in segregate
genera of Cyperus. Tax on 60 (3): 885–895.
Reynders, M., Vrijdaghs, A., Muasya, A.M., Larridon, I., Goetghebeur, P. & Smets E. (2012) Gynoecial anatomy and
development in Cyperoideae (Cyperaceae, Poales): congenital fusion of carpels facilitates evolutionary modifications in
pistil structure. Plant ecology and evolution 145 (1): 96–125.
http://dx.doi.org/10.5091/plecevo.2012.675
Richard, A. (1850–51) Tentamen florae abyssinicae 2. Bertrand, Paris, 518 pp.
Ridley, H.N. (1884) The Cyperaceae of the west coast of Africa in the Welwitsch herbarium. Transactions of the Linnean
Society of London. Botany 2: 121–172.
http://dx.doi.org/10.1111/j.1095-8339.1884.tb00007.x
Rottbøll, C.F. (1773) Descriptionum et Iconum Rariores et pro maxima parte novas plantas illustrantium liber primus.
Copenhagen.
http://dx.doi.org/10.5962/bhl.title.9538
Roxburgh, W. (1820) Flora Indica 1. Mission press, Serampore.
Samain, M.-S., Reynders, M. & Goetghebeur, P. (2006) Kyllinga beninensis (Cyperaceae), a new species from Bénin. Novon
16: 516–519.
http://dx.doi.org/10.3417/1055-3177(2006)16[516:kbcans]2.0.co;2
Schumacher, H.C.F. (1827) Beskrivelse af Guineeiske Planter. Hartv. Frid. Popps Bogtrykkerie, Copenhagen, 466 pp.
Schumann, K.M. (1900) Neue Arten der Siphonogamen 1898. Just's Botanischer Jahresbericht 26 (1): 323–396.
Simpson, D.A., Muasya, A.M., Alves, M., Bruhl, J.J., Dhooge, S., Chase, M.W., Furness, C.A., Ghamkhar, K., Goetghebeur, P.,
LARRIDON ET AL.48 Phytotaxa 166 (1) © 2014 Magnolia Press
Hodkinson, T.R., Marchant, A.D., Nieuborg, R., Reznicek, A.A., Roalson, E.H., Smets, E., Starr, J.R., Thomas, W.W.,
Wilson, K.L. & Zhang, X. (2007, publ. 2008) Phylogeny of Cyperaceae based on DNA sequence data—a new rbcL
analysis. In: Monocots III/Grasses IV. Aliso 23: 72–83. Rancho Santa Ana Botanic Garden. Claremont, CA.
Soják, J. (1979 publ. 1980) Fragmenta phytotaxonomica et nomenclatorica 2. Casopsis Narodniho Musea – rada
prirodnovedna 148: 193–197.
Steudel, E.G. von (1829) Einige Bemerkungen über Kap'sche Cyperaceen und Gramineen 1 (2). Flora 12 (10): 145–154.
Steudel, E.G. von (1842) Ueber die Arten von Cyperus, Mariscus und Kyllingia, welche in der zweiten Sendung von Pflanzen
aus Abyssinien von dem Reisenden des Vereins Hrn. W. Schimper enthalten sind (Schluss). Flora 25, 2 (38): 593–599.
Steudel, EG. von (1854–55) Synopsis plantarum glumacearum 2. Cyperaceae. J.B. Metzler, Stuttgart.
http://dx.doi.org/10.5962/bhl.title.471
Stokes, J. (1812) A botanical materia medica 1. J. Johnson and Co., London, 503 pp.
Symoens, J.J. (1953) Travaux du Laboratoire de botanique systématique et de phytogéographie de l'Université libre de
Bruxelles nº 6: Note sur la végétation des salines de Mwashya (Katanga). Laboratoire de botanique systématique,
Bruxelles, 121 pp.
Thunberg, C.P. (1825) Florula Javanica. Palmblad et C., Uppsala, 23 pp.
Tropicos.org (2013). Missouri Botanical Garden, Saint Louis, U.S.A. Available from: http://www.tropicos.org/ (accessed: 20
August 2013).
Turrill, W.B. (1915) Ascolepis gracilis Turrill. Hooker's Icones Plantarum 31: t. 3020.
Vahl, M. (1805) Enumeratio plantarum 2. Copenhagen, Impenis auctoris, & prostat apud J.H. Schubothe.
Van der Veken, P. (1955) Trois taxa nouveaux de Cypéracées du Congo belge. Bulletin du Jardin Botanique de l'État à
Bruxelles 25 (1): 143–147.
http://dx.doi.org/10.2307/3667104
Vrijdaghs, A., Reynders, M., Muasya, A.M., Larridon, I., Goetghebeur, P. & Smets, E. (2011) Spikelet and Floral Morphology
and Development in Cyperus and Pycreus (Cyperaceae). Plant Ecology and Evolution 144 (1): 44–63.
http://dx.doi.org/10.5091/plecevo.2011.436
Welwitsch, F.M.J. (1859) Apontamentos Phytogeographicos. pp. 527–593.
Welwitsch, F.M.J. (1869) Sertum Angolense. Transactions of the Linnean Society of London 27: 1–97.
http://dx.doi.org/10.1111/j.1096-3642.1869.tb00202.x
Wood, A. (1861) Class book of botany: being outlines of the structure, physiology and classification of plants; with a flora of
all parts of the United States and Canada. Troy: A.S. Barnes & Burr, Moore & Nims, New York, 832 pp.
http://dx.doi.org/10.5962/bhl.title.32648
Zollinger, H. (1854) Systematisches Verzeichniss der im indischen Archipel in den Jahren 18421848 gesammelten sowie der
aus Japan empfangenen Pflanzen. Zürich, 63 pp.
http://dx.doi.org/10.5962/bhl.title.80000
... Species belonging to the Cyperaceae family represent an important source of active constituents with therapeutic properties (Gamal, Hani, Sabrin, 2015). In taxonomic terms, Cypereae is the second most diverse tribe and its largest genus, Cyperus (Linnaeus 1753:44) includes about 600 species, some of which are used in popular medicine (Larridon et al., 2014;Reid, Carter, Urbatsch, 2014). The activities antioxidant and antimicrobial of some species of this genus have been widely studied and scientifically proven (Soumaya et al., 2014;Essaidi et al., 2014;Nassar et al., 2015;Jing et al., 2016). ...
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This study was conducted to investigate the complete sequences of chloroplast maturase K (matK) gene, the study were determined for five species of Cyperus L. Cyperaceae in Iraq to infer phylogenetic relationships. Dipsacus laciniatus were used as out groups.. The phylogenetic tree based on Maximum Parsimony (MP) included three major clades the first major clade included C.alternifolius in the base of trees which is a sister clade to C.iria , was the third major clade gathered the C.aucheri, C.rotandus, and C.longus respectively with C.iria. These clades are subdivided into two secondary clades, C.aucheri descendent in single clade alone with the C.rotandus and C.longus. the clades of Bayesian analysis are as follow: First Major clade includes consist only C.alternifolius with the Second Major clade C.aucheri and the last Major clades subdivided into two subclades the first on descending C.iria alone as a sister clade with C.longus and C.rotandus subclade .
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The morphological diversity and the presence of several convergent evolutionary lineages in the Cypereae tribe (Cyperaceae) have given rise to various conflicting classifications. These conflicts do not only arise in the delimitation of genera and their subdivisions, but also in the use of similar subdivisional names for different species groups and different names for similar species groups. This has resulted in the publication of approximately 350 names of genera and subdivi sions of genera to accommodate the ca. 950 species in Cyperus and its segregate genera. This complex nomenclature has led to an accumulation of errors in the assessment of valid publication, priority and legitimacy, and in typification in almost all existing taxonomic treatments for the group. Renewed interest in the phylogeny and taxonomy of Cypereae reveals the need to evaluate the nomenclature of generic names and names of subdivisions of genera. In this paper, the first in a series of four papers, we present a survey of all generic names described in the Cyperus clade, designate types where needed, and evaluate priority and legitimacy.
Thesis
Cypereae form one of the largest and most complex tribes of the sedge family (Cyperaceae). Recently, two clades have been revealed within the tribe, the largest of which includes the giant genus Cyperus and its closest allies. However, thirteen genera of the generally accepted classification of Goetghebeur (1998) appear to be nested within Cyperus. The taxonomic status of many of these taxa has been under discussion since they are based on different combinations of a limited set of derived characters. Pycreus, the largest of these segregate genera, is characterised by laterally compressed dimerous pistils of which the derivation from the general trimerous situation was not yet understood. It shares this pistil with Kyllinga and Queenslandiella that both are, as is Pycreus, embedded in the Cyperus clade which uses C4 photosynthesis. The recent insights from molecular phylogenetics make a reevaluation possible of the taxonomic status of the thirteen different segregate genera of Cyperus and of the taxonomic value of the characteristics that have been used to delimitate these taxa. This is currently tackled in a joint international research effort, using a combination of molecular phylogenetics, ontogeny, anatomy and morphology, to understand evolutionary patterns in Cyperaceae and to build a modern classification of sedges. This research strategy is situated on three taxonomic levels: family to tribal level (macro-scale), tribal to generic level (meso-scale) and infrageneric level (micro-scale). The current thesis is embedded in this international research context and focusses mainly on meso-scale objectives (C4 Cyperus and the position and taxonomic state of its segregate genera, including Pycreus) and micro-scale objectives (the infrageneric taxonomy of Pycreus). At first, a complete nomenclatural survey is presented of all generic and subdivisional names that have been published for the taxa now included in the Cyperus clade (around 350 names), along with an evaluation of their validity, legitimacy and priority. Types are indicated and where necessary lectotypes are designated. This nomenclatural survey serves as a base for the selection of representative taxa in the molecular, ontogenetic, anatomical and morphological studies. In addition it forms an essential tool when building a modern revision for the clade. In the current thesis only names for taxa in which Pycreus species have been placed are included. Next, to be able to reevaluate the taxonomical value of derived pistils in the Cyperus clade, especially the laterally compressed dimerous pistils of Pycreus, an elaborate ontogenetic study of Pycreus and Cyperus species was performed. This study shows that both taxa follow the general ontogenetic patterns of spikelets and flowers found throughout Cyperoideae. In addition, the ontogeny and anatomy of the different types of pistils was reviewed with addition of new ontogenetic and anatomical data. These demonstrate that in Cyperoideae the pistil wall starts from an annular primordium (which evolved from congenitally fused carpels) on top of which the stigma primordia develop. The development of the central ovule is decoupled from the ovary wall development. Vascular patterns follow the development of the primordia and vascular bundles are formed where necessary. The presence of an annular gynoecial wall primordium appears to have opened new possibilities for the development of the stigma primordia in new positions independent from the constraints of individual carpels. An elaborate molecular phylogenetic study was performed on the C4 Cyperus clade using ETS1f, rpl32-trnL, trnH-psbA. Although relationships within the C4 Cyperus clade are still largely unresolved in a large polytomy, early emerging branches show better resolution than in previous studies. Pycreus appears to be para- or polyphyletic and in addition no relationships have been found between Pycreus, Kyllinga and Queenslandiella. Therefore, we have to admit, laterally compressed dimerous pistils have most likely originated multiple times in the clade. Subsequently, the most appropriate classification strategy for these taxa is sinking them into Cyperus. This also seems to be the most appropriate strategy for all other segregate genera based on a reevaluation of the taxonomical value of their key characters. Only for the C4 Cyperus clade (accommodated in C. subgenus Cyperus), which is nested within a grade of species using C3 photosynthesis (accommodated in C. subgenus Anosporum), an evolutionary classification strategy has been adopted. This is based on the evolutionary value of the origin of C4 photosynthesis which had led to a major radiation of species. On the micro-scale, it is not yet possible to present a modern classification for Pycreus since molecular phylogenetic relationships are largely unresolved. Therefore, results are presented as several case studies. First, in an elaborate SEM study, the taxonomical value of the nutlet epidermis was reevaluated. Next, the reestablishment of P. sect. Tuberculati is discussed. Finally, the new classification strategy for the Cyperus clade was applied on Pycreus and necessary combinations and nomina nova under Cyperus are listed along with some critical notes on synonymisations of several taxa.