Lebrun, J.-P. & A.L. Stork (2019). New combinations in Cyperaceae for continental Africa. In English, English abstract. Candollea 74: 145–151. DOI: http://dx.doi.org/10.15553/c2019v742a4
The generic concept adopted in our forthcoming volume of the family Cyperaceae for the Tropical African Flowering Plants series requires some new combinations in Cyperaceae. Our compilation of Cyperaceae is a compromise between traditional and the most recent treatments as regards Cyperus. We therefore propose 18 new combinations and a replacement name for taxa first described in Cyperus L. transferred to the genera Kyllinga Vahl, Mariscus Rottb. and Pycreus P. Beauv.: Kyllinga brunneofibrosa (Lye) J.-P. Lebrun & Stork, Kyllinga inselbergensis (Lye) J.-P. Lebrun & Stork, Kyllinga microcristata (Lye) J.-P. Lebrun & Stork, Kyllinga rheophytica (Lye) J.-P. Lebrun & Stork, Mariscus absconditicoronatus (Bauters, Reynders & Goetgh.) J.-P. Lebrun & Stork, Mariscus baobab (Lye) J.-P. Lebrun & Stork, Mariscus baoulensis (Kük.) Hutch. ex J.-P. Lebrun & Stork, Mariscus boreochrysocephalus (Lye) J.-P. Lebrun & Stork, Mariscus cundudoensis (Chiov.) J.-P. Lebrun & Stork, Mariscus gypsophilus (Lye) J.-P. Lebrun & Stork, Mariscus kitaleensis J.-P. Lebrun & Stork, Mariscus micromedusaeus (Lye) J.-P. Lebrun & Stork, Mariscus ossicaulis (Lye) J.-P. Lebrun & Stork, Mariscus pluricephalus (Lye) J.-P. Lebrun & Stork, Mariscus recurvispicatus (Lye) J.-P. Lebrun & Stork, Mariscus somalidunensis (Lye) J.-P. Lebrun & Stork, Mariscus soyauxii subsp. pallescens (Lye) J.-P. Lebrun & Stork, Mariscus unispicatus (Bauters, Reynders & Goetgh.) J.-P. Lebrun & Stork, and Pycreus micropelophilus (Lye) J.-P. Lebrun & Stork.
Received: June 24, 2019. Accepted: July 22, 2019. First published online: August 19, 2019.
Introduction
Cyperus L. (Cyperaceae) is a large genus of c. 950 species (s.l.) or 550 (s.str.) distributed in temperate and tropical regions worldwide. Until recently the circumscription of the genus was problematic. Early molecular studies resolved a well-supported clade that included Cyperus s.str., but also inferred a paraphyletic genus containing about 13 segregated genera, thus rendering Cyperus monophyletic. But most of the Cyperus species have not been included in phylogenetic studies. “Due to the massive size of the genus, a comprehensive phylogeny of Cyperus is not imminent” (Reid et al., 2017: 434–344).
The generic concept adopted in our forthcoming volume of the family Cyperaceae for the Tropical African Flowering Plants series (Lebrun & Stork, in press) is the same as that used in several traditional floras, and in our Énumération (Lebrun & Stork, 1995). In its main lines it corresponds to the classification proposed by Goetghebeur (1998). This system differs from the one adopted by, for instance, the World Checklist of Selected Plant Families (WCSP, 2019). The differences of opinion mainly concern the genus Cyperus where Goetghebeur recognizes several segregate genera. These plants “are difficult to classify due to the complex structure of their inflorescences, which leads to different interpretations and to establishing uncertain hypotheses of homology” (Reutemann et al., 2012: 184).
Our compilation of Cyperaceae is a compromise between traditional and the most recent treatments as regards Cyperus. We think that a more narrowly defined genus is preferable to a broad generic concept with subgeneric divisions. These segregate genera are often morphologically distinct and the species easily recognized in the field. Examples can be found in the genera such as Courtoisina Soják with deciduous intact spikelets, Kyllingiella R.W. Haines & Lye with spirally arranged glumes, and Oxycaryum Nees with spirally arranged glumes and dorsiventrally flattened dimerous gynoecia.
There is also the question of a “very complex generic and subdivisional nomenclature with approximately 350 generic and subdivisional names to accommodate the roughly 950 species present in the group” (Larridon et al., 2011: 868). When BODARD (1952) described a new species of Cyperus as “C. (Mariscus) plurinervosus” from Chad he followed the trend of his time, i.e. to merge Mariscus Vahl into Cyperus, and he noticed on the large number of new combinations that would be necessary but not called for this taxonomic concept.
It is true that the distinction between Cyperus and Mariscus is sometimes doubtful, e.g. in Cyperus distans L. f. Certain specimens have some spikelets falling off as intact units (Mariscus character), whereas other spikelets on the same plant have glumes breaking off from a persistent rachilla (Cyperus character) (Haines & Lye, 1983).
Mariscus was included in Cyperus by Goetghebeur (1998). He wrote: “Mariscus is kept separate by several authors, is rarely maintained at subgeneric level when included in Cyperus, and is often divided into sections of widely scattered affinities” (Goetghebeur, 1998: 170). For the latter author the polyphy-letic nature of Mariscus has convincingly been demonstrated by Lye (1992) and is recognized as a separate genus by some authors (e.g. RAYNAL, 1973).
A historical review of the treatment of Mariscus is given by REYNDERS et al. (2011), with the names of subdivisions cited. In older floras, such as Flora of West Tropical Africa (Hutchinson & Dalziel, 1972), Mariscus is maintained as separate. However, the recent Flora of Tropical East Africa on Cyperaceae (Beentje, 2010) includes Mariscus in Cyperus, although other segregate genera are maintained. On writing up our compilation we checked more recent treatments. It seems that authors of local floras or checklists keep Mariscus as a distinct genus (e.g. Lisowski, 2009; Malaisse, 2010; Chatelain et al., 2011; Thiombiano et al., 2012). Even Browning & Goetghebeur (2017: 61) present Mariscus as a separate entity.
We can also cite Gordon-Gray (1995: 125) who made a pragmatic decision for Cyperaceae in Natal. This latter author wrote: “From anatomy, physiology, karyology and phytochem-istry, information is steadily accumulating that Mariscus species are more naturally positioned within Cyperus and Pycnostachys than collectively in a taxon Mariscus at either generic or subgeneric rank […] Nevertheless, in the present work Mariscus is maintained as Natal species are well known under that genus and Cyperus is already cumbersome with the greatest number of species in Natal for Cyperaceae as a whole”.
Lowe & Stanfield (1974: 93, 95) described Mariscus as “a large and difficult genus [c. 200 species]; in characters of habit and inflorescence the species are variable, and no clear feature links them. They resemble Cyperus in having 3 stigmas, and Kyllinga in that the spikelets fall entire when mature. […] Some species are much like Torulinium in that the rhachis of the inflorescence bears persistent scales after the spikelets have fallen.”
Browning & Goetghebeur (2017) give a summary of characters previously used to differentiate Mariscus from Cyperus: 1. spikelets disarticulating as a unit; 2. leaf blades usually well developed; 3. less than 5 nutlets per spikelet; 4. winged rachilla.
Our main list (Lebrun & Stork, in press) comprises 65 species of Mariscus, and 9 species of Cyperus are added as probably belonging to that genus.
For Muasya et al. (2010: 65–66) the diagnostic characters for Kyllinga include “capitate inflorescences; spikelets with distichous glume arrangement which are shed intact; bifid style and lenticular nutlets; and laterally flattened nutlets”. For these authors, Kyllinga is either recognized as a distinct genus or ranked as a subgenus of Cyperus.
Although Kyllinga is now often treated under Cyperus (e.g., WCSP, 2019), some authors maintain Kyllinga as a separate genus. This is the case in our treatment, and we follow Govaerts & Simpson (2007), Goetghebeur (1998), Flora of China (WU et al., 2010), Beentje (2010), and Browning & Goetghebeur (2017). We therefore propose four new combinations in Kyllinga. This in spite of Lye & Cheek (2006: 276) arguing that “[a]lthough the genus Kyllinga Rottb. was incorporated in Cyperus more than 100 years ago […] it was not until 100 years later […] that it was proven beyond doubt that this is actually correct”.
Pycreus is a rather large pantropical genus (c. 120 species), so also in tropical Africa (63 species). Recent molecular phylogenetic studies indicate that it is included in Cyperus s.l., and also in culm and leaf anatomy Pycreus species have a high resemblance to the Cyperus C4 taxa. However, “Pycreus is characterized by a combination of bicarpellate ovaries, bifid styles, laterally flattened achenes, and multi-flowered spikelets” (Pereira-Silva et al., 2018: 741). It can be added that the achenes are arranged in a single row down the two opposite sides, like small discs set on edge, the rachilla is 4-sided (in Cyperus flattened). The achenes and their bracts are shed gradually from the bottom up, exposing the scarred rachilla (Reynders et al., 2011).
Pycreus is recognised as a segregate genus by several recent authors and floras, e.g. Goetghebeur (1998), Govaerts & Simpson (2007), Beentje (2010), Flora de Guinea Ecuatorial (Velayos et al., 2014), and Browning & Goetghebeur (2017). We follow this genus concept.
Taxonomy
Kyllinga brunneofibrosa (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus brunneofibrosus Lye in Candollea 51: 423. 1996.
Holotypus: Somalia. Reg. Bakool: 17 km E of Wojid (Uegit) on road to Oddur, 22.V.1983, Gillett & Hemming 24356 (K [K000321403] image seen).
Notes. – Cyperus brunneofibrosus was first introduced by Lye in Thulin (1995) but lacking a Latin diagnosis. LYE (1996a) validated the name in his revision of Cyperus subg. Kyllinga (Roth) J.V. Suringar in Somalia.
The species is illustrated in Thulin (1995: 141) and LYE (1996a: 427). A distribution map is also provided by LYE (1996a: 431).
Kyllinga inselbergensis (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus inselbergensis Lye in Nord. J. Bot. 31: 574. 2013.
Holotypus: Cameroon: Ako-akas Rock, 23 km on the road from N’Koemvone to Ambam (old road), 29.IX.1974, de Wilde 7771 (WAG; iso-: BR, K).
Notes. – This taxon is mentioned under Cyperus in Cameroon (ONANA, 2013), in Equatorial Guinea (VELAYOS, 2014: 116–117) and Gabon (LYE & THERY, 2012: 84, who also provide a photograph of the nutlet on p. 85).
Kyllinga microcristata (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus microcristatus Lye in Nord. J. Bot. 24: 269. 2005.
Holotypus: Cameroon. Reg. Sud Ouest: Kupe village, 11.VII.1995, Patterson 11 (K; iso-: BR, MO, NAG, NLH, YA).
Notes. – Cyperus microcristatus was first introduced by Lye in Cheek et al. (2004: 190) but lacking a Latin diagnosis, and then validated by Lye & Pollard (2005: 269) with iconography. Other citations figure under Cyperus in Onana (2011: 365, 2013: 223).
Kyllinga rheophytica (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus rheophyticus Lye in Nord. J. Bot. 24: 273. 2006.
Holotypus: Cameroon. Reg. Sud Ouest: Kupe-Muanenguba, Muambong, bank of River Chide, 8.II.1998, Onana 585 (K; iso-: O, YA).
Notes. – Cyperus rheophyticus was first introduced by Lye under the name “Cyperus rheophytorum” in Cheek et al. (2004: 190) but lacking a Latin diagnosis, and then validated as C. rheophyticus by Lye & Cheek (2006). It is mentioned under Cyperus in Onana (2011: 161; 2013) and Onana & Cheek (2011: 365).
A map is provided by Kuetegue et al. (2019: 88) and an illustration by Lye & Cheek (2006: 274–275).
Mariscus absconditicoronatus (Bauters, Reynders & Goetgh.)
J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus absconditicoronatus Bauters, Reynders & Goetgh. in Novon 20: 133. 2010.
Holotypus: Angola. Prov. Bié: falls of Cutato River, S of [Kuvango], Cuchi Rd., 15.IX.1952, H. & E. Hess 52/262 (GENT image seen).
Note. – An illustration is available in Bauters et al. (2010: 134).
Mariscus baobab (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus baobab Lye in Nord. J. Bot. 16: 371. 1996.
Holotypus: Somalia: Shabeelaha Dhexe, 1.VIII.1959, Moggi & Bavazzano 344 (FT [FT000610] image seen).
Note. – Iconography is available in Thulin (1995: 131) and Lye (1996b: 368–372).
Mariscus baoulensis (Kük.) Hutch. ex J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus baoulensis Kük. in Engl., Pflanzenr. 38: 467. 1936.
Holotypus: Ivory Coast: Cercle de Baoulé Nord, entre Tiégouakro et Kodiokoffi, 8.VIII.1909, Chevalier 22336 (P [P00568889] image seen).
– Mariscus baoulensis Hutch. in Hutchinson & Dalziel, Fl. W. Trop. Afr. 2: 486. 1936 [nom. inval.]. = Cyperus baoulensis (A. Chev.) Kük. in Repert. Spec. Nov. Regni Veg. 29: 199. 1931 [nom. nud.]. = Pycreus baoulensis A. Chev., Explor. Bot. Afrique Occ. Franç. 1: 695. 1920 [nom. nud.].
Note. – The taxon is mentioned under Cyperus as such by Hooper in Hutchinson & Dalziel (1972: 287), Brunel et al. (1984: 537) and Chatelain et al. (2011: 220).
Mariscus boreochrysocephalus (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus boreochrysocephalus Lye in Nord. J. Bot. 3: 216. 1983.
Holotypus: Uganda. Distr. Karamoja: 5–6 km N of Lothaa, 1180 m, 10.IV.1970, Lye 5462 (MTTU; iso-: C, EA [EA000002703] image seen, K, P, UPS).
Notes. – Cyperus boreochrysocephalus is illustrated in LYE (1983: 216) and Haines & Lye (1983: 219).
This taxon is mentioned under Cyperus by Beentje (2010: 149) and Darbyshire et al. (2015: 104).
Mariscus cundudoensis (Chiov.) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus cundudoensis Chiov. in Malpighia 35: 65. 1939.
Holotypus: ETHIOPIA: Harar, cima del Gara Cun-duduo, 2800 m, 5.XII.1937, Gortani & Jaboli n. I. 18 (FT [FT000630] image seen).
Notes. – The spelling of the species epithet varies. It was described by Chiovenda in Malpighia (vol. 35: 65. 1939) as C. cundudoensis. This issue of Malpighia is present in the K library, and we have seen a copy of the species description. However, the usual citation is C. cunduduensis, following Index Kewensis (Suppl. X (1936–1940: 68, 1947) with reference to Atti della Reale Accademia d’Italia. Memorie delle classe di scienze fisiche, matematiche e naturali (vol. 11: 60, 1940). A copy of this article, Plantae novae aut minus notae ex Aethiopia (pp. 17–67), is present at the G Library, dated on cover page “1940-XVIII”, but at the end of the article “Roma, 1941-XIX”. The article arrived at the Academy on 22 August 1939, and was presented on 18 November 1939.
An illustration is available in Thulin (1995: 135) and the taxon is cited under Cyperus in EDWARDS et al. (1997: 465).
Mariscus gypsophilus (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus gypsophilus Lye in Nord. J. Bot. 16: 374. 1996.
Holotypus: Somalia. Reg. Nugal: 3 km E of Anod, 30.VI.1979, Hansen & Heemstra 6323 (K [K000321405] image seen; iso-: C, EA, WAG).
Notes. – Cyperus gypsophilus was first introduced by Lye in Thulin (1995: 137) but lacking a Latin diagnosis and validated the following year by LYE (1996b).
The taxon is illustrated in Thulin (1995: 137) and cited under Cyperus by Davies (1998: 15) and Lye (2001: 204).
Mariscus kitaleensis J.-P. Lebrun & Stork, nom. nov.
≡ Cyperus kyllingiformis Lye in Nord. J. Bot. 3: 218. 1983.
Holotypus: Kenya: Kitale, 1400 m, 12.V.1953, Bogdan 3726 (K; iso-: EA).
Notes. – The name Mariscus kyllingiiformis Boeckeler (in Flora 42: 443. 1859) was given to a plant the name of which is now considered a synonym of Cyperus dubius Rottb. The compound of the species epithet is based on an orthographic variant of Kyllinga Rottb. 1773, nom. cons. The replacement name refers to the collecting site of the type, i.e. Kitale (Kenya).
The taxon is illustrated in Lye (1983: 218) and Haines & Lye (1983: 224) and cited under Cyperus by Lye (2001) and Beentje (2010: 185).
Mariscus micromedusaeus (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus micromedusaeus Lye in Nord. J. Bot. 16: 373. 1996.
Holotypus: Somalia. Reg. Nugal: gorge of Wadi Nogal, 5 km from Eil, 4.I.1973, Bally & Melville 15547 (K [K000321406] image seen).
Notes. – Cyperus micromedusaeus was first introduced by Lye in Thulin (1995: 134) but lacking a Latin diagnosis and validated the following year by Lye (1996b). The taxon is cited under Cyperus by Lye (2001: 204).
Mariscus ossicaulis (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus ossicaulis Lye in Kew Bull. 51: 205. 1996.
Holotypus: Somalia: 20 km W of Xarardheere, 10.VI.1979, Beckett 202 (K [K000321408] image seen).
Notes. – Cyperus ossicaulis was first introduced by Lye in Thulin (1995: 135) but lacking a Latin diagnosis and validated and illustrated the following year by LYE (1996c: 206–208). The taxon is cited under Cyperus by LYE (2001: 204).
Mariscus pluricephalus (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus pluricephalus Lye in Nord. J. Bot. 16: 133. 1996.
Holotypus: Somalia. Reg. Hiran: Jalalaqsi distr., ± 12 km NE of Ceel Baraf, 14.XII.1987, Kuchar 17635 (UPS).
Notes. – Cyperus pluricephalus was first introduced by Lye in Thulin (1995: 131) with an illustration but lacking a Latin diagnosis and validated the following year by Lye (1996b).The taxon is cited under Cyperus by Lye (2001: 204).
Mariscus recurvispicatus (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus recurvispicatus Lye in Nord. J. Bot. 16: 376. 1996.
Holotypus: Somalia. Reg. Mudug: 28 km S of Jeriban, 27.V.1979, Gillett et al. 22100 (K [K000321409] image seen).
Notes. – Cyperus recurvispicatus was first introduced by Lye in Thulin (1995: 137) with an illustration but lacking a Latin diagnosis and validated the following year by Lye (1996b). The taxon is cited under Cyperus by Lye (2001: 204).
Mariscus somalidunensis (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus somalidunensis Lye in Nord. J. Bot. 16: 374. 1996.
Holotypus: Somalia: Shabeelaha Dhexe, 1.VIII.1959, Moggi & Bavazzano 377 (FT [FT000653] image seen).
Notes. – Cyperus somalidunensis was first introduced by Lye in Thulin (1995: 137) with an illustration but lacking a Latin diagnosis and validated the following year by Lye (1996b). The taxon is cited under Cyperus by Lye (2001: 204).
Mariscus soyauxii subsp. pallescens (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus soyauxii subsp. pallescens Lye in Nord. J. Bot. 3: 227. 1983.
Holotypus: Kenya: Garissa, 300 m, 14.XII.1977, Stannard & Gilbert 1061 (EA; iso-: K [K000321463] image seen).
Notes. – Mariscus soyauxii (Boeckeler) C.B. Clarke is mentioned by Simpson & Inglis (2001: 308), Beentje (2010: 217) and Chatelain et al. (2011: 226) under Cyperus.
The taxon is illustrated by Haines & Lye (1983: 207–208), Berhaut (1988: 278), Hedberg et al. (2009: 265) and Lye & Thery (2012: 21).
Mariscus unispicatus (Bauters, Reynders & Goetgh.) J.-P.
Lebrun & Stork, comb. nova.
≡ Cyperus unispicatus Bauters, Reynders & Goetgh. in Novon 20: 137. 2010.
Holotypus: Angola. Prov. Huíla: near Mupa Catholic Mission in Cuanhama, 1250 m, 9.IX.1952, H. & E. Hess 52/34 (GENT).
Note. – Bauters et al. (2010: 135) propose a comparative table of morphologically similar species, all cited under Cyperus (C. absconditicoronatus, C. rhynchosporoides, C. stramineoferrugineus).
Pycreus micropelophilus (Lye) J.-P. Lebrun & Stork, comb. nova.
≡ Cyperus micropelophilus Lye in Willdenowia 26: 233. 1996.
Holotypus: Somalia. Reg. Bay: Bur Akaba inselberg, 20.VI.1983, Gillett & Hemming 24892 (K [K000321407] image seen).
Notes. – Cyperus micropelophilus was first introduced by Lye in Thulin (1995: 144) but lacking a Latin diagnosis, and then validated by LYE (1996d: 233). The species is mentioned by LYE (2001: 204) under Cyperus.
Acknowledgements
The authors are much indebted to Martin Callmander for his interest in their work on the African flora and for his remarks on and editing of this text. They also wish to thank Cyrille Chatelain for his important contribution to this article, as well as Nicolas Fumeaux who verified items of nomenclature. Finally, they thank Lorenzo Ramella and Patrick Bungener for their help in edition.
