See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/233597555 Nomenclature and typification of names of genera and subdivisions of genera in Cypereae (Cyperaceae): 2. Names of subdivisions of Cyperus Article in Taxon · June 2011 CITATIONS READS 26 338 7 authors, including: Isabel Larridon Wim Huygh 64 PUBLICATIONS 282 CITATIONS 20 PUBLICATIONS 184 CITATIONS Royal Botanic Gardens, Kew SEE PROFILE Ghent University SEE PROFILE Marc Reynders A. Muthama Muasya 36 PUBLICATIONS 300 CITATIONS 260 PUBLICATIONS 2,359 CITATIONS Botanic Garden Meise SEE PROFILE University of Cape Town SEE PROFILE Some of the authors of this publication are also working on these related projects: Flora of Thailand Poaceae View project Biodiversity and Conservation of Chilean cacti View project All content following this page was uploaded by Isabel Larridon on 23 May 2014. The user has requested enhancement of the downloaded file. Larridon & al. • Names of subdivisions of Cyperus TAXON 60 (3) • June 2011: 868–884 Nomenclature and typification of names of genera and subdivisions of genera in Cypereae (Cyperaceae): 2. Names of subdivisions of Cyperus Isabel Larridon,1 Wim Huygh,1 Marc Reynders,1 A. Muthama Muasya, 2 Rafaël Govaerts, 3 David A. Simpson3 & Paul Goetghebeur1 1 Ghent University, Department of Biology, Research Group Spermatophytes, K.L. Ledeganckstraat 35, 9000 Gent, Belgium 2 Botany Department, University of Cape Town, Rondebosch 7701, Cape Town, South Africa 3 Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K. Author for correspondence: Isabel Larridon, Isabel.Larridon@UGent.be Abstract This paper is the second in a series of papers on the nomenclature of the names of genera (Huygh & al., 2010) and subdivisions of genera in Cypereae (Cyperaceae). Here we present a survey of all published names of subdivisions of Cype­ rus, designate types where needed, and evaluate priority and legitimacy. Cyperus is the largest genus in Cypereae and is the second-largest genus in the family. The high diversity in Cyperus and the complex relationships with its segregate genera resulted in the accumulation of more than 220 names of subdivisions in Cyperus of which 18 are not validly published and 33 are illegitimate. Keywords Cyperaceae; Cyperus; subdivisions of genera; nomenclature; typification INTRODUCTION Cyperus is the second-largest genus in Cyperaceae and is its most important genus in the tropics. It is the core taxon of the Cypereae tribe. Recent molecular phylogenetic studies of Cyperaceae (Muasya & al., 2002, 2009a; Simpson & al., 2007), based on markers from the chloroplast genome, reveal the paraphyletic character of Cyperus s.str. In the Cyperus clade, several taxa are nested which have diverged from the “typical” Cyperus vegetative, floral and anatomical characters. Spikelets with distichously arranged glumes have traditionally been the diagnostic character for Cyperus and its closest allies. However, Muasya & al. (2002) conclude that several taxa with spirally arranged glumes are present in the Cyperus clade (e.g., Cyperus michelianus, Kyllingiella, Oxy­ caryum). Species with distichously arranged glumes within Ficinia and Isolepis (I. levynsiana) also occur, blurring the generic delimitations of Cypereae genera. These findings urge a recircumscription of the genera in this tribe (Muasya & al., 2009b). The Cyperus clade has a very complex generic and subdivisional nomenclature with approximately 350 generic and subdivisional names to accommodate the roughly 950 species present in the group. The subdivisional classification of Cype­ rus alone comprises more than 220 names of which 7% are not validly published and 12% are found to be illegitimate after evaluation in accordance with the International Code of Bo­ tanical Nomenclature (ICBN , McNeill & al., 2006). The lack of overview and the limited availability of literature sources in the past have led to the erroneous use of subdivisional names with regards to valid publication and legitimacy in almost all recent and past classifications of the group (e.g., Kükenthal, 1935–36; Chermezon, 1937; Kern, 1974; Haines & Lye, 1983). 868 An overview of all published names of subdivisions of Cyperus and the nomenclatural priority and typification of each is a major necessity when creating a modern classification of the Cyperus clade based on the ongoing molecular and morphological studies. This paper is part of our effort to provide the first complete nomenclatural survey of the names of genera and subdivisions of genera in Cypereae. In this paper, the second in a series of four papers, we present a survey of all published names of subdivisions of Cyperus, designate types where needed, and evaluate priority and legitimacy. A previous paper treated all names published at generic rank in the Cyperus clade (Huygh & al., 2010). A third (Reynders & al., 2011) and fourth paper will respectively present a synopsis of all infrageneric names published in the segregate genera of Cyperus, and a synopsis of all names published in taxa of the Ficinia clade. History of infrageneric taxonomy of Cyperus. — The morphological diversity and the presence of several convergent evolutionary lines in Cyperus result in various controversial classifications. Furthermore, there is no unanimity on the delimitation of the genus. Metcalfe (1971) already realised that to study relationships within Cyperaceae using only morphological characters is problematic: “The main difficulty about the classification of the Cyperaceae when the subject is approached solely along traditional lines is that the flowers are very small, the exact morphology of their parts is often obscure, and the morphology of the inflorescences is difficult to interpret.” This leads to uncertain homologies and conflicting interpretations. The first infrageneric classification of the genus (and of related genera which were later included in the genus) is that of Nees (1834). He was the first to apply a sectional division in the genus Cyperus, and created eight sections. Often, his sections have incorrectly been assigned to Kunth (1837) as stated by Väre & Kukkonen (2006). Bentham (1881) comments TAXON 60 (3) • June 2011: 868–884 on Nees’s work that “he created some confusion, as well by his usual tendency to raise species to the rank of genera, as by a want of reference to the original papers or work where Cyperaceae had been described, and by using a terminology occasionally founded on mistaken views of the homology of floral organs.” Most genera created by Nees in Cyperaceae are no longer accepted as such. Kunth (1837) greatly expanded our knowledge of Cyper­ aceae. He classified the species of Cyperus with a trifid style into 19 groups of unspecified rank, now usually treated at sectional level. He discussed the species of Cyperus with a bifid style separately. Kunth (1837) accepted Mariscus and Kyllinga as distinct genera. His main error was, according to Bentham (1881), “a tendency to give as characters rather what in theory we ought to see than what we actually do see, and, in his later works, to describe specimens rather than species”. Steudel (1854) divided Cyperus into three unnamed sections. The first section included species with a bifid style and corresponds to the genus Pycreus. His “Sectio II”, split into 17 groups of unspecified rank, comprised the species of Cyperus with a trifid style. Many of these groups are now treated at sectional level. The third section held the species with uncertain affinities. Furthermore, Steudel (1854) recognised Mariscus and Kyllinga (“Kyllingia”) as segregate genera. Böckeler (1868–77) studied Cyperaceae in the Berlin Herbarium. Like Steudel (1854), Böckeler (1868) treated Kyllinga (“Kyllingia”) as a separate genus, and included Pycreus at sectional level in the genus Cyperus alongside a section “Eucype­ rus”. However, he did not accept Mariscus at generic rank. Böckeler (1868) divided his section “Eucyperus” into 20 groups of unspecified rank, now usually treated at sectional level. Clarke (1883, 1884, 1893, 1897, 1900, 1902, 1908), the founder of modern cyperology, originally considered Cype­ rus in a wide sense, but in his later publications he proposed the subdivision of Cyperus s.l. into seven distinct genera, i.e., Cyperus, Courtoisina, Juncellus, Kyllinga, Mariscus, Pycreus, and Torulinium. The last complete revision of Cyperus was published in the generic monograph of Kükenthal (1935–36). His infrageneric classification is the one most commonly used at present. As in Suringar’s (1898) excellent revision of Cyperus in the Malaysian Archipelago, the genus is taken in its wide sense. Kükenthal (1935–36) divided Cyperus s.l. into six subgenera (C. subgg. “Eucyperus”, Juncellus, Pycreus, Mariscus, Kyl­ linga, and Torulinium), 61 sections and eight subsections. The sections were primarily delineated by the nature of branching of the compound inflorescence, extent of development of the rhizomes, and the number of stamens and carpels per flower. Obviously, many of Kükenthal’s sections need revision before a natural classification of the genus can be proposed. Van der Veken (1965) studied the embryos of 132 species of Cyperus belonging in different subdivisions of the genus. The uniformity of the embryos appears to support the wide concept of Cyperus. Furthermore, the study revealed the presence of embryos of the Cyperus type in many taxa previously placed near Scirpus (e.g., Ascolepis, Ficinia, Isolepis, Lipocarpha, Kyllingiella, and Oxycaryum). Since Van der Veken’s (1965) Larridon & al. • Names of subdivisions of Cyperus publication, the inclusion of these genera in Cypereae led to a more natural circumscription of that tribe. All of these genera have, until now, been treated separately from Cyperus, except for the treatment of Lipocarpha under Cyperus by Koyama (1961). In his treatment of the genus, Kern (1974) only accepted C. subg. Cyperus, Pycreus and Kyllinga at subgeneric rank, as he considered Juncellus, Mariscus, and Torulinium too ill defined to be recognised as subgenera. Later publications are only regional studies of Cyperus (e.g., Kern, 1974; Haines & Lye, 1983). Although they present extensive knowledge of the genus, they are decidedly less valuable in assessing species relationships in a large genus such as Cyperus, the members of which have successfully utilised long-distance dispersal as well as more gradual short-distance migration and in situ diversification. Goetghebeur (1989) discussed the problems in the lectotypification of names relating to the infrageneric classification of Cyperus. He specifically excluded all taxa with laterally compressed nutlets, i.e. Kyllinga, Pycreus and Queenslandiella, and several highly specialised taxa, i.e., Alinula, Courtoisina, Kyllingiella, Oxycaryum, Remirea and Sphaerocyperus, from Cyperus s.str. However, he included such segregates as Ano­ sporum, Galilea, Juncellus, Mariscus, Sorostachys and Tor­ ulinium. Goetghebeur (1998) retained this perspective in his treatment of the family Cyperaceae. Current views on the infrageneric classification of Cyperus. — Cyperus s.str. is commonly divided into two units, determined by the alternative character states of an anatomical and of an inflorescence character set. The presence of Kranz anatomy, correlated with C4 photosynthesis, has been used in the classification of Cyperus since Rikli (1895). The vegetative anatomy is eucyperoid or chlorocyperoid, characterising “C. subg. Eucyperus” (Grisebach, 1846) and “C. subg. Chloro­ cyperus” (Schischkin, 1935). The inflorescence is composed of digitately clustered spikelets and is often condensed, or the inflorescence is an anthela composed of spikes of spikelets, characterising C. subg. Pycnostachys and C. subg. Choris­ tachys (Clarke, 1893). As demonstrated by Goetghebeur (1989), the two classification systems are not completely congruent. Either a few sections of subg. Chlorocyperus (with predominantly open inflorescence) exhibit a (partly) condensed and depauperate inflorescence (Kükenthal, 1935–36), or a few sections of the predominantly eucyperoid subgenus Pycnostachys reveal a chlorocyperoid type of anatomy (Druyts-Voets, 1970) linked to C4 photosynthesis. Inevitably, this means that at least one, or perhaps both of these presumed apomorphic character states (chlorocyperoid anatomy, condensed inflorescence) has or have evolved several times. This raises questions about the switch from C3 to C4 anatomy either having evolved repeatedly with no significant morphological divergence or only once with subsequent convergence in the morphology of unrelated species. Clearly these are problems that cannot be resolved using morphological data alone. As already suggested by Raynal (1973) and Goetghebeur (1989), Soros & Bruhl (2000) confirmed that the chlorocyperoid anatomy type arose only once within Cyperus. Soros & Bruhl (2000) stated that the C4 photosynthetic pathway arose at least four separate times in 869 Larridon & al. • Names of subdivisions of Cyperus Cyperaceae. However, in the tribe Cypereae only the chlorocyperoid anatomical variant occurs. Ongoing molecular research suggests that the Cyperus C3 species form a paraphyletic group with a monophyletic C4 subclade nested within the C3 group (Muasya & al., 2002, 2009a; Simpson & al., 2007; Larridon & al., in press). On the other hand, Goetghebeur (1989) remarked that a multiple origin of the condensed inflorescence within Cy­ perus is much more probable. There are many instances of species exhibiting both inflorescence types (open vs. condensed), in Cyperaceae in general, and in Cyperus in particular. Consequently, within Cyperus at least two main infrageneric groups have been recognised. An eucyperoid subgenus, uniting plants without the Kranz syndrome, and with an inflorescence composed of digitately clustered spikelets; and a chlorocyperoid subgenus, uniting plants with a chlorocyperoid type of Kranz syndrome, and an inflorescence composed of spikes of spikelets or condensed spikes. Within Cyperus, two subgenera thus circumscribed were recognised by Chermezon (1937), Raynal (1973), Tucker (1983) and Hooper (1985). However, Hooper (1985) also upheld C. subg. Juncellus, accommodating the chlorocyperoid species with a condensed inflorescence and dorsiventrally flattened nutlets. Even more subgenera were proposed by Haines & Lye (1983), who considered Cyperus in a broad sense, although in Lye’s earlier work generic segregates such as Anosporum, Sorostachys (Lye, 1981b), Kyllinga (Lye, 1982), and Pycreus (Lye, 1981c) were recognised. Goetghebeur (1989) remarked that he failed to see the advantage of merging such easily recognisable taxa with obvious synapomorphies, like Alinula, Ascolepis, Courtoisina, Kyllinga, Kyllingiella, Lipocarpha, Oxycaryum, Pycreus and Queenslandiella. On the other hand Lye (1992) convincingly demonstrated the polyphyly of Mariscus within Cyperus. Most authors now accept Mariscus as an element of Cyperus (e.g., Goetghebeur, 1998; Simpson & Koyama, 1998). Goetghebeur (1989) proposed the recognition within Cyperus of only two subgenera, respectively composed of the eucyperoid and of the chlorocyperoid species, further arranged in sections. The recent molecular studies (Muasya & al., 2002, 2009a; Simpson & al., 2007; Larridon & al., in press) reveal the paraphyletic character of Cyperus s.str. including several highly derived lineages, urging a redefinition of generic delimitations in Cypereae. MATERIALS AND METHODS The typifications were made after careful review of the original descriptions of the taxa, examination of herbarium specimens, and of live specimens in the Ghent University Botanical Garden. They are founded on expertise in Cyperaceae obtained through long-standing research into the systematics of this family by the Research Group Spermatophytes of the Ghent University. References to Articles refer to the Interna­ tional Code of Botanical Nomenclature (ICBN , McNeill & al., 2006). References to accepted names refer to the World Checklist of Cyperaceae (Govaerts & al., 2007, 2010). The classification used is that of Goetghebeur (1998). 870 TAXON 60 (3) • June 2011: 868–884 The type of each name of a subdivision of Cyperus is indicated. If in the protologue only one validly published species name is cited or referred to (Art. 10.3 of the ICBN), the entry appears simply as “Type: …” Where a type has been selected later, the entry appears as “Lectotype” with a parenthetical reference to the publication in which the selection was made. Names that are homotypic, e.g., under Art. 7.5, are indicated appropriately. If, when first published, the name of a subdivision of Cyperus had been assigned to a generic subdivision of higher rank, that assignment is given in parentheses even although it has no nomenclatural significance. The large number of nomina nuda published by Chermezon are a consequence of the fact that Chermezon (1922, 1931) did not provide descriptions for his intended new names, and later Chermezon (1937) only provided a French diagnosis; therefore his names are not validly published (Art. 32.1(d) and 36.1). Of the names of subdivisions of Cyperus with epithets that are derived from the epithet of a constituent species and hence typified under Art. 22.6 by the type of that species name, a significant number, when published, included the already established type of a name of another generic subdivision of the same rank, thereby making them superfluous and under a strict reading of Art. 52, also illegitimate. As Art. 22.6 establishes that such a name has a different type from that of the name that ought to have been adopted, automatic typification under Art. 7.5 does not apply. The resultant situation is very unsatisfactory in that perfectly appropriate epithets of subdivisions of Cyperus based on the names of familiar species, are blocked from use even when the taxon is circumscribed to exclude the type of any earlier name at that rank. One might suppose by analogy with superfluous names formed from legitimate basionyms (Art. 52.3) that such names would be available for use when the cause of the superfluity was removed, but a strict reading of Art. 52.3 does not permit this. As it would seem logical that the Code should extend the provisions of Art. 52.3 to such cases, which are particularly important in large genera such as Cyperus, we have noted the superfluous situation in the listings below, but have refrained from adding the word “illegitimate”. NOMENCLATURE Cyperus [unranked] Alati C.B. Clarke (see Cyperus L. sect. Cyperus). Cyperus sect. Alati Kük., nom. illeg. (see Kyllinga sect. Alatae (Kük.) Tang & F.T. Wang; Reynders & al., 2011). Cyperus (subg. Pycreus) sect. Albomarginati Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 359. 1936 – Type: Cyperus albomarginatus (Mart. & Schrad. ex Nees) Steud. [= Pycreus macrostachyos (Lam.) J. Raynal] (Art. 22.6). This sectional epithet has not been combined in Pycreus. Cyperus subg. Alinula (J. Raynal) Lye in Nordic J. Bot. 3: 230. 1983 ≡ Alinula J. Raynal in Adansonia, ser. 2, 17: 43. 1977 TAXON 60 (3) • June 2011: 868–884 Larridon & al. • Names of subdivisions of Cyperus – Type: Cyperus lipocarphioides (Kük.) Lye [≡ Alinula lipocarphioides (Kük.) J. Raynal (Art. 7.4)]. ≡ Alinula J. Raynal (see Huygh & al., 2010). Repentes Kunth are validly published under Art. 34.2. Of the two names Jaubert & Spach (1844) selected the epithet of the former for this section. Cyperus sect. Allagostachyi Nees in Linnaea 9: 284. 1834 – Lectotype (designated here): Cyperus hexastachyos Rottb. [= C. rotundus L.]. Nees (1834) noted “ex affinitate Cyperi hexastachyi ” in his description of C. sect. Allagostachyi. According to Kükenthal (1936), however, Nees interpreted this species incorrectly, his plant being C. zollingeri Steud. Cyperus sect. Aristati Nees in Linnaea 9: 284. 1834 – Type: Cyperus aristatus Rottb., nom. illeg. (Art. 52.1) (≡ Scirpus intricatus L., non C. intricatus Schrad.) [= C. squarro­ sus L.] (Art. 22.6). Often, this section has been incorrectly assigned to Kunth (1837) as stated by Väre & Kukkonen (2006). Cyperus sect. Alopecuroidei Nees in Linnaea 9: 285. 1834 – Type: Cyperus alopecuroides Rottb. (Art. 22.6). Väre & Kukkonen (2006) remarked that this sectional name has often been incorrectly assigned to Kunth (1837). Cyperus sect. Alternifolii (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 20: 288. 1883 ≡ C. [unranked] Alternifolii Kunth, Enum. Pl. 2: 31. 1837 – Type: Cyperus alternifolius L. (Art. 22.6). Cyperus sect. Amabiles C.B. Clarke in Hooker, Fl. Brit. India 6: 598. 1893 – Type: Cyperus amabilis Vahl (Art. 22.6). Clarke (1893) originally includes three species in this section, i.e., Cyperus amabilis, C. castaneus Willd., and C. cus­ pidatus Kunth. Cyperus (sect. Diffusi) [unranked] Angustifolii C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 98. 1908 (‘Angusti­ foliae’) – Lectotype (designated here): Cyperus derreilema Steud. We designate C. derreilema as lectotype since its characters (narrow leaves) fit very well the description of C. [unranked] Angustifolii. Kükenthal (1936) later organised the 11 species of this group of unspecified rank into the sections Diffusi (4 spp.), Fusci (2 spp.), Haspani (1 sp.), and Incurvi (4 spp.). Cyperus subg. Anosporum (Nees) C.B. Clarke in J. Linn. Soc., Bot. 21: 34. 1884 ≡ Anosporum Nees in Linnaea 9: 287. 1834 ≡ C. sect. Anosporum (Nees) Pax in Engler, Nat. Pflanzenfam. 2, 2: 107. 1887 – Type: Cyperus monocepha­ lus Roxb. [= C. cephalotes Vahl]. Cyperus subsect. Apteri Nakai (see Cyperus sect. Compressi Nees). Cyperus sect. Arenarii (Kunth) Jaub. & Spach, Ill. Pl. Orient. 2, 11: t. 101. 1844 ≡ C. [unranked] Arenarii sive Repentes Kunth, Enum. Pl. 2: 46. 1837 – Type: Cyperus arenarius Retz. (Art. 22.6). Väre & Kukkonen (2006) remark that as to the name of the section, Kunth (1837) caused some confusion, because he simultaneously proposed two different names based on the same type for the same taxon, viz. “Arenarii sive Repentes” both the names C. [unranked] Arenarii Kunth and C. [unranked] Cyperus subg. Aristomariscus Lye in Nordic J. Bot. 3: 213. 1983 ≡ C. (“sect. Eucyperus”) [unranked] Squarrosi Benth., Fl. Austral. 7: 255. 1878 – Type: Cyperus squar­ rosus L. Bentham (1878) published C. [unranked] Squarrosi without mention of rank. Lye (1983) published C. subg. Aristomariscus based on the same type. “Cyperus sect. Bernieriani Cherm.” in Humbert, Fl. Madagasc. 29: 104. 1937, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. bernieri Cherm. Cyperus sect. Bobartia (L.) C.B. Clarke in J. Linn. Soc., Bot. 21: 110. 1884 – Type: Bobartia indica L. (Art. 7.4). ≡ Bobartia L. Clarke (1884) based C. sect. Bobartia on the monotypic genus Bobartia L. The only species described by Linnaeus (1753), Bobartia indica L., has since been typified with a specimen in the Hermann Herbarium (BM), which applies the name to an iridaceous plant from southern Africa, traditionally placed in the genus Bobartia (Strid, 1974; Väre & Kukkonen, 2006). Although Väre & Kukkonen (2006) stated that this means that the name of C. sect. Bobartia should be attributed to Clarke (1884) alone, according to Art. 7.4 a new name formed from a previously published legitimate name is, in all circumstances, typified by the type of the basionym, even though it may have been applied erroneously to a taxon now considered not to include that type. Clarke (1884) considered Bobartia indica L. to represent C. pachyrrhizus, but later Clarke (1893) identified it with C. arenarius Retz. (a species he explicitly excluded when he established C. sect. Bobartia) when he established C. sect. Con­ glomerati, nom. illeg., a section uniting the species C. arenar­ ius, C. conglomeratus, and C. pachyrrhizus. Kükenthal (1936) and Koyama (1985) cite Bobartia indica L. as a synonym of C. arenarius. Cyperus [unranked] Bracteati Boeck., nom. illeg. (see Cyperus sect. Luzuloidei (Kunth) C.B. Clarke). Cyperus sect. Brevifoliati C.B. Clarke, nom. illeg. (see Cyperus sect. Corymbosus (Kunth) C.B. Clarke). Cyperus subsect. Brevifolii Kük. (see Kyllinga sect. Brevifoliae (Kük.) Kukkonen; Reynders & al., 2011). 871 Larridon & al. • Names of subdivisions of Cyperus Cyperus (subg. Mariscus) subsect. Brevistyli Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 522. 1936 – Lectotype (designated here): Cyperus cylindrostachyus Boeck. Cyperus cylindrostachyus is the type of the name C. sect. Umbellati, nom. illeg. in which C. subsect. Brevistyli is included. Cyperus subg. Bulbocaules (C.B. Clarke) Lye in Nordic. J. Bot. 3: 230. 1983 (‘Bulbocaulis’) ≡ Mariscus subg. Bulbocaules C.B. Clarke in Hooker, Fl. Brit. India 6: 619. 1893 ≡ M. sect. Bulbocaules (C.B. Clarke) C.B. Clarke in Dyer, Fl. Cap. 7: 185. 1897 ≡ C. (subg. Mariscus) sect. Bulbocaules (C.B. Clarke) Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 538. 1936 – Lectotype (designated here): Mariscus dregeanus Kunth [= Cyperus dubius Rottb.]. Clarke (1893) originally included two species in this subgenus, i.e., M. dregeanus and M. bulbosus C.B. Clarke, nom. illeg., non Steud. [≡ C. clarkei T. Cooke]. Lye (1983) was incorrect to indicate C. dubius as lectotype of his new combination C. subg. Bulbocaules (C.B. Clarke) Lye because Clarke (1893: 620) definitely excluded the type of C. dubius from Mariscus dregeanus Kunth (1837: 120). Based on Lye’s choice for C. dubius in 1983, we here correctly designate Mariscus dregeanus as lectotype of the basionym M. subg. Bulbocaules C.B. Clarke (1893) and by extension as type of all names formed from this previously published legitimate name (Art. 7.4). However, the currently accepted name in Cyperus for Mariscus dregeanus is still Cyperus dubius Rottb. (Govaerts & al., 2007, 2010). It is important to note that Cyperus dregeanus Kunth (1837: 31) is different from Mariscus dregeanus Kunth (1837: 120). Clarke (1908) added 29 species (including M. bulbocaulis Hochst. ex A. Rich. [= C. plateilema (Steud.) Kük.]) and published this taxon at sectional level. He divided C. sect. Bulbo­ caules in two unnamed series differing in having either one or more nutlets ripening per spikelet. Cyperus subg. Bulbomariscus Lye in Nordic J. Bot. 3: 214. 1983 – Type: Cyperus amauropus Steud. Cyperus sect. Bulbosi C.B. Clarke in Hooker, Fl. Brit. India 6: 611. 1893 – Type: Cyperus bulbosus Vahl (Art. 22.6). Chermezon also lists a C. sect. Bulbosi in 1922 and 1937 without any author citation nor description and with C. escu­ lentus L. as only species. It is unclear whether Chermezon intended this as a new section, but in that case the name would be a nomen nudum and would also not be validly published (Art. 22.2), as C. esculentus was selected as type of Cyperus (Britton, 1907). Cyperus subg. Caespitosi Nakai (see Cyperus sect. Irioidei Nees). Cyperus (subg. Mariscus) subsect. Capenses Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 538. 1936 – Type: Cyperus capensis (Steud.) Endl. (Art. 22.6). 872 TAXON 60 (3) • June 2011: 868–884 Cyperus [unranked] Capitati Boeck. (see Cyperus sect. Leuco­ cephali Cherm. ex Kük.). “Cyperus [unranked] Capitati multinervii Steud.”, Syn. Pl. Glumac. 2: 27. 1854, nom. inval. Steudel (1854) used two words to name this proposed subdivision. The Code does not directly address the use of such names of subdivisions of genera. However, we conclude that this name is not in accordance with Art. 21.2, which states that the epithet of a subdivision of a genus is either of the same form as a generic name, or a noun in the genitive plural, or a plural adjective agreeing in gender with the generic name, but not a noun in the genitive singular. In addition, Art. 20.3 states that the name of a genus may not consist of two words, unless these words are joined by a hyphen, and Steudel did not link the words in this way. Cyperus sect. Capituligeri Nees in Linnaea 9: 284. 1834 ≡ C. sect. Tenelli C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 96. 1908 (‘Tenellae’) – Type: Cyperus tenel­ lus L. f. [≡ Isolepis levynsiana Muasya & D.A. Simpson] (lectotype C. sect. Capituligeri designated here – type C. sect. Tenelli Art. 22.6). We designate C. tenellus applicable to a species now included in Isolepis (Muasya & al., 2006, 2007), as lectotype of this name. Nees (1834) included 29 species in C. sect. Capitu­ ligeri based on the presence of a head-like inflorescence or an umbel with head-like inflorescences. Although Nees mentioned C. tenellus in both his C. sect. Capituligeri and C. sect. Allagostachyi, since it has head-like spikelet clusters it clearly belongs in C. sect. Capituligeri. Since Nees (1834), C. sect. Capituligeri has not been used by any later author; his work from 1834 has been overlooked by most authors as mentioned by Väre & Kukkonen (2006). The combination in Isolepis has not been made. Cyperus sect. Cephalotes J.V. Suringar, Cyperus: 76. 1898, nom. superfl. – Type: Cyperus cephalotes Vahl (Art. 22.6). = Cyperus sect. Anosporum (Nees) Pax. When published, Cyperus sect. Cephalotes included the type of C. sect. Anosporum (Nees) Pax (1887) and is therefore superfluous. “Cyperus subg. Chlorocyperus (Rikli) Schischk.” in Komarov, Fl. URSS 3: 17. 1935, nom. inval. (Art. 22.2). “Cyperus subg. Chlorocyperus” is not validly published (Art. 22.2), as it included C. esculentus which was selected as type of Cyperus (Britton, 1907). This proposed name was based on: Chlorocyperus Rikli, Jahrb. Wiss. Bot. 27: 563. 1895, nom. illeg. (see Huygh & al., 2010). Cyperus subg. Choristachys C.B. Clarke, Cyperus sect. Chori­ stachys (C.B. Clarke) C.B. Clarke (see Cyperus L. sect. Cyperus). Cyperus sect. Chrysanthi (C.B. Clarke) J. Kern (see Pycreus sect. Chrysanthi C.B. Clarke; Reynders & al., 2011). TAXON 60 (3) • June 2011: 868–884 “Cyperus sect. Compacti Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 31. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. compactus Retz. Cyperus sect. Compressi Nees in Linnaea 9: 284. 1834 ≡ C. (“sect. Eucyperus”) [unranked] Compressi (Nees) Benth., Fl. Austral. 7: 255. 1878 ≡ C. [unranked] Sub­ exalati C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 313. 1901 (‘Subexalatae’) ≡ C. (sect. Pseudohaspani) subsect. Apteri Nakai in Bot. Mag. (Tokyo) 26: 189. 1912 – Type: Cyperus compressus L. (type C. sect. Compressi Art. 22.6 – type C. subsect. Apteri – lectotype Subexalati designated here). Väre & Kukkonen (2006) remarked that C. sect. Com­ pressi has often been incorrectly assigned to Kunth (1837). We designate C. compressus as lectotype of C. [unranked] Subexalati as this species clearly has an unwinged rachilla. Clarke (1901) created this group of unspecified rank, based on his former C. subg. Choristachys series A from 1893, and having a wingless or scarcely winged rachilla to differentiate it from C. [unranked] Alati in which the rachilla is obviously winged. Clarke (1901) divided C. [unranked] Subexalati into three sections but named only the first section (i.e., C. sect. Compressi). Cyperus subsect. Apteri Nakai has the same type as C. sect. Compressi Nees (1834) because C. compressus is the only species included by Nakai (1912), but C. subsect. Apteri is legitimate with priority at the rank of subsection. Cyperus sect. Congesti C.B. Clarke in J. Linn. Soc., Bot. 20: 285. 1883 – Lectotype (designated here): Cyperus exilis Willd ex Kunth. Originally, Clarke (1883) included five species in this section based on the presence of a simple to congested anthela: C. dubius Rottb., C. obtusiflorus Vahl [= C. niveus var. leuco­ cephalus (Kunth) Fosberg], C. exilis, C. stoloniferus Retz., and C. pervillei Boeck [= Pycreus pervillei (Boeck.) C.B. Clarke]. Cyperus sect. Conglomerati C.B. Clarke in Hooker, Fl. Brit. India 6: 602. 1893, nom. illeg. – Type: Cyperus conglo­ meratus Rottb. (Art. 22.6). = Cyperus sect. Arenarii (Kunth) Jaub. & Spach. Cyperus sect. Conglomerati C.B. Clarke (1893) is illegitimate (Art. 52.1) since it includes the type of the older name C. sect. Arenarii (Kunth) Jaub. & Spach (1844) (Art. 11.4). Cyperus sect. Corymbosi (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 21: 153. 1884 ≡ C. [unranked] Corymbosi Kunth, Enum. Pl. 2: 53. 1837 ≡ C. ([unranked] Corym­ bosi) [unranked] Subaphylli Boeck. in Linnaea 36: 271. 1870 ≡ C. sect. Brevifoliati C.B. Clarke in Hooker, Fl. Brit. India 6: 611. 1893 (‘Brevefoliati’), nom. illeg. (Art. 52.1) ≡ C. sect. Tunicati C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 314. 1901 (‘Tunicatae’), nom. illeg. (Art. 52.1) – Type: Cyperus corymbosus Rottb. (type of C. sect. Corymbosi Art. 22.6 – lectotype of C. [unranked] Subaphylli designated here – type of C. sect. Brevifoliati and C. sect. Tunicati Art. 7.5). = Cyperus L. sect. Cyperus (by lectotypification). Larridon & al. • Names of subdivisions of Cyperus Bentham (1878: 256) mentioned Kunth’s name Corymbosi as a group of unspecified rank under C. sect. Papyrus. Böckeler (1870) described C. [unranked] Subaphylli as a subgroup under C. [unranked] Corymbosi characterised by reduced leaves. Several authors published similar names of unspecified rank for the same group of species. These names were not validly published and might not have been intended as names of subdivisions of Cyperus, i.e., “Aphylli ” (Kunth, 1837: 53), “Culmo aphyllo” (Steudel, 1854: 30), “Folia abbreviata aut subnulla” (Clarke, 1883: 291) and “Sub-efoliati ” (Suringar, 1898: 121). All these authors also published names for an analogous group of species at unspecified rank characterised by well-developed leaves, i.e., “Foliati ” (Kunth, 1837: 57), “Culmo foliato” (Steudel, 1854: 32), “Foliati ” (Böckeler, 1870: 277), “Foliati ” (Clarke, 1883: 292) and “Foliati ” (Suringar, 1898: 107). However, all these names were published without a diagnosis or description (Art. 32.1(d) and 36.1). Cyperus sect. Brevifoliati C.B. Clarke (1893) and C. sect. Tunicati C.B. Clarke (1901) are illegitimate (Art. 52.1) since they include the type of the older name C. sect. Corymbosi (Kunth) C.B. Clarke (1884). Cyperus sect. Corymbosi Nakai, nom. illeg. (see Cyperus sect. Irioidei Nees). Cyperus subg. Courtoisia (Nees) Lye in Nordic J. Bot. 3: 230. 1983 ≡ Courtoisia Nees in Linnaea 9: 286. 1834, nom. illeg. (Art. 53.1), non Marchand (1830) ≡ Cyperus subg. Courtoisina (Soják) Lye in Lidia 3, 2: 52. 1992, nom. illeg. (Art. 52.1) – Type: Cyperus pseudokyllingioides Kük. as nomen novum of Courtoisia cyperoides (Roxb.) Nees (Kyl­ linga cyperoides Roxb.) [≡ Courtoisina cyperoides (Roxb.) Soják] (Art. 7.4). ≡ Courtoisina Soják (see Huygh & al., 2010). Although the generic name Courtoisia Nees is an illegitimate later homonym, the subgeneric name Cyperus subg. Courtoisia is legitimate with priority from its date of publication (1983) (Art. 58.1). Consequently, it is the correct name for the taxon at subgeneric rank in Cyperus, although Courtoisina Soják is the correct name at generic rank. “Cyperus sect. Cuspidati Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 4. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. cuspidatus Kunth. Cyperus L. sect. Cyperus, autonym automatically established by Nees in Linnaea 9: 284. 1834 (Art. 22.3) ≡ Cyperus L., Sp. Pl. 1: 44. 1753 (Huygh & al., 2010) ≡ Cyperus subg. Cyperus, autonym automatically established in Petermann, Deutschl. Fl. 11: 595. 1849 (Art. 22.3) ≡ C. subg. Pterocy­ perus Peterm., Deutschl. Fl. 11: 595. 1849 ≡ Pterocyperus (Peterm.) Opiz, Seznam: 80. 1852 ≡ C. sect. Solubiles C.B. Clarke in J. Linn. Soc., Bot. 20: 291. 1883 ≡ C. subg. Chori­ stachys C.B. Clarke in Hooker, Fl. Brit. India 6: 605. 1893 ≡ C. sect. Choristachys (C.B. Clarke) C.B. Clarke” in Urban, Symb. Antill. 2: 22. 1900 (‘Choristachyae’) ≡ C. [unranked] 873 Larridon & al. • Names of subdivisions of Cyperus Alati C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 314. 1901 (‘Ala­ tae’) ≡ Cyperus subsect. Cyperus, autonym automatically established by Nakai in Bot. Mag. (Tokyo) 26: 188. 1912 (Art. 22.3) – Lectotype: Cyperus esculentus L. (lectotype of Cyperus designated by Britton (1907) – lectotype of autonyms Art. 7.6 – lectotypes of C. subg. Pterocyperus, C. sect. Solubiles, C. subg. Choristachys, C. sect. Choris­ tachys and C. [unranked] Alati designated here). Because the lectotype of Cyperus was only designated by Britton (1907), Art. 22.2 does not apply on the names of subdivisions of Cyperus including C. esculentus published before 1907. Kern (1974) was the first to effectively use the autonyms of Cyperus at sectional and subgeneric rank. Väre & Kukkonen (2006) erroneously indicated C. rotundus L. as the type of C. sect. Cyperus. Five species were mentioned in the original description of C. subg. Pterocyperus, Opiz (1852) later established the genus Pterocyperus including only C. es­ culentus (Huygh & al., 2010). “Cyperus sect. Debilissimi Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 18. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. debilissimus Baker. Cyperus (subg. Mariscus) sect. Decidui Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 471. 1936 – Type: Cyperus deciduus Boeck. (Art. 22.6). “Cyperus sect. Denudati Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 12. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. denudatus L. f. Cyperus sect. Dichostylis (P. Beauv. ex T. Lestib.) Baill., Hist. Pl. 12: 338. 1893 (see Dichostylis P. Beauv. ex T. Lestib.; Huygh & al., 2010). “Cyperus sect. Dichroostachyi Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 9. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. dichrostachyus Hochst. ex A. Rich. Cyperus sect. Diclidium (Schrad. ex Nees) Griseb., Fl. Brit. W. I.: 565. 1864 ≡ Diclidium Schrad. ex Nees in Martius, Fl. Bras. 2, 1: 51. 1842 ≡ C. subg. Diclidium (Schrad. ex Nees) C.B. Clarke in J. Linn. Soc., Bot. 21: 191. 1884 ≡ Mariscus subg. Torulinium (Desv. ex Ham.) C.B. Clarke in Hooker, Fl. Brit. India 6: 624. 1893 ≡ C. subg. Torulinium (Desv. ex Ham.) Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 614. 1936 ≡ C. (subg. Torulinium) sect. Feraces Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 614. 1936 – Type: Cyperus ferax Rich. [= Cyperus odoratus L. subsp. odo­ ratus] (lectotype of Diclidium and Torulinium designated by Koyama (1961) – type of C. sect. Feraces Art. 22.6). Although Torulinium Desv. ex Ham. has priority at generic level (see Huygh & al., 2010), at subgeneric rank the epithet Diclidium has priority. 874 TAXON 60 (3) • June 2011: 868–884 Cyperus sect. Difformes C.B. Clarke in J. Linn. Soc., Bot. 20: 289. 1883, nom. superfl. ≡ Cyperus (“sect. Eucyperus”) [unranked] Nigricantes Benth., Fl. Austral. 7: 255. 1878 – Type: Cyperus difformis L. (type of C. sect. Difformes Art. 22.6 – lectotype of C. [unranked] Nigricantes designated here). When published, Cyperus sect. Difformes included the type of C. sect. Irioidei Nees (1834) and is therefore superfluous. Bentham’s older name of unspecified rank does not have priority (Art. 35.3). Although most of the epithets in Bentham’s named subdivisions of Cyperus in Flora Australiensis appear to be derived from species epithets and from the context some, such as C. [unranked] Compressi (above) clearly are, there seems no possibility that Bentham intended his C. [unranked] Nigricantes to include either C. nigricans (L.) With. (1796) (≡ Schoenus nigricans L.) or C. nigricans Steud. (1842), nom. illeg. (≡ Pycreus nigricans C.B. Clarke). Cyperus sect. Diffusi (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 21: 124. 1884 ≡ C. [unranked] Diffusi Kunth, Enum. Pl. 2: 25. 1837 ≡ C. subg. Pycnostachys C.B. Clarke in Hooker, Fl. Brit. India 6: 597. 1893 ≡ C. sect. Pycnostachys (C.B. Clarke) C.B. Clarke in Urban, Symb. Antill. 2: 22. 1900 (‘Pycnostachyae’) ≡ C. (Diffusi) [unranked] Eudiffusi C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 98. 1908 (‘Eu­Diffusae’) – Type: Cyperus diffusus Vahl. (type of C. sect. Diffusi Art. 22.6 – lectotype of C. subg. Pycno­ stachys (≡ C. sect Pycnostachys) designated by Tucker (1987) – type of C. [unranked] Eudiffusi Art. 22.6). Clarke (1908) mistakenly describes two sections XII: the Diffusi and the Eudiffusi, the latter containing the type, C. dif­ fusus. Possibly, this error can be blamed on the posthumously publishing of this work and Eudiffusi was meant as a group of unspecified rank under sectional rank to contrast with C. [unranked] Simplices and C. [unranked] Angustifolii, which have either a simple inflorescence or narrower leaves. Although the epithet in this subdivision begins with the prefix Eu-, Art. 21.3 does not apply here. Cyperus [unranked] Digitati J.V. Suringar, Cyperus: 133. 1898 – Type: Cyperus digitatus Roxb. (Art. 22.6). Cyperus [unranked] Digitati includes the type of C. sect. Exaltati (Kunth) C.B. Clarke (1883), which is the correct name at sectional rank. Cyperus sect. Distantes C.B. Clarke in J. Linn. Soc., Bot. 20: 290. 1883 ≡ C. sect. Longispicati J.V. Suringar, Cyperus: 123. 1898, nom. illeg. (Art. 52.1) ≡ C. [unranked] Distantes (C.B. Clarke) J.V. Suringar, Cyperus: 123. 1898 – Type: Cyperus distans L. f. (Art. 22.6). Suringar (1898: 123) included C. sect. Distantes as a group of unspecified rank in his new, but illegitimate C. sect. Long­ ispicati in which he published two more groups of unspecified rank based on the species C. digitatus Roxb. and C. pilosus Vahl. Cyperus sect. Elegantes C.B. Clarke in J. Linn. Soc., Bot. 20: 288. 1883 – Type: Cyperus elegans L. (Art. 22.6). TAXON 60 (3) • June 2011: 868–884 “Cyperus sect. Esculenti Kük.” in Engler, Pflanzenr. IV, 20 (Heft 101): 116. 1935, nom. inval. (Art. 22.2). “Cyperus subg. Eucyperus Griseb.”, Spic. Fl. Rumel. 2: 420. 1846, nom. inval. (Art. 21.3) – “Cyperus sect. Eucyperus (Griseb.) Boeck.” in Linnaea 35: 493. 1868, nom. inval. (Art. 21.3). Cyperus (Diffusi) [unranked] Eudiffusi C.B. Clarke (see Cype­ rus sect. Diffusi (Kunth) C.B. Clarke). Cyperus sect. Eukyllinga Kük. (see Kyllinga sect. Brevifoliae (Kük.) Kukkonen; Reynders & al., 2011). Cyperus [unranked] Eupycreus Boeck. (see Pycreus P. Beauv. ser. Pycreus; Reynders & al., 2011) Cyperus sect. Exaltati (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 20: 293. 1883 ≡ C. [unranked] Exaltati Kunth, Enum. Pl. 2: 70. 1837 ≡ C. subsect. Exaltati (Kunth) Nakai in Bot. Mag. (Tokyo) 26: 192. 1912 – Type: Cyperus exaltatus Retz. (Art. 22.6). Bentham (1878: 257) adopted Kunth’s epithet Exaltati for a group of unspecified rank under C. sect. Papyrus. Nakai (1912) included C. subsect. Exaltati in his illegitimate C. sect. Corymbosi Nakai. “Cyperus sect. Exiles Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 33. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. exilis Willd. ex Kunth. Cyperus [unranked] Fasciculati Boeck. in Linnaea 36: 342. 1870 – Lectotype (designated here): Cyperus congestus Vahl. The characters of C. congestus fit very well the description of this rankless group (e.g., “spiculae … elongatae lineares … Rhacheola flexuosa compressa angusta, internodiis elongates squamis decurrentibus alatis”). Although this name is validly published under Art. 35.3, its epithet has not been adopted in any ranked subdivision of Cyperus. Cyperus sect. Fastigiati Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 53. 1935 – Type: Cyperus fastigiatus Rottb. (Art. 22.6). Cyperus sect. Feraces Kük. (see Cyperus sect. Diclidium (Schrad. ex Nees) Griseb.). Cyperus (subg. Torulinium) sect. Filiformes Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 624. 1936 – Type: Cyperus filiformis Sw. (Art. 22.6). Cyperus subg. Fimbricyperus Lye in Nordic J. Bot. 1: 60. 1981 – Type: Cyperus subparadoxus Kük. [≡ Alinula paradoxa (Cherm.) Goetgh. & Vorster]. Cyperus sect. Flabelliformes (C.B. Clarke) J. Kern in Fl. Males., ser. 1, 7(3): 638. 1974 ≡ Mariscus sect. Flabelli­ Larridon & al. • Names of subdivisions of Cyperus formes C.B. Clarke in Hooker, Fl. Brit. India 6: 624. 1893 – Type: Mariscus microcephalus J. Presl & C. Presl. [= Cyperus compactus Retz.]. Mariscus microcephalus was the only species included in M. sect. Flabelliformes by Clarke (1893). Kern (1974) considers C. sect. Subulati C.B. Clarke as a synonym of C. sect. Flabelliformes (C.B. Clarke) J. Kern and in consequence erroneously indicated C. subulatus R. Br. as the type of C. sect. Flabelliformes (C.B. Clarke) J. Kern (Art. 7.4). Cyperus sect. Flavescentes Kük. (see Pycreus ser. Zonati (C.B. Clarke) C.B. Clarke; Reynders & al., 2011). Cyperus (sect. Pseudohaspani) subsect. Flavidi Nakai in Bot. Mag. (Tokyo) 26: 188. 1912 – Type: Cyperus flavidus Retz. [= Pycreus flavidus (Retz.) T. Koyama] (Art. 22.6). Nakai (1912) misinterpreted C. flavidus as being closely related to C. haspan L. and C. tenuispica Steud.; he mentioned the type of his C. sect. Pseudohaspani, i.e., C. pseudohaspan Makino [= C. tenuispica Steud.] as a synonym under C. flavi­ dus. This subsection has not been combined in Pycreus. Cyperus (subg. Pycreus) sect. Fontinales Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 340. 1936 – Type: Cyperus fontinalis (Cherm.) Kük. [≡ Pycreus fontinalis Cherm.] (Art. 22.6). This epithet has not been adopted for any ranked subdivision of Pycreus. Cyperus (“sect. Eucyperus”) [unranked] Fulvi Benth., Fl. Austral. 7: 255. 1878 – Type: Cyperus fulvus R. Br. (Art. 22.6). Bentham (1878) published this name without mention of rank. This epithet has not been adopted in any ranked subdivision of Cyperus. Cyperus sect. Fusci (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 21: 131. 1884 ≡ C. [unranked] Fusci Kunth, Enum. Pl. 2: 37. 1837 ≡ C. subsect. Fusci (Kunth) Nakai in Bot. Mag. (Tokyo) 26: 190. 1912 – Type: Cyperus fuscus L. (Art. 22.6). Steudel (1854) synonymised C. sect. Irioidei Nees (1834) and C. [unranked] Fusci Kunth (1837) under the rankless group “Iriae et fusci ”. Nakai (1912) included C. subsect. Fusci in his illegitimate C. sect Corymbosi Nakai. Cyperus sect. Galilea (Parl.) T.V. Egorova in Novosti Sist. Vyssh. Rast. 34: 21. 2002 ≡ Galilea Parl., Fl. Palerm. 1: 297. 1845 (Huygh & al., 2010) – Type: Galilea mucronata (L.) Parl. (Art. 7.4) [= Cyperus capitatus Vand.]. Cyperus sect. Globosi (C.B. Clarke) Kük. (see Pycreus sect. Globosi C.B. Clarke; Reynders & al., 2011). Cyperus sect. Glomerati (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 21: 140. 1884, nom. superfl. ≡ C. [unranked] Glom­ erati Kunth, Enum. Pl. 2: 75. 1837 ≡ Cyperus ([unranked] 875 Larridon & al. • Names of subdivisions of Cyperus Leptolepides) [unranked] Umbellati Boeck. in Linnaea 35: 591. 1868 ≡ Cyperus subsect. Glomerati (Kunth) Nakai in Bot. Mag. (Tokyo) 26: 191. 1912 – Type: Cyperus glomeratus L. (type of C. [subdivisions] Glomerati Art. 22.6 – lectotype of C. [unranked] Umbellati designated here). = Cyperus sect. Distantes C.B. Clarke. When published, Cyperus sect. Glomerati included the type of C. sect. Distantes C.B. Clarke (1883) and is therefore superfluous. The older name of unspecified rank does not have priority (Art. 35.3). Böckeler (1868) distinguishes C. [unranked] Umbellati in his C. [unranked] Leptolepides and included four species: C. semiradiciflorus Boeck. [= C. glomeratus L.], C. kappleri Hochst. ex Steud., C. glomeratus L. and C. prolixus Kunth. Nakai (1912) included C. subsect. Glomerati in his illegitimate C. sect. Corymbosi Nakai. Cyperus sect. Graciles (Benth.) Kük., in Engler, Pflanzenr. IV, 20 (Heft 101): 202. 1936 ≡ Cyperus (“sect. Eucyperus”) [unranked] Graciles Benth., Fl. Austral. 7: 254. 1878 – Type: Cyperus gracilis R. Br. (Art. 22.6). Cyperus sect. Graciles C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 311. 1901, nom. illeg. (Art. 35.3, Art. 53.4), non C. [unranked] Graciles Benth. (1878) – Type: Cyperus cancel­ latus Ridl. [= C. haspan L.]. = Cyperus sect. Haspani (Kunth) C.B. Clarke. Cyperus sect. Haspani (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 20: 287. 1883 ≡ C. [unranked] Haspani Kunth, Enum. Pl. 2: 34. 1837 – Type: Cyperus haspan L. (Art. 22.6). Cyperus sect. Humiles (Kunth) J. Kern in Fl. Males., ser. 1, 7(3): 631. 1974, nom. superfl. ≡ C. [unranked] Humiles Kunth, Enum. Pl. 2: 51. 1837 – Type: Cyperus humilis Kunth (Art. 22.6). When published, Cyperus sect. Humiles included the type of C. sect. Rupestres C.B. Clarke (1908) and is therefore superfluous. The older name of unspecified rank does not have priority (Art. 35.3). Kern (1974) considers C. sect. Rupestres as a synonym of C. [unranked] Humiles and in so incorrectly indicated C. ru­ pestris Kunth as the type species of C. sect. Humiles (Kunth) J. Kern. Cyperus sect. Hymenolepides Nees in Linnaea 9: 284. 1834 ≡ C. [unranked] Platystachyi Kunth, Enum. Pl. 2: 44. 1837 ≡ C. sect. Platystachyi (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 21: 105. 1884 ≡ C. sect. Nivei C.B. Clarke in Hooker, Fl. Brit. India 6: 601. 1893 – Type: Cyperus niveus Retz. (lectotype of C. sect. Hymenolepides designated by Väre & Kukkonen (2006) – lectotype of C. sect. Platystachyi designated here – type of C. sect. Nivei Art. 22.6). Although Kern (1974) lectotypified C. sect. Platystachyi with C. leucocephalus Retz., Kunth (1837) did not include this species in his C. [unranked] Platystachyi (but in his C. [unranked] Kyllingioides) and consequently it is not eligible as 876 TAXON 60 (3) • June 2011: 868–884 type. Instead we select C. niveus. Cyperus niveus was included in the original circumscription of C. [unranked] Platystachyi given by Kunth (1837), and its characters fit very well the description of this group, in contrast to C. leucocephalus. Simpson (1990) clearly states the differences between C. sect. Leucocephali Cherm. ex Kük. and C. sect. Platystachyi. Kern’s choice of type is based on Clarke (1884), who incorrectly placed C. leucocephalus in C. sect. Platystachyi based on the presence of C. sphaerocephalus var. leucocephalus Kunth [= C. niveus var. leucocephalus (Kunth) Fosberg] in Kunth’s Platystachyi. Furthermore, Kunth’s Platystachyi and C. sect. Leucocephali form natural groups in Cyperus and following the subgeneric classification of Goetghebeur (1998) they respectively belong in the C. subg. Cyperus (C4 photosynthesis—chlorocyperoid anatomy) and C. subg. Anosporum (C3 photosynthesis—eucyperoid anatomy) and are in so not closely related. Cyperus sect. Incurvi Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 216. 1936 – Lectotype (designated here): Cyperus disjunctus C.B. Clarke. The characters of C. disjunctus fit very well the description of this section; especially the curved glumes, dilated filaments and well-developed connective of the anthers are clearly visible in this species. Kükenthal (1936) also added an illustration of this species showing the typical characteristics. Cyperus sect. Irioidei Nees in Linnaea 9: 285. 1834 ≡ C. subg. Caespitosi Nakai in Bot. Mag. (Tokyo) 26: 188. 1912 ≡ C. sect. Corymbosi Nakai in Bot. Mag. (Tokyo) 26: 190. 1912, nom. illeg. (Art. 53.4), non C. sect. Corymbosi (Kunth) C.B. Clarke (1884) ≡ C. subsect. Irioidei (Nees) Nakai in Bot. Mag. (Tokyo) 26: 191. 1912 (‘Iriae’) – Type: Cyperus iria L. (type of C. [subdivisions] Irioidei Art. 22.6 – type of C. sect. Corymbosi Nakai, nom. illeg. Art. 7.5 – lectotype of C. subg. Caespitosi designated here). Väre & Kukkonen (2006) remarked that this sectional name has often been incorrectly assigned to Kunth (1837). Kunth (1837: 38) published this name as C. [unranked] Iriae. Steudel (1854) synonymised C. sect. Irioidei Nees (1834) and C. [unranked] Fusci Kunth (1837) under the rankless group Iriae et fusci. Cyperus iria is the type of the name C. subsect. Irioidei and C. sect. Corymbosi Nakai, nom. illeg. which are included in C. subg. Caespitosi. Cyperus sect. Corymbosi Nakai (1912) is illegitimate under Art. 52.1 since it includes the type of the older name C. sect. Irioidei Nees (1834) (Art. 11.4). Nakai (1912) included Nees’s Irioidei as a subsection in his illegitimate section Corymbosi. Like most authors he incorrectly assigned the section based on C. iria to Kunth (1837). Cyperus (subg. Pycreus) [unranked] Isodiametrici Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 327. 1936 – Lectotype (designated here): Cyperus polystachyos Rottb. [≡ Pycreus polystachyos (Rottb.) P. Beauv.]. Cyperus polystachyos is the type of the name C. subg. Pycreus (P. Beauv.) J. Carey in which C. [unranked] Isodia­ TAXON 60 (3) • June 2011: 868–884 metrici is included. Kükenthal (1936) used this name at a level between subgenus and section to replace Pycreus [unranked] Puncticulati C.B. Clarke and P. subg. Reticulati C.B. Clarke (both characterised by nutlets with isodiametric epidermal cells). Although this name is validly published under Art. 35.3, its epithet has not been adopted in any ranked subdivision of Pycreus. Cyperus sect. Ixiocarpi S.T. Blake in Proc. Roy. Soc. Queensland 58: 37. 1947 – Type: Cyperus ixiocarpus F. Muell. (Art. 22.6). Cyperus sect. Juncellus Griseb., Fl. Brit. W. I.: 562. 1864 ≡ C. subg. Juncellus (Griseb.) C.B. Clarke in J. Linn. Soc., Bot. 20: 282. 1883 ≡ Juncellus (Griseb.) C.B. Clarke in Hooker, Fl. Brit. India 6: 594. 1893 ≡ J. sect. Juncellus, autonym automatically established by Nakai in Bot. Mag. (Tokyo) 26: 199 1912 (Art. 22.3) – Type: Cyperus mu­ cronatus Rottb. [= C. laevigatus subsp. distachyos (All.) Maire & Weiller]. Cyperus subg. Kyllinga (Rottb.) J.V. Suringar, Cyperus (subg. Kyllinga) sect. Kyllinga (Rottb.) J. Kern (see Kyllinga Rottb. sect. Kyllinga; Reynders & al., 2011). Cyperus (subg. Mariscus) subsect. Kyllingiformes Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 553. 1936 – Lectotype (designated here): Cyperus dubius Rottb. The characters of C. dubius fit very well the description of this subsection (“Anthela capitato contracta”). Cyperus sect. Kyllingioides (Kunth) J. Kern in Fl. Males., ser. 1, 7(3): 643. 1974 ≡ C. [unranked] Kyllingioides Kunth, Enum. Pl. 2: 94. 1837 – Type: Cyperus kyllingioides Vahl [= C. dubius Rottb.] (Art. 22.6). Cyperus (subg. Juncellus) sect. Laevigati Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 319. 1936 ≡ C. (sect. Py­ creus) [unranked] Pseudopycreus Boeck. in Linnaea 35: 485. 1868 – Type: Cyperus laevigatus L. (type of C. sect. Laevigati Art. 22.6 – lectotype of C. [unranked] Pseudopy­ creus Boeck. designated here). = Cyperus sect. Juncellus Griseb. Cyperus [unranked] Pseudopycreus Boeck. (1868) is different from C. sect. Pseudopycreus (C.B. Clarke) Kük., nom. illeg. (1936). Cyperus sect. Lancei Kük., nom. superfl. (see Pycreus sect. Lancei L.K. Dai; Reynders & al., 2011). Cyperus sect. Latespicati Kük. (see Pycreus sect. Latespicati (Kük.) L.K. Dai; Reynders & al., 2011). Cyperus sect. Latifolii C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 99. 1908 (‘Latifoliae’), nom. superfl. – Type: Cyperus latifolius Poir. (Art. 22.6). = Cyperus sect. Proceri (Kunth) C.B. Clarke. Larridon & al. • Names of subdivisions of Cyperus When published, Cyperus sect. Latifolii included the type of C. sect. Proceri (Kunth) C.B. Clarke (1883) and is therefore superfluous. Cyperus (subg. Mariscus) sect. Laxiglumi (C.B. Clarke) Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 457. 1936 ≡ Mariscus (“sect. Eumariscus”) subsect. Laxiglumi C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 103. 1908 (‘Laxiglumae’) – Lectotype (designated here): Mariscus ischnos (Schltdl.) C.B. Clarke [≡ Cyperus ischnos Schltdl.]. The characters of C. ischnos fit very well the original description of this subsection (Clarke, 1908). Cyperus [unranked] Leptolepides Boeck. (see Cyperus sect. Leucocephali Cherm. ex Kük.). Cyperus sect. Leptostachyi (Boeck.) C.B. Clarke in J. Linn. Soc., Bot. 20: 282. 1883 ≡ C. [unranked] Leptostachyi Boeck. in Linnaea 36: 353. 1870 – Lectotype (designated here): Cyperus nitidulus Boeck. [= C. odoratus L. subsp. odoratus]. = Cyperus sect. Diclidium (Schrad. ex Nees) Griseb. This name is not based on C. leptostachyus Griff. [= C. ro­ tundus L.]. Böckeler included 19 species in this section; Kükenthal (1936) later divided them among nine of his sections in the subgenera Mariscus and Torulinium. Clarke (1883) published the name on sectional level including only C. ferax Rich. [= C. odoratus L. subsp. odoratus]. Böckeler (1870: 353) included this species in C. [unranked] Diclidium, but included C. nitidulus, a synonym of C. ferax in C. [unranked] Lepto­ stachyi. Cyperus sect. Diclidium (Schrad. ex Nees) Griseb. had already been published in 1864 with C. odoratus as type. Clarke (1883) clearly remarked on the relationship of the latter species with C. ferax, and definitely considered both taxa as belonging to the same section, writing: “C. odoratus, Vahl; … paullo differt spiculis 14–21-floris, sed mihi varietas videtur”. Cyperus sect. Leucocephali Cherm. ex Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 276. 1936 ≡ C. [unranked] Leptolepides Boeck. in Linnaea 35: 588. 1868 ≡ C. ([unranked] Leptolepides) [unranked] Capitati Boeck. in Linnaea 35: 588. 1868 – Type: Cyperus leucocephalus Retz. (type of C. sect. Leucocephali Art. 22.6 – lectotype of C. [unranked] Leptolepides and C. [unranked] Capitati designated here). Cyperus sect. Leucocephali was first published as a nomen nudum by Chermezon (1931), Kükenthal (1936) added a valid description. The characters of C. leucocephalus fit very well the description of C. [unranked] Leptolepides (e.g., “Inflorescentia saepissime capitato-contracta”). Böckeler (1868) distinguished two lower groups in C. [unranked] Leptolepides, i.e., C. [unranked] Capitati (incl. C. leucocephalus) and C. [unranked] Umbellati. Although these names are validly published under Art. 35.3, they do not have priority. Consequently, the correct name of this taxon at sectional level is C. sect. Leuco­ cephali Cherm. ex Kük. (1936) (Art. 11.4). 877 Larridon & al. • Names of subdivisions of Cyperus Cyperus sect. Limosi T.V. Egorova in Novosti Sist. Vyssh. Rast. 34: 27. 2002 – Type: Cyperus limosus Maxim. (Art. 22.6). Cyperus subg. Lipocarpha (R. Br.) T. Koyama in Quart. J. Taiwan Mus. 14: 163. 1961 ≡ Lipocarpha R. Br. in Tuckey, Narr. Exped. Zaire: 459. 1818 – Type: Cyperus lipocarpha T. Koyama [= Lipocarpha chinensis (Osbeck) J. Kern] (Art. 7.4). ≡ Lipocarpha R. Br. (see Huygh & al., 2010). Cyperus sect. Longispicati J.V. Suringar, nom. illeg. (see Cype­ rus sect. Distantes C.B. Clarke). Cyperus (subg. Mariscus) subsect. Longistyli Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 507. 1936 – Lectotype (designated here): Cyperus cyperoides (L.) Kuntze. We designate C. cyperoides as lectotype of this subsection as it is a common species and it fits very well to the description given by Kükenthal (1936). Cyperus (sect. Papyrus) [unranked] Lucidi Benth., Fl. Austral. 7: 256. 1878 – Type: Cyperus lucidus R. Br. (Art. 22.6). Bentham (1878) published this name without mention of rank. This epithet has not been adopted in any ranked subdivision of Cyperus. Cyperus sect. Luzuloidei (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 21: 115. 1884 ≡ C. [unranked] Luzuloidei Kunth, Enum. Pl. 2: 39. 1837 (‘Luzuloideae’) ≡ C. [unranked] Bracteati Boeck. in Linnaea 35: 549. 1868, nom. illeg. (Art. 52.1) – Type: Cyperus luzulae (L.) Retz. (type of C. sect. Luzuloidei Art. 22.6 – type of C. [unranked] Bracteati Art. 7.5). Kunth (1837) was the first to describe this group (as C. [unranked] Luzuloidei). Böckeler (1868) referred to the same group as C. [unranked] “Bracteati s. Luzuloidei ” but excluding several species. According to Art. 34.2 both of Böckeler’s names are validly published, but C. [unranked] Bracteati Boeck. is nomenclaturally superfluous and illegitimate (Art. 52.1). Cyperus sect. Mariscoidei Nees in Linnaea 9: 285. 1834 – Type: Cyperus mariscoides Nutt., nom. illeg. [= C. fili­ culmis Vahl] (Art. 22.6). Steudel (1854) incorrectly assigned this section to Kunth (1837: 84 as ‘Cyperus [unranked] Mariscoides’). Cyperus subg. Marisculus (Goetgh.) Lye in Lidia 3, 4: 132. 1994 ≡ Marisculus Goetgh., Bull. Jard. Bot. Natl. Belg. 47: 444. 1977 (Huygh & al., 2010) – Type: Alinula peteri (Kük.) Goetgh. & Vorster (Ascolepis peteri Kük.) (Art. 7.4) [≡ Cyperus microaureus Lye]. = Alinula J. Raynal (see Huygh & al., 2010). Cyperus sect. Mariscus (Vahl) Griseb., Fl. Brit. W. I.: 566. 1864 ≡ Mariscus Vahl, Enum. Pl. 2: 372. 1805, nom. cons. ≡ C. subg. Mariscus (Vahl) C.B. Clarke in J. Linn. Soc., Bot. 20: 296. 1883 ≡ M. subg. Mariscus, autonym automatically 878 TAXON 60 (3) • June 2011: 868–884 established by C.B. Clarke in Hooker, Fl. Brit. India 6: 619. 1893 (Art. 22.3) ≡ M. sect. Mariscus, autonym automatically established by C.B. Clarke in Hooker, Fl. Brit. India 6: 620. 1893 (Art. 22.3) ≡ M. ser. Mariscus, autonym automatically established by C.B. Clarke in Urban, Symb. Antill. 2: 39. 1900 (Art. 22.3) ≡ M. subsect. Mariscus, autonym automatically established by C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 103. 1908 (Art. 22.3) – Typ. cons.: Mariscus capillaris (Sw.) Vahl (Schoenus capillaris Sw.) [= Cyperus nanus Willd. var. nanus] (Art. 7.4). Endlicher (1836: 119) was the first to use Mariscus as an infrageneric taxon of Cyperus as C. [unranked] Mariscus (Vahl) Endl. In this group, Böckeler (1870: 372) distinguished several unnamed subgroups based on the inflorescence morphology from a single capitate inflorescence (group a), suborbiculate spikes (b), an umbel with sessile spikes (group b1), pedunculated spikes (b2), cylindrical spikes (c), simple umbel (c1), compound with sessile umbels (c2) and compound umbel with stalked umbels (c3). Cyperus subg. Micheliani (C.B. Clarke) Kukkonen in Ann. Bot. Fenn. 32, 3: 161. 1995 ≡ Scirpus sect. Micheliani C.B. Clarke in Hooker, Fl. Brit. India 6: 662. 1893 – Type: Scir­ pus michelianus L. (Art. 22.6) [≡ Cyperus michelianus (L.) Delile subsp. michelianus]. Clarke (1893) described only two species in S. sect. Micheliani, S. michelianus and S. kyllingioides (A. Rich.) Boeck. [= Kyllingiella microcephala (Steud.) R.W. Haines & Lye] the type of the genus Kyllingiella. In 1902, Clarke added a third species S. steudneri Boeck. [= Kyllinga ambigua Steud.]. Cyperus subg. Micromariscus Lye in Nordic J. Bot. 3: 215. 1983 – Type: Cyperus micromariscus Lye (Art. 22.6). Cyperus (subg. Juncellus) sect. Minuti Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 325. 1936 – Type: Cyperus minutus (C.B. Clarke) Kük, nom. illeg., non J.F. Gmel. (Art. 22.6). The type specimen of C. minutus, nom. illeg. clearly shows affinities to C. meeboldii Kük. and relatives which are treated by Kükenthal (1936) in C. sect. Rupestres which is characterised by a tendency towards monostigmatic gynoecia, distinctive obovoid nutlets and swollen culm bases. The exact status of C. minutus, nom. illeg. is unclear and needs further attention. Cyperus sect. Monocephali C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 310. 1901 (‘Monocephalae’) ≡ C. sect. Rupestres C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 96. 1908 – Type: Cyperus rupestris Kunth (lectotype of C. sect. Monocephali designated here – type of C. sect. Rupestres Art. 22.6). The characters of C. rupestris fit very well the description of C. sect. Monocephali, especially in having a constant single head of spikelets. Cyperus (subg. Pycreus) sect. Muricati Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 394. 1936, nom. superfl. TAXON 60 (3) • June 2011: 868–884 Larridon & al. • Names of subdivisions of Cyperus – Type: Cyperus muricatus Kük. [≡ Pycreus muricatus (Kük.) Napper] (Art. 22.6). When published, Cyperus sect. Muricati included the type of Pycreus sect. Tuberculati Cherm. (1919) and is therefore superfluous. However, Reynders & Goetghebeur (2010) misinterpreted Art. 52.3 in stating that the name C. sect. Muricati can be used when C. divulsus Ridl. [≡ Pycreus divulsus (Ridl.) C.B. Clarke] is excluded. Cyperus sect. Pennati (Kunth) J. Kern in Fl. Males., ser. 1, 7(3): 635. 1974, nom. superfl. ≡ C. [unranked] Pennati Kunth, Enum. Pl. 2: 79. 1837 – Type: Cyperus pennatus Lam. (Art. 22.6). = Cyperus (subg. Mariscus) sect. Turgiduli (C.B. Clarke) Kük. When published, Cyperus sect. Pennati included the type of C. sect. Turgiduli (C.B. Clarke) Kük. (1936) and is therefore superfluous. Cyperus sect. Nivei C.B. Clarke (see Cyperus sect. Hyme­ nolepides Nees). Cyperus (subg. Mariscus) sect. Phleoidei Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 426. 1936 – Type: Cyperus phleoides (Nees ex Kunth) H. Mann. (Art. 22.6). “Cyperus sect. Nudicaules Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 3. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. nudicaulis Poir. [= C. pectinatus Vahl]. Cyperus (subg. Mariscus) sect. Ochrocephali Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 537. 1936 – Type: Cyperus ochrocephalus C.B. Clarke, nom. illeg. [= C. rhynchospor­ oides Kük.] (Art. 22.6). Cyperus (“sect. Eucyperus”) [unranked] Ornati Benth., Fl. Austral. 7: 256. 1878 – Type: Cyperus ornatus R. Br. (Art. 22.6). Cyperus [unranked] Ornati includes the type of C. sect. Irioidei Nees (1834), which is the correct name at sectional rank. “Cyperus sect. Pandanophylli Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 27. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. pandanophyllum C.B. Clarke. Cyperus sect. Papyrus (Willd.) Thouars in Griseb., Fl. Brit. W. I.: 566. 1864 ≡ Papyrus Willd., Abh. Königl. Akad. Wiss. Berlin 1812–1813: 70. 1816 (Huygh & al., 2010) – Type: Cyperus papyrus subsp. antiquorum (Willd.) Kük. [= C. papyrus L. subsp. papyrus] (Art. 7.4). Kunth (1837: 64) was the first to treat Papyrus as an infrageneric taxon of Cyperus as C. [unranked] Papyrus (Willd.) Kunth (‘Papyri’). Cyperus subsect. Parvuli Kük. (see Kyllinga subsect. Parvulae (Kük.) Tang & F.T. Wang; Reynders & al., 2011). “Cyperus sect. Paucispiculati Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 17. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. paucispiculatus Cherm., nom. illeg. [≡ C. limiticola Larridon & Reynders]. Recently, Larridon & al. (2008) proposed a new name for C. paucispiculatus Cherm. since the latter is an illegitimate later homonym of C. paucispiculatus Boeck. [= Pycreus di­ vulsus (Ridl.) C.B. Clarke]. Cyperus sect. Pilosi (J.V. Suringar) Nakai in Bot. Mag. (Tokyo) 26: 187. 1912, nom. superfl. ≡ C. [unranked] Pilosi J.V. Suringar, Cyperus: 128. 1898 – Type: Cyperus pilosus Vahl (Art. 22.6). = Cyperus sect. Proceri (Kunth) C.B. Clarke. When published, Cyperus sect. Pilosi included the type of C. sect. Proceri (Kunth) C.B. Clarke (1883) and is therefore superfluous. Cyperus subsect. Pingues Kük. (see Kyllinga subsect. Pingues (Kük.) Tang & F.T. Wang; Reynders & al., 2011). Cyperus (subg. Mariscus) sect. Pinnati Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 451. 1936 – Lectotype (Kern, 1974): Cyperus angustatus R. Br. “Cyperus sect. Platyphylli Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 59. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. platyphyllus Roem. & Schult. Cyperus sect. Platystachyi (Kunth) C.B. Clarke (see Cyperus sect. Hymenolepides Nees). Cyperus sect. Polystachyi (C.B. Clarke) Kük., nom. illeg. (see Pycreus P. Beauv. ser. Pycreus; Reynders & al., 2011). Cyperus sect. Proceri (Kunth) C.B. Clarke in J. Linn. Soc., Bot. 20: 291. 1883 ≡ C. [unranked] Proceri Kunth, Enum. Pl. 2: 72. 1837 ≡ C. [unranked] Marginati Boeck. in Linnaea 35: 598. 1868 ≡ C. sect. Marginati (Boeck.) C.B. Clarke in J. Linn. Soc., Bot. 21: 146. 1884, nom. illeg. (Art. 52.1) ≡ C. subg. Rhizomati Nakai in Bot. Mag. (Tokyo) 26: 186. 1912 – Type: Cyperus procerus Rottb. (type of C. sect. Pro­ ceri Art. 22.6 – type of C. sect. Marginati Art. 7.5 – lectotype of C. subg. Rhizomati designated here). Kunth (1837) cited the species as C. procerus Vahl and Böckeler (1868) considered the Wallich 3355a material as mixed, partly being C. pilosus Vahl, and partly being C. heynei Boeck. (with reference to C. procerus Kunth). According to Art. 7.4 this has no influence on C. procerus Rottb. being the type of Kunth’s group of unspecified rank. Clarke (1884) and Suringar (1898) considered the descriptions and illustrations as sufficiently resembling each other and assigned them all to C. procerus Rottb. 879 Larridon & al. • Names of subdivisions of Cyperus Cyperus sect. Propinqui (C.B. Clarke) Kük. (see Pycreus sect. Propinqui C.B. Clarke; Reynders & al., 2011). Cyperus subg. Protocyperus Lye in Nordic J. Bot. 1: 57. 1981 – Type: Cyperus difformis L. = Cyperus subg. Pycnostachys C.B. Clarke (by lectotypification). According to Lye (1981a), his C. subg. Protocyperus includes all species of Cyperus with an eucyperoid culm-anatomy. This includes C. diffusus Vahl, the lectotype (Tucker, 1987) of C. subg. Pycnostachys C.B. Clarke (1893). Cyperus sect. Pseudanosporum C.B. Clarke in J. Linn. Soc., Bot. 21: 117. 1884 ≡ C. sect. Natantes C.B. Clarke in Hooker, Fl. Brit. India 6: 597. 1893, nom. illeg. (Art. 52.1) – Type: Cyperus platystylis R. Br. Cyperus sect. Pseudohaspani (‘Pseudo-Haspani ’) Nakai in Bot. Mag. (Tokyo) 26: 188. 1912 – Type: Cyperus pseudo­ haspan Makino [= C. tenuispica Steud.] (Art. 22.6). Nakai (1912) mentioned C. pseudohaspan as a synonym under C. flavidus Retz. [= Pycreus flavidus (Retz.) T. Koyama]. Cyperus sect. Pseudomariscus C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 99. 1908 (‘Pseudo­Mariscus’) – Lectotype (designated here): Cyperus schweinitzii Torr. The characters of C. schweinitzii fit very well the description of this section (Clarke, 1908). Clarke (1908) included C. sect. Pseudomariscus in C. subg. Choristachys as its species have subcaduceus spikelets, in contrast to the completely caduceus spikelets in Mariscus (which he treats as a separate genus). Cyperus (sect. Pycreus) [unranked] Pseudopycreus Boeck. (see Cyperus sect. Laevigati Kük.). Cyperus sect. Pseudopycreus (C.B. Clarke) Kük., nom. illeg. (see Kyllinga subg. Pseudopycreus C.B. Clarke; Reynders & al., 2011). Cyperus subg. Pterocyperus Peterm. (see Cyperus L. sect. Cyperus). Cyperus sect. Pulchrae C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 312. 1901 – Lectotype: Cyperus apricus Ridl. [C. semi­ trifidus Schrad.]. Cyperus sect. Pumili Kük. (see Pycreus sect. Pumili (Kük.) L.K. Dai; Reynders & al., 2011). TAXON 60 (3) • June 2011: 868–884 Cyperus sect. Pycreus (P. Beauv.) Griseb., Cyperus subg. Pycreus (P. Beauv.) J. Carey (see Pycreus P. Beauv. ser. Pycreus; Reynders & al., 2011). Cyperus sect. Pygmaei Cherm. ex Y.L. Chang in Fl. Pl. Herb. Chin. Bor.-Or. 11: 204. 1976 ≡ Juncellus sect. Caespitosi Nakai in Bot. Mag. (Tokyo) 26: 200. 1912 – Type: Cype­ rus pygmaeus Rottb. [≡ C. michelianus (L.) Delile subsp. pygmaeus (Rottb.) Asch. & Graebn.] (Art. 22.6). The epithet of the name Juncellus sect. Caespitosi Nakai (1912), which has the same type being the only species referred to in its protologue, has priority under Art. 11.4. However, at the same time Nakai (1912) also published a C. subg. Caespitosi in Cyperus with a different type, i.e., C. iria L. Consequently, Chang’s name is the correct name for this section when included in Cyperus. Cyperus (subg. Kyllinga) sect. Queenslandiella (Domin) J. Kern in Fl. Males., ser. 1, 7(3): 654. 1974 ≡ Queenslandiella Domin in Biblioth. Bot. 85, 4: 415. 1915 ≡ C. subg. Queens­ landiella (Domin) Govind. in Reinwardtia 9: 194. 1975 – Type: Queenslandiella hyalina (Vahl) Ballard (Art. 7.4) [≡ Cyperus hyalinus Vahl]. ≡ Queenslandiella Domin (see Huygh & al., 2010). Cyperus sect. Radiantes J.V. Suringar, Cyperus: 84. 1898 – Type: Cyperus radians Nees & Meyen ex Kunth (Art. 22.6). Cyperus sect. Remirea (Aubl.) T. Koyama in Quart. J. Taiwan Mus. 14: 162. 1961 ≡ Remirea Aubl., Hist. Pl. Guiane 1: 44. 1775 ≡ C. subg. Remirea (Aubl.) Lye in Nordic J. Bot. 3: 230. 1983 – Type: Remirea maritima Aubl. (Art. 7.4). ≡ Remirea Aubl. (see Huygh & al., 2010). The name Remirea maritima Aubl. has not been combined in Cyperus; Lye (1983) used the heterotypic synonym: Cyperus pedunculatus (R. Br.) J. Kern. Cyperus sect. Remotiglumi C.B. Clarke (see Cyperus sect. Subimbricati C.B. Clarke). Cyperus subg. Rhizomati Nakai (see Cyperus sect. Proceri (Kunth) C.B. Clarke). Cyperus sect. Rhizomatosi Kük. (see Pycreus sect. Rhizoma­ tosi (Kük.) J. Raynal; Reynders & al., 2011). Cyperus sect. Rotundi C.B. Clarke in Hooker, Fl. Brit. India 6: 614. 1893 – Type: Cyperus rotundus L. (Art. 22.6). = Cyperus L. sect. Cyperus (by lectotypification). “Cyperus sect. Pustulati Cherm.” in Arch. Bot. Mem. 4, Mém. 7: 19. 1931, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. pustulatus Vahl. Cyperus sect. Rupestres C.B. Clarke (see Cyperus sect. Mono­ cephali C.B. Clarke). Cyperus subg. Pycnostachys C.B. Clarke (see Cyperus sect. Diffusi (Kunth) C.B. Clarke). Cyperus sect. Serotini Kük., nom. illeg. (see Juncellus sect. Stoloniferi Nakai; Reynders & al., 2011). 880 TAXON 60 (3) • June 2011: 868–884 Cyperus (sect. Diffusi) [unranked] Simplices C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 98. 1908 (‘Dif­ fusae’) – Type: Cyperus simplex Kunth (Art. 22.6). This epithet has not been adopted in any ranked subdivision of Cyperus. Cyperus sect. Solubiles C.B. Clarke (see Cyperus sect. Cype­ rus). Cyperus subg. Sorostachys (Steud.) Lye in Nordic J. Bot. 3: 230. 1983 ≡ Sorostachys Steud., Syn. Pl. Glumac. 2: 71. 1854 (Huygh & al., 2010) – Type: Sorostachys kyllingioides Steud. (Art. 7.4). [= Cyperus pulchellus R. Br.]. “Cyperus sect. Sphacelati Cherm.” in Ann. Mus. Colon. Marseille, ser. 3, 10(1): 39. 1922, nom. nud. (Art. 32.1(d) and 36.1). This intended name was based on C. sphacelatus Rottb. Cyperus [unranked] Sphaerolepides Boeck. in Linnaea 35: 595. 1868 – Type: Cyperus sphaerolepis Boeck. (Art. 22.6). On first sight, Böckeler (1868) included only three species in C. [unranked] Sphaerolepides: C. iria L., C. eleusi­ noides Kunth [≡ C. nutans var. eleusinoides (Kunth) R.W. Haines] and C. nutans Vahl. However, Böckeler’s description of C. [unranked] Sphaerolepides ends in “Inflorescentia interdum (in C. spectabili—C. filiculmi—C. leucolepidi) ad capitulum solitarum redacta” but these species are only mentioned under his next group C. [unranked] Marginati. Under C. [unranked] Marginati, Böckeler (1868) also published the new species C. sphaerolepis Boeck. on which C. [unranked] Sphaerolepides was probably based. Most likely, these species were erroneously placed under C. [unranked] Marginati while editing Böckeler’s notes. Although C. [unranked] Sphaerolepi­ des is validly published under Art. 35.3, its epithet has not been adopted in any ranked subdivision of Cyperus. Cyperus sect. Spicati (Boeck.) C.B. Clarke in J. Linn. Soc., Bot. 20: 294. 1883 ≡ C. [unranked] Spicati Boeck. in Linnaea 36: 322. 1870 – Lectotype (designated here): Cyperus ligularis L. Originally, Böckeler (1870) included 21 species in C. [unranked] Spicati ; Clarke (1897) and Kükenthal (1936) later placed 15 of these species in C. sect. Thunbergiani. Cyperus (“sect. Eucyperus”) [unranked] Squarrosi Benth. (see Cyperus subg. Aristomariscus Lye). Cyperus (subg. Mariscus) sect. Strigosi Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 404. 1936 ≡ Mariscus ser. Multiflori C.B. Clarke in Urban, Symb. Antill. 2: 40. 1900 (‘Multiflorae’) ≡ M. sect. Multiflori (C.B. Clarke) C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 103. 1908 (‘Multiflorae’) ≡ M. (sect. Multiflori) subsect Oblongispiculosi C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 103. 1908 (‘Oblongi­spiculosae’) ≡ M. (sect. Multiflori subsect. Oblongispiculosi) ser. Neogei C.B. Larridon & al. • Names of subdivisions of Cyperus Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 104. 1908 (‘Neogeae’) – Type: Cyperus strigosus L. (type of C. sect. Strigosi Art. 22.6 – lectotype of M. sect. Multi­ flori, M. subsect. Oblongispiculosi and M. ser. Neogei designated here). Mariscus sect. Multiflori includes species with manyflowered spikelets in contrast to Clarke’s other Mariscus sections, which only have a few nutlets ripening in each spikelet. Clarke (1908) further divided this section into two subsections and two series based on the shape of the spikelets and their new vs. old world origin. Kükenthal (1936) used the epithet Strigosi, instead of Multiflori when he published a section with a similar circumscription in Cyperus. Cyperus ([unranked] Corymbosi) [unranked] Subaphylli Boeck. (see Cyperus sect. Corymbosi (Kunth) C.B. Clarke). Cyperus [unranked] Subexalati C.B. Clarke (see Cyperus sect. Compressi Nees). Cyperus sect. Subimbricati C.B. Clarke in Hooker, Fl. Brit. India 6: 612. 1893 (‘Subimbricatae’) ≡ C. sect. Remotiglumi C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 314. 1901 (‘Remoti­ glumae’) ≡ C. sect. Zollingeriani C.B. Clarke in Kew Bull. Addit. Ser. 8: 100. 1908 (‘Zollingerianae’) (Art. 22.6) ≡ C. sect. Subquadrangulares Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 127. 1935 – Lectotype: Cyperus zollingeri Steud. (lectotype of C. sect. Subimbricati designated by Kern (1974) – lectotype of C. sect. Remotiglumi and C. sect. Subquadrangulares designated here). Clarke (1893) included three species in C. sect. Subim­ bricati: C. macer C.B. Clarke, C. tegetum Roxb. [= C. pan­ gorei Rottb.] and C. zollingeri. Clarke (1908) placed all tree species in different sections: C. macer into C. sect. Brevifo­ liati, C. tegetum was put into the new C. sect. Tegetales and C. zollingeri in the new C. sect. Zollingeriani. Kükenthal (1935) included the first two species in C. sect. Brevifoliati, nom. illeg. and the third species in his new C. sect. Subquad­ rangulares. Cyperus sect. Subquadrangulares Kük. (see Cyperus sect. Subimbricati C.B. Clarke). Cyperus sect. Subulati C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 100. 1908 (‘Subulatae’) – Type: Cyperus subulatus R. Br. (Art. 22.6). Cyperus (subg. Pycreus) sect. Sulcati Kük., nom. illeg. non C. [unranked] Sulcati Boeck. (see Pycreus sect. Vestiti C.B. Clarke; Reynders & al., 2011). Cyperus sect. Tegetales C.B. Clarke in Kew Bull. Addit. Ser. 8: 100. 1908 – Type: Cyperus tegetum Roxb. [= C. pangorei Rottb.] (Art. 22.6). Cyperus sect. Tenelli C.B. Clarke (see Cyperus sect. Capitu­ ligeri Nees). 881 Larridon & al. • Names of subdivisions of Cyperus Cyperus (subg. Mariscus) sect. Tetragoni Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 486. 1936 – Type: Cyperus tetragonus Elliott (Art. 22.6). Cyperus sect. Textiles C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 312. 1901, nom. superfl. – Type: Cyperus textilis Thunb. (Art. 22.6). = Cyperus sect. Alternifolii (Kunth) C.B. Clarke. When published, Cyperus sect. Textiles included the type of C. sect. Alternifolii (Kunth) C.B. Clarke (1883) and is therefore superfluous. Cyperus (subg. Mariscus) sect. Thunbergiani (C.B. Clarke) Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 430. 1936 ≡ Mariscus sect. Thunbergiani C.B. Clarke in Dyer, Fl. Cap. 7: 186. 1897 – Type: Mariscus thunbergii (Vahl) Schrad. (Art. 7.4, Art. 22.6) [≡ Cyperus thunbergii Vahl]. Cyperus subg. Torulinium (Desv. ex Ham.) Kük. (see Cyperus subg. Diclidium (Schrad. ex Nees) C.B. Clarke). Cyperus sect. Tunicati C.B. Clarke (see Cyperus sect. Corym­ bosi (Kunth) C.B. Clarke). Cyperus (subg. Mariscus) sect. Turgiduli (C.B. Clarke) Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 473. 1936 ≡ Mariscus sect. Turgiduli C.B. Clarke in Hooker, Fl. Brit. India 6: 623. 1893 – Type: Mariscus albescens Gaudich. [= C. ja­ vanicus Houtt.]. Cyperus [unranked] Umbellati Boeck. (see Cyperus sect. Dis­ tantes C.B. Clarke). Cyperus sect. Umbellati (C.B. Clarke) Kük., nom. illeg. (see Mariscus sect. Umbellati C.B. Clarke; Reynders & al., 2011). Cyperus sect. Uncinati C.B. Clarke in Oliver, Fl. Trop. Afr. 8: 311. 1901 (‘Uncinatae’), nom. superfl. – Type: Cyperus uncinatus Poir. [= C. cuspidatus Kunth] (Art. 22.6). = Cyperus sect. Amabiles C.B. Clarke. When published, Cyperus sect. Uncinati included the type of C. sect. Amabiles C.B. Clarke (1893) and is therefore superfluous. Cyperus sect. Vaginati (Boeck.) Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 187. 1936, nom. superfl. ≡ C. [unranked] Vaginati Boeck. in Linnaea 35: 565. 1868 – Type: Cyperus vaginatus R. Br. (Art. 22.6). = Cyperus sect. Alternifolii (Kunth) C.B. Clarke. When published, Cyperus sect. Vaginati included the type of C. sect. Alternifolii (Kunth) C.B. Clarke (1883) and is therefore superfluous. The older name of unspecified rank does not have priority (Art. 35.3). Cyperus sect. Vegeti C.B. Clarke in Bull. Misc. Inform. Kew, Addit. Ser. 8: 98. 1908 (‘Vegetae’), nom. superfl. – Type: Cyperus vegetus Willd. [= C. eragrostis Lam.] (Art. 22.6). 882 TAXON 60 (3) • June 2011: 868–884 = Cyperus sect. Luzuloidei (Kunth) C.B. Clarke. When published, Cyperus sect. Vegeti included the type of C. sect. Luzuloidei (Kunth) C.B. Clarke (1883) and is therefore superfluous. Cyperus (subg. Mariscus) subsect. Vestiti Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 540. 1936 – Type: Cyperus vestitus Hochst. ex C. Krauss (Art. 22.6). Kükenthal (1936) created this subsection in C. sect. Bulbo­ caules C.B. Clarke (Kük.). Cyperus subsect. Vestiti Kük. (1936) is different from C. sect. Vestiti (C.B. Clarke) J. Kern, nom. illeg. (1974). Cyperus sect. Vestiti (C.B. Clarke) J. Kern, nom. illeg. (see Pycreus sect. Vestiti C.B. Clarke; Reynders & al., 2011). Cyperus (sect. Pseudohaspani) subsect. Virides Nakai in Bot. Mag. (Tokyo) 26: 189. 1912 – Type: Cyperus hakonensis Franch. & Sav. Cyperus sect. Viscosi C.B. Clarke in J. Linn. Soc., Bot. 21: 114. 1884 ≡ C. [unranked] Sulcati Boeck. in Linnaea 35: 547. 1868 ≡ C. [unranked] Glutinosi Boeck. in Linnaea 35: 547. 1868 ≡ C. sect. Glutinosi (Boeck.) Kük. in Engler, Pflanzenr. IV, 20 (Heft 101): 163. 1936 – Type: Cyperus viscosus Sw. [= C. elegans L. subsp. elegans] (type of C. sect. Viscosi Art. 22.6 – lectotype of C. [unranked] Glutinosi (≡ C. sect. Glutinosi) and C. [unranked] Sulcati designated here). = Cyperus sect. Elegantes C.B. Clarke. As Clarke (1884) is a local treatment of Cyperus (India), he only described C. viscosus and C. oxylepis Nees ex Steud. in C. sect. Viscosi. Kükenthal (1936) chose the epithet Gluti­ nosi when he treated Böckeler’s rankless group “C. [unranked] Sulcati s. Glutinosi Boeck.” at sectional rank. According to Art. 34.2 both of Böckeler’s names are validly published, but C. [unranked] Sulcati Boeck. is nomenclaturally superfluous and illegitimate (Art. 52.1). The older names of unspecified rank do not have priority (Art. 35.3). Böckeler (1868) included C. viscosus, but not C. elegans. Kükenthal (1936) included C. viscosus in synonymy under C. elegans. Cyperus subg. Xerocyperus Lye in Nordic J. Bot. 3: 214. 1983 – Type: Cyperus rubicundus Vahl. Cyperus sect. Zollingeriani C.B. Clarke (see Cyperus sect. Subimbricati C.B. Clarke). Cyperus (subg. Pycreus) [unranked] Zonati (see Pycreus ser. Zonati (C.B. Clarke) C.B. Clarke; Reynders & al., 2011). ACKNOWLEDGEMENTS Nicole Hanquart (Library, National Botanic Garden of Belgium) and the staff of the library of the Royal Botanic Gardens, Kew are gratefully acknowledged to help with tracing several historical publications. Many thanks also to the editor John McNeill (Royal Botanic TAXON 60 (3) • June 2011: 868–884 Garden, Edinburgh, Scotland, U.K.) and the reviewers for the careful and critical review of the manuscript. Specimens were studied from the following herbaria: B, BM, BR, GENT, K, P, UPS. This study financed by the Special Research Fund (BO5622, BO7418, BOF, Ghent University), and the Department of Biology, Ghent University. LITERATURE CITED Bentham, G. 1878. Flora australiensis, vol. 7. London: Reeve. Bentham, G. 1881. Notes on Cyperaceae. J. Linn. Soc., Bot. 18: 281–360. Böckeler, O. 1868. Die Cyperaceen des Königlichen Herbariums zu Berlin. 1. Theil. Die Cypereen, Scirpeen und Hypolytreen. Lin­ naea 35: 397–612. Böckeler, O. 1868–77. Die Cyperaceen des Königlichen Herbariums zu Berlin. Linnaea 35: 397–612, 36: 271–512, 691–768, 37: 1–142, 520–647, 38: 223–544, 39: 1–152, 40: 327–452, 41: 145–356. Böckeler, O. 1870. Die Cyperaceen des Königlichen Herbariums zu Berlin. 1. Theil. Die Cypereen, Scirpeen und Hypolytreen. Lin­ naea 36: 271–512. Britton, N.L. 1907. The sedges of Jamaica (Cyperaceae). Bull. Dept. Agric. Jamaica 5, Suppl. 1: 1–19. Chermezon, H. 1919. Révision des Cypéracées de Madagascar. Ann. Mus. Colon. Marseille, ser. 3, 7(2): 29–87. Chermezon, H. 1922. Révision des Cypéracées de Madagascar. Ann. Mus. Colon. Marseille, ser. 3, 10(1): 1–62. Chermezon, H. 1931. Les Cypéracées de Haut-Oubangui. Arch. Bot. Mem. 4, Mém. 7: 1–56. Chermezon, H. 1937. Cypéracées. Pp. 1–335 in: Humbert, H. (ed.), Flore de Madagascar, vol. 29. Tananarive: Imprimerie officièle. Clarke, C.B. 1883. Cyperus. Pp. 279–296 in: Baker, J.G. (ed.), Contributions to the flora of Madagascar. — Part 3. J. Linn. Soc., Bot. 20: 237–304. Clarke, C.B. 1884. On the Indian species of Cyperus; with remarks on some others that specially well illustrate the subdivisions ot the genus. J. Linn. Soc., Bot. 21: 1–202, pl. 1–4. Clarke, C.B. 1893. Cyperaceae. Pp. 585–672 in: Hooker, J.D. (ed.), The flora of British India, vol. 6, part 19. London: Reeve. Clarke, C.B. 1897. Cyperaceae. Pp. 149–192 in: Thiselton-Dyer, W.T. (ed.), Flora capensis, vol. 7. London: Reeve. Clarke, C.B. 1900. Cyperaceae. Pp. 8–169 in: Urban, I. (ed.), Symbolae antillanae, vol. 2, part 1. Leipzig: Fratres Borntraeger. Clarke, C.B. 1901–02. Cyperaceae. Pp. 266–524 in: Oliver, D. (ed.). Flora of tropical Africa, vol. 8. London: Reeve. Clarke, C.B. 1908. New genera and species of Cyperaceae. Bull. Misc. Inform. Kew, Addit. Ser. 8: 1–196. Druyts-Voets, E. 1970. Types van stengel- en bladstrukturen in het genus Cyperus L. Natuurw. Tijdschr., Antwerp. 52: 28–49. Endlicher, S.F.L. 1836. Genera plantarum secundum ordines naturales disposita, vol. 2. Vienna: Fr. Beck. Goetghebeur, P. 1989. Studies in Cyperaceae 9. Problems in the lectotypification and infrageneric taxonomy of Cyperus. Bull. Soc. Roy. Bot. Belgique 122: 103–114. Goetghebeur, P. 1998. Cyperaceae. Pp. 141–190 in: Kubitzki, F., Huber, H., Rudall, P.J., Stevens, P.S. & Stützel, T. (eds), The families and genera of vascular plants, vol. 4. Berlin: Springer. 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. Govaerts, R., Simpson, D.A., Goetghebeur, P., Wilson, K.L., Egorova, T. & Bruhl, J. 2010. World checklist of Cyperaceae. The Board of Trustees of the Royal Botanic Gardens, Kew. http:// www.kew.org/wcsp/monocots/ (accessed 20 May 2010). Grisebach, A.H.R. 1846. Spicilegium florae Rumelicae, vol. 2. Brunswick: F. Vieweg et filium. Larridon & al. • Names of subdivisions of Cyperus Grisebach, A.H.R. 1864. Flora of the British West Indian Islands. London: Reeve. Haines, R.W. & Lye, K.A. 1983. The sedges and rushes of East Africa. Nairobi: East African Natural History Society. Hooper, S.S. 1985. Cyperaceae. Pp. 1682–1710 in: Meikle, R.D. (ed.), Flora of Cyprus, vol. 2. Kew: Bentham-Moxon Trust. Huygh, W., Larridon, I., Reynders, M., Muasya, A.M., Govaerts, R., Simpson, D.A. & Goetghebeur, P. 2010. Nomenclature and typification of names of genera and subdivisions of genera in Cy­ pereae (Cyperaceae): 1. Names of genera in the Cyperus clade. Taxon 59: 1883–1890. Jaubert, H.F. & Spach, E. 1844. Illustrationes plantarum orientalium, ou Choix de plantes nouvelles ou peu connues de l’Asie occiden­ tale. Paris: Roret. Kern, J.H., 1974. Cyperaceae. Pp. 435–753 in: Van Steenis, C.G.G.J. (ed.), Flora Malesiana, ser. 1, vol. 7, part 3. Leiden: National Herbarium. Koyama, T. 1961. Classification of the family Cyperaceae 3. Quart. J. Taiwan Mus. 14: 159–194. Koyama, T. 1985. Cyperaceae. Pp. 125–405 in: Dassanayake, M.D. & Fosberg, F.R. (eds), A revised handbook to the flora of Ceylon, vol. 5. New Delhi: Amerind Publishing. Kükenthal, G. 1935–36. Cyperaceae—Scirpoideae—Cypereae. Pp. 1–671 in: Engler, A. (ed.), Das Pflanzenreich, vol. 4, 20. Stuttgart: Engelmann (J. Cramer). Kunth, C.S. 1837. Enumeratio plantarum, vol. 2. Stuttgart & Tübingen: Sumtibus J.G. Cottae. Larridon, I., Reynders, M. & Goetghebeur, P. 2008. Cyperus limiti­ cola, a new name for a Madagascan Cyperus (Cyperaceae). Novon 18: 187–188. Larridon, I., Reynders, M., Huygh, W., Bauters, K., Van de Putte, K., Muasya, A.M., Boeckx, P., Simpson, D.A., Vrijdaghs, A. & Goetghebeur P. In press. Affinities in C3 Cyperus lineages (Cyperaceae) revealed using molecular phylogenetic data and carbon isotope analysis. Bot. J. Linn. Soc. Linnaeus, C. 1753. Species plantarum. Stockholm: Impensis Laurentii Salvii. Lye, K.A. 1981a. Studies in African Cyperaceae 18. Tho new subgenera of Cyperus. Nordic J. Bot. 1: 57–61. Lye, K.A. 1981b. Studies in African Cyperaceae 19. The genera Ano­ sporum Nees and Sorostachys Steudel. Nordic J. Bot. 1: 186–191. Lye, K.A. 1981c. Studies in African Cyperaceae 20. New taxa and combinations in Pycreus Beauv. Nordic J. Bot. 1: 617–622. Lye, K.A. 1982. Studies in African Cyperaceae 21. New taxa and combinations in Kyllinga Rottb. Nordic J. Bot. 1: 741–747. Lye, K.A. 1983. Studies in African Cyperaceae 25. New taxa and combinations in Cyperus L. Nordic J. Bot. 3: 213–232. Lye, K.A. 1992. The history of the genus Mariscus (Cyperaceae). Lidia 3: 37–72. McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Hawksworth, D.L., Marhold, K., Nicolson, D.H., Prado, J., Silva, P.C., Skog, J.E., Wiersema, J.H. & Turland, N.J. (eds). 2006. International code of botanical nomenclature (Vienna Code): Adopted by the Seventeenth International Botanical Congress Vienna, Austria, July 2005. Regnum Vegetabile 146. Ruggell, Liechtenstein: Gantner. Metcalfe, C.R. 1971. Anatomy of the Monocotyledons, vol. 5, Cypera­ ceae. Oxford: Clarendon Press. Muasya, A.M., Simpson, D.A. & Chase, M.W. 2002. Phylogenetic relationships in Cyperus L. s.l. (Cyperaceae) inferred from plastid DNA sequence data. Bot. J. Linn. Soc. 138: 145–153. Muasya, A.M., Simpson, D.A. & Smets, E. 2006. Isolepis tenella, a new combination in Cyperaceae. Novon 16: 89–90. Muasya, A.M., Simpson, D.A. & Smets, E. 2007. Isolepis levynsiana, a new name for Cyperus tenellus (Cyperaceae). Novon 17: 59. Muasya, A.M., Simpson, D.A., Verboom, G.A., Goetghebeur, P., Naczi, R.F.C., Chase, M.W. & Smets, E. 2009a. Phylogeny of 883 Larridon & al. • Names of subdivisions of Cyperus Cyperaceae based on DNA sequence data: Current progress and future prospects. Bot. Rev. 75: 2–21. 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. Bot. Rev. 75: 52–66. Nakai, T. 1912. Cyperaceae-Cyperinae Japono-Coreanae. Bot. Mag. (Tokyo) 26: 185–207. Nees von Esenbeck, C.G. 1834. Uebersicht der Cyperaceengattungen. Linnaea 9: 273–308. Opiz, F.M. 1852. Seznam Rostlin Kveteny Ceske. Prague. Raynal, J. 1973. Notes cypérologiques: 19. Contribution à la classification de la sous-famille des Cyperoideae. Adansonia, ser. 2, 13(2): 145–171. Reynders M. & Goetghebeur P. 2010. Reestablishment of Pycreus sect. Tuberculati Cherm. Blumea 55: 226–230. 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 Cy­ pereae (Cyperaceae): 3. Names of subdivisions in segregate genera of Cyperus. Taxon 60: 885–895. Rikli, M. 1895. Beiträge zur vergleichenden Anatomie der Cyperaceen mit besonderer Berücksichtigung der inneren Parenchymscheide. Jahrb. Wiss. Bot. 27: 485–580. Schischkin, B.K. 1935. Cyperaceae and Juncaceae. Pp. 1–578 in: Komarov, V.L. (ed.), Flora unionis rerumpublicarum sovieticarum socialisticarum, vol. 3. Leningrad: Institutum Botanicum Academiae Scientiarum URSS. Simpson, D.A. 1990. A revision of Cyperus sect. Leucocephali. Kew Bull. 45: 485–501. 884 View publication stats TAXON 60 (3) • June 2011: 868–884 Simpson, D.A. & Koyama, T. 1998. Cyperaceae. Pp. 345–387 in: Santisuk, T. & Larsen, K. (eds), Flora of Thailand. Bangkok: Royal Forest Department. Simpson, D.A., Muasya, A.M., Alves, M., Bruhl, J.J., Dhooge, S., Chase, M.W., Furness, C.A., Ghamkhar, K., Goetghebeur, P., 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. Pp. 72–83 in: Columbus, J.T., Friar, E.A., Porter, J.M., Prince, L.M. & Simpson, M.G. (eds.), Monocots: Comparative biology and evolution: Poales. Aliso 23. Claremont: Rancho Santa Ana Botanic Garden. Soros, C.L. & Bruhl, J.J. 2000. Multiple evolutionary origins of C4 photosynthesis in the Cyperaceae. Pp. 629–636 in: Wilson, K.L. & Morrison, D.A. (eds.), Monocots: Systematics and evolution. Melbourne: CSIRO. Steudel, E.G. von. 1854–1855. Synopsis plantarum glumacearum, vol. 2, Cyperaceae. Stuttgart: Metzler. Strid, A. 1974. On the typification of Bobartia L. Taxon 23: 422–424. Suringar, J.V. 1898. Het geslacht Cyperus. Leeuwarden: Hugo Suringar. Tucker, G.C. 1983. The taxonomy of Cyperus (Cyperaceae) in Costa Rica and Panama. Syst. Bot. Monogr. 2: 1–85. Tucker, G.C. 1987. The genera of Cyperaceae in the SE United States. J. Arnold Arbor. 68: 361–445. Van der Veken, P. 1965. Contribution à l’embryographie systématique des Cyperaceae-Cyperoideae. Bull. Jard. Bot. État Brux­ elles 35: 285–354. Väre, H. & Kukkonen, I. 2006. Typification of names in Cyperus section Arenarii (Cyperaceae). Nordic J. Bot. 24: 279–294