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What is a genus in Cypereae: phylogeny, character homology assessment and generic circumscription in Cypereae.


Taxa included in tribe Cypereae are annual or perennial herbs that vary in stature from minute to 5 m tall. Leaves generally have well-developed blades, but are reduced to lobes in some species; there also may be a ligule. Inflorescences are capitate or anthelate. They all have hermaphrodite, trimerous flowers, with each subtended by a papery glume. Glumes are spirally or distichously arranged in the spikelets, apart from some reduced species in which the arrangement is obscure.

Generic classification in tribe Cypereae and subfamily Cyperoideae dates back to Linneaus (1753), who described the genera Scirpus and Cyperus to include all species of Cyperaceae with bisexual flowers, and distinguished by the spiral versus distichous glume arrangement in Scirpus and Cyperus respectively. The broad circumscription of Scirpus, based on common and widespread characters, resulted in a heterogeneous assemblage which was treated by subsequent workers as one genus (e.g. Boeckeler, 1870; Clarke, 1894, 1898, 1902; Hitchcock et al., 1969) or split into a number of smaller genera (e.g. Brown, 1810; Raynal 1973; Wilson, 1981; Goetghebeur, 1998).

Classification of genera into tribes in Cyperoideae has differed widely among authors, depending on which character(s) were emphasized. Therefore, there is a need to revise generic and tribal circumscriptions and especially incorporate new evidence from morphology and DNA sequence data.

Taxonomic History of Tribe Cypereae

Tribal concepts in Cyperoideae have varied over the years. A large number of legitimate tribal names have been published in Cyperoideae, including Cypereae, Scirpeae, Fuireneae, Ficinieae, Schoenoplecteae, Abildgaardieae, Lipocarpheae, and Eleocharideae (Goetghebeur, 1985). Cypereae and Scirpeae have been the most frequently used tribal names (e.g. Haines & Lye, 1983; Bruhl, 1995; Goetghebeur, 1998). The main difference has traditionally been that Scirpeae have spirally arranged glumes whereas in Cypereae glumes are two-ranked (e.g. Lye, 1971). However, this tribal classification has resulted in genera such as Oxycaryum and Isolepis being classified in Scirpeae even though these genera show closer affinity to Cyperus, as pointed out by Raynal (1973).

Embryological data (e.g. Van der Veken, 1965; Haines & Lye, 1971, 1976, 1983; Raynal, 1973, 1977; Wilson, 1981; Goetghebeur, 1996, 1998; Bruhl, 1995) have contributed significantly to generic and tribal circumscription in Cyperoideae. Heterogeneous Scirpus sensu lato has embryo types characteristic of Cyperus, Carex, Bulbostylis, Fimbrislylis, Schoenus and Schoenoplectus, whereas Cyperus has mainly the Cyperus-type embryo. Based on the interpretation that several genera could have the same type of embryo but a single genus should have only a single type of embryo, Scirpus sensu lato was split into several genera. Currently (Geotghebeur, 1998), tribes in Cyperoideae are classified to include genera sharing a single embryo type. For example, Cypereae have the Cyperus-type embryo and the similar Ficinia-type, whereas Scirpeae sensu stricto have only the Fimbristylis-type embryo.

Two recent classifications of Cyperoideae, based predominantly on morphological data, have differed in placement of genera in Cypereae. Goetghebeur (1998) classified all taxa characterised by Cyperus-type embryo in Cypereae, whereas Bruhl (1995) placed genera having spiral glume arrangement (i.e. Isolepis, Ficinia, Desmoschoenus, Scirpoides, Kyllingiella, Oxycaryum) in Scirpeae. Hellmuthia, bearing spirally arranged glumes but having an additional pair of scales in flowers subtended by the most proximal glumes, was placed in Scirpeae by Bruhl (1995) but in Chrysitricheae by Goetghebeur (1998). Bruhl (1995) did not recognise the tribes Eleocharideae and Fuireneae (sensu Goetghebeur, 1998), but included these taxa in Scirpeae.

Phylogeny, Character Homology Assessment and Generic Circumscription

Phylogenetic Relationship Based on Molecular Data

Over the last ten years, molecular systematic data have been used in the classification of ranks above family (e.g. APG, 2003) and in supraqeneric classification within Cyperaceae (Muasya et al., 1998, 2000a; Simpson et al., 2007). Studies on Cyperoideae have targeted phylogenetic relationships in Scirpeae (Muasya et al., 2000b, Dhooge et al., 2003) and Abildgaardieae (Ghamkhar et al., 2007), and focussed on genera Eleocharis (Roalson & Friar, 2000; Yano et al., 2004), Isolepis (Muasya et al., 2001a), Cyperus sensu lato (Muasya et al., 2002) and Schoenoplectus (Yano & Hoshino, 2005).

Total DNA was extracted from leaves or culms collected in the field or from herbarium specimens (Table 1). DNA extraction, amplification and sequencing were performed according to published procedures (e.g. Muasya et al., 2001a, 2002), and the resulting sequences aligned manually. We present and discuss here results of a maximum parsimony analysis of representatives of 18 of the 19 genera in Cypereae recognised by us; no material of the monotypic genus Ascopholis was available. The DNA data matrix (rbcL gene, rps16 intron, trnL intron and trnL-F intergenic spacer) comprises 3,721 characters among which 625 are potentially parsimony-informative. The matrix was analysed using the heuristic algorithm in PAUP* (Swofford, 2002), random addition for 10,000 replicates with tree-bisection-reconnection (TBR). Bootstrap analysis was performed for 1,000 replicates under maximum parsimony criterion (random taxon addition, twin replicates, TBR).

The strict consensus tree generated from the maximum parsimony analysis is presented in Fig. 1, with the bootstrap values for the various branches mapped. Cypereae are resolved into the Cyperus and Ficinia clades. The Ficinia clade comprises Scirpoides, Hellmuthia, Isolepis, Ficinia, Desmoschoenus and two Scirpus species (S. falsus and S. ficinioides). The Cyperus clade has Cyperus sensu stricto as the core genus, in which the thirteen derived genera (Alinula, Androtrichum, Ascolepis, Courtoisina, Kyllinga, Kyllingiella, Lipocarpha, Oxycaryum, Pycreus, Queenslandiella, Remirea, Sphaerocyperus, and Volkiella) are embedded.

Assessment of Morphological Character Homology

Using the DNA phylogenetic framework (Fig. 1), we evaluate the homology of key morphological characters used in classification of the Cypereae. The morphological characters are manually plotted on the DNA topology, majority of characters can be unambiguously reconstructed on the phylogeny. Ascopholis, a monotypic genus restricted to India (Goetghebeur, 1998), has not been included in this study due to unavailability of material. Generic status of Ascopholis is not accepted by all, and it has been suggested to be conspecific to the widespread Cyperus mollipes (C. B. Clarke) K. Schum (Govaerts et al., 2007).

Mature Embryo Morphology

Cypereae are characterised by the presence of a Cyperus-type embryo (Van der Veken, 1965; Haines & Lye, 1971, 1976; Raynal 1973; Wilson, 1981; Goetghebeur, 1985). In the Ficinia clade, species of Ficinia have a Ficinia-type embryo which is similar to Cyperus-type, but Isolepis, Hellmuthia and Scirpoides have a typical Cyperus-type embryo (Van der Veken, 1965; Haines & Lye, 1971). The embryo type in Scirpus falsus and S. ficinioides has not been studied, mainly because mature nutlets were not available.


Concepts of mature embryo morphological states are subject to individual interpretation of homology, and it may be difficult to distinguish similar embryo types in some cases. For example, Isolepis humillima, placed in Isolepis due to the presence of spiral glume arrangement, has been interpreted as having an embryo similar to Scirpoides (Wilson, 1981). The phylogenetic position of this taxon in molecular analyses is within Schoenoplectus subgen. Actaeogeton, a group possessing a Schoenoplectus-type embryo. The mature embryo in Cypereae is less complex when compared to state is the sister tribe Fuireneae (Schoenoplectus type), hence our study does not support Juguet's contention (as reported in Raynal, 1973) that the embryogeny of Cypereae is very evolved compared to the rest of the family.

Annual Versus Perennial Life Form

Annual and perennial growth forms are observed among members of tribe Cypereae (Haines & Lye, 1983; Goetghebeur, 1998; Table 2). In the Ficinia clade, an annual life form has evolved only in Isolepis (which also has some perennial species) whereas all other taxa are perennial. In the Cyperus clade, an annual life form is exclusively found in Courtoisina, Queenslandiella and Alinula; a predominantly perennial life form is observed in Oxycaryum, Kyllingiella, Remirea, Sphaerocyperus, Kyllinga and Ascolepis); while both annual and perennial life forms are recorded in Cyperus sensu stricto, Pycreus and Lipoearpha.

Glume Arrangement

Spiral glume arrangement is a plesiomorphic state in Cyperoideae (Muasya et al., 2001b). In Cypereae (Table 2), the Ficinia clade has predominantly a spiral glume arrangement, except in few species of Ficinia (e.g. F. distans and F. angustifolia) and Isolepis (I. levynsiana and I. venustula). In the Cyperus clade, distichous glume arrangement is usual especially in Androtrichum, Cyperus sensu stricto, Courtoisina, Pycreus, Kyllinga, Queenslandiella, Sphaerocyperus, Remirea, and Volkiella. Oxycaryum, Kyllingiella and Alinula have a spiral glume arrangement, while the spikelet is too reduced in Ascolepis and Lipocarpha for interpretation of glume arrangement (Goetghebeur, 1998). Distichous glume arrangement has evolved more than once in Cypereae, occurring in both the Ficinia and Cyperus clades, and is therefore not unique in Cyperus sensu stricto. The unreliability of distichous arrangement as a diagnostic character has been previously shown (e.g. Raynal, 1973), and evident from our study where taxa with the Cyperus-like distichous glume arrangement (e.g. Isolepis levynsiana) are resolved in the Ficinia clade.

Hypogynous Scales

Hypogynous scales, a character considered plesiomorphic in Cyperoideae, are found in Scirpeae, Fuireneae, Eleocharideae, Dulichieae, and Schoeneae but are absent from Abildgaardieae and Cypereae (Goetghebeur, 1998). Scirpus falsus and S. ficinioides, resolved in Cypereae in molecular phylogenetic analyses (Fig. 1), have bristle-like perianth segments. Similar perianth segments, some well developed and others rudimentary, have been observed in Ficinia material (Muasya et al., unpublished results).

Some florets in Hellmuthia have two scales, which have been suggested to be homologous to scales in Mapanioideae (Haines & Lye, 1976; Goetghebeur, 1998). Recent floral ontogenetic studies (Vrijdaghs et al., 2006) have revealed an adaxially situated third scale in some proximal flowers in spikelets of Hellmuthia, and these are interpreted to be perianth segments and not glumes of reduced florets as in Mapanioideae. Hellmuthia is resolved in the DNA phylogeny among the Ficinia clade and closely related to Scirpus falsus and S. ficinioides.


The gynophore in Cypereae, formed by the development of the hypogynous stalk, is characterised by a lobed cup that envelops the basal part of the nutlet (Vrijdaghs et al., 2005). This structure is absent from the rest of Cyperoideae except for Ficinia, in which variation is observed in size and shape of the gynophore. However, some Ficinia species lack a gynophore, while on the other hand some Isolepis species (e.g. I. marginata) have a rudimentary gynophore (Clarke, 1898; Levyns, 1950; Muasya et al., 2000c, 2001a). A gynophore is present in Alinula lipocarphoides, a taxon previously described in Ficinia and later transferred to Alinula (Kukenthal, 1936; Raynal, 1977), here resolved in the Cyperus clade as sister to Lipocarpha.

Kranz Anatomy

As in most angiosperms families, the plesiomorphic photosynthetic system in most of Cyperaceae is [C.sub.3] type. Multiple origins of Kranz anatomy are recorded in several lineages including Rhynchospora, Eleocharis, Fimbristylis and Cyperus (Raynal, 1973; Estelita, 1993; Goetghebeur, 1998; Soros & Bruhl, 2000; Muasya et al., 2002; Bruhl & Wilson, 2007). Among Cypereae, Kranz anatomy has evolved once among Cyperus clade and is recorded in Fig. 1 between Cyperus cuspidatus to Alinula lipocarphoides. Bruhl & Wilson (2007) erroneously reported Volkiella to be [C.sub.3], while in the supporting references they show isotopic carbon reading (-13.6) which is typical for [C.sub.4].

Samples of Alinula paradoxa and Lipocarpha rehmannii, reported to be Ca (Stock et al., 2004), might have been based on wrongly identified material, especially since there are four other records as [C.sub.4] for L. rehmannii (Bruhl and Wilson, 2007), and recent carbon isotope studies have confirmed other samples of these taxa to be [C.sub.4] (Muasya, unpublished results).

Inflorescence Morphology

Inflorescence morphology varies greatly in Cypereae. The basic inflorescence has spikelets in a panicle (Raynal, 1971), which is often modified into an anthela or contracted into a capitate head, spike or reduced to a single spikelet (Goetghebeur, 1998). In Cyperus, Ca taxa tend to have the spikelets arranged in digitate clusters, which is one of the few morphological characters to distinguish the [C.sub.3] and [C.sub.4] taxa (which are usually spicately arranged), apart from those species that have the inflorescence reduced to a head (Goetghebeur, 1998). Kukenthal (1935-1936) used this (only partly correctly) to subdivide his subgenus 'Eu-cyperus', while Raynal (1973) also noted this (as not being a simple dividing character) particularly in discussing the origins of the 'Mariscus' group of species.

Spikelets in a majority of Cypereae have many flowers. Several genera (e.g. Lipocarpka, Aseolepis, Alinula) have pseudo-spikelets, in which spikelets are reduced to single flowers (glumes lost) arranged in cones, each single-flower spikelet subtended by a glume-like bract. The resulting cone resembles a spikelet (Haines & Lye, 1983; Goetghebeur & Vorster, 1988) hence the use of the term 'pseudo-spikelet'.

Elongation of Filaments

Stamen filaments in most members of Cyperoideae are nearly as long as the glumes and inconspicuous after anthesis. Androtriehum trigynum and A. giganteum have filaments strongly elongating after anthesis, giving the inflorescence a cotton-like look. Such elongation of filaments is not observed in any other species in Cyperoideae.

Dispersal unit

Nutlets (also called achenes by some authors, e.g. Goetghebeur, 1998) in members of Cypereae are dispersed singly or together with elongated filaments, one to a few glumes, or parts of the spikelet axis, or even as complete spikelets (Kukenthal, 1935-1936; Raynal, 1973; Haines & Lye, 1983; Goetghebeur, 1998; Table 2). Courtoisina, Queenslandiella, Kyllinga, Remirea, Sphaeroeyperus, Lipocarpha, and Ascolepis have spikelets dispersing as intact units, whereas all taxa in the Ficinia clade, Kyllingiella, Pycreus, Oxycaryum, and Remirea have nutlets dispersed singly. Cyperus has nutlets dispersed either singly or as whole spikelets or variants thereof (notably in Cyperus odoratus).

Nutlet Orientation

Two kinds of nutlet orientation are observed in Cypereae (Table 2). Dorsiventral nutlet orientation is the most common and plesiomorphic state (Kukenthal, 1935-1936; Goetghebeur, 1998; Muasya et al., 2001b). Within Cypereae and Cyperaceae, species with distigmatic styles and dorsiventrally compressed nutlets are observed. Only the genera Kyllinga, Pycreus, and Queenslandiella have lateral nutlet orientation with distigmatic styles and laterally compressed nutlets.

Generic Circumscription

Cypereae are defined here as including all taxa sharing the Cyperus-type of embryo. We expand the tribal circumscription to include characters states such as the occasional presence of floral scales and bristle-like perianth segments, observed in the Ficinia clade.

The Ficinia Clade

Taxa in this clade have a predominantly spiral glume arrangement, but note the presence of distichous glume arrangement in Ficinia and Isolepis. All the genera share ficinioid morphology, e.g. tufted perennials, spiral glume arrangement, and have a center of diversity in the Cape floristic region of South Africa (Goetghebeur, 1998; Archer, 1998; Muasya & Simpson, 2002; Muasya, 2005). The individual genera are diagnosed by a combination of several characters (Table 2), the most notable being the presence of a gynophore and ligule in Ficinia (including Desmoschoenus), presence of two or three scales in the lower florets in Hellmuthia, and perennial growth form and spiral glume arrangement in Scirpoides, whereas Isolepis includes predominantly annual species with a spiral glume arrangement. Two annual species (Isolepis leucoloma and I. levynsiana) with distichous glumes previously described in Cyperus have been transferred to Isolepis, based on morphological and molecular data (Archer, 1998; Muasya et al., 2006, 2007).

There is overlap in generic limits between Isolepis and Ficinia as presently recognised, whereas Desmoschoenus is embedded in Ficinia (Fig. 1). An annual species with rudimentary gynophore described as Isolepis (I. marginata) is resolved in DNA analysis as more closely related to Ficinia. Desmoschoenus and Sickmannia (Ficinia radiata), taxa with a gynophore but with additional unique features, have been recognised as distinct from Ficinia. Phylogenetic results presented here (Fig. 1) show that these taxa are embedded in Ficinia, and should be recognized as members of Ficinia. Sickmannia has already been recognised as Ficinia (F. radiata) in recent treatments (Goetghebeur, 1998; Archer, 2000), whereas Desmoschoenus spiralis, a New Zealand endemic growing in the same coastal habitat as Ficinia nodosa, has no name in Ficinia. More studies are in progress to resolve relationships in the Ficinia clade.

Two of the Scirpus species, S. falsus and S. ficinioides from southern Africa, have the gross morphology of the Ficinia clade, including perennial habit, scapose culms, pseudolateral inflorescences, and spiral glumes. Presence of perianth segments has been used to include these taxa in Scirpus (e.g. Kunth, 1837; Clarke, 1898; Gordon-Gray, 1995) even though typical Scirpus has paniculate inflorescences and nodded culms. So far no embryo studies have been done on these taxa, and attempts to locate appropriate material have not been successful as the taxa rarely produce mature nutlets. Phylogenetic studies resolve these taxa as sister to Hellmuthia (Fig. 1), a pattern that suggests evolution from a southern African ancestor, unlike Scirpus, which is Holarctic. A new genus should be erected to include these two taxa, and more studies are in progress to formalise the recognition of this genus.

The Cyperus Clade

Genera in the Cyperus clade are circumscribed by a combination of morphological characters including spikelet morphology, unit of dispersal, and nutlet orientation (Table 2). Although these genera can be grouped into [C.sub.3] and [C.sub.4] anatomical types, there are few observable gross morphological characters to separate the species of Cyperus sensu stricto with the two kinds of anatomy.

Among [C.sub.3] genera, Androtrichum is diagnosed by the presence of elongated stamen filaments that are persistent and dispersed with the nutlets. However, the two taxa, A. giganteum and A. trigynum, are not sister (Fig. 1) and their shared character state, presence of elongated filaments, may be a parallel adaptation to dispersal in swampy coastal dunes. Kyllingiella and Oxycaryum, previously classified in Scirpeae (e.g. Bruhl, 1995), have a spiral glume arrangement unlike [C.sub.3] species of Cyperus sensu stricto, which have a distichous glume arrangement (Lye, 1971; Haines & Lye, 1978). Courtoisina has similar morphology to [C.sub.3] species of Cyperus, but the whole spikelet is dispersed intact.

The [C.sub.4] genera include a number that are monotypic or with few species (i.e. Queenslandiella, Sphaerocyperus, Remirea, Volkiella, and Alinula), which are separated from the larger genera by a combination of characters. Among the clearly recognizable larger genera are Kyllinga, and Pycreus (together with monotypic Queenslandiella), which have laterally flattened nutlets. Alinula, Volkiella, Ascolepis, and Lipocarpha have highly reduced spikelets. The [C.sub.4] species of Cyperus sensu stricto have spikelets comprising more than one floret and dorsiventrally compressed nutlets.

There are differences in opinion on whether to recognise Cyperus sensu lato, in a very broad sense with a number of subgenera (e.g. subgenus Kyllinga, and [C.sub.3] and [C.sub.4] species of Cyperus sensu stricto in different subgenera; e.g. Kukenthal, 19351936), or in a narrow sense with various segregate genera (with Cyperus sensu stricto including [C.sub.3] and [C.sub.4] species; e.g. Goetghebeur, 1998). Our results show Cyperus sensu stricto to be polyphyletic, and merging all the segregate taxa into broadly circumscribed Cyperus sensu lato and recognizing various segregates as subgenera would make a monophyletic entity. However, this option is not favored because it would result in a big genus (c. 900 species) and reduce taxonomic clarity. Other partial merging of the taxa into Cyperus, recognizing Oxycaryum, Kyllingiella, Sphaerocyperus, Remirea, Lipocarpha, and Ascolepis as distinct, but treating Courtoisina, Kyllinga, Pycreus, Queenslandiella, and Alinula as subgenera of Cyperus (e.g. Haines & Lye, 1983; Lye, 1997) is not supported by this study.

We follow Goetghebeur (1998) in recognizing Cyperus sensu stricto and recognizing the segregate taxa at generic rank (Table 2) pending more intensive phylogenetic studies to get a full resolution of their relationships.

Future Research

Molecular phylogenetic studies have focused more attention on the Ficinia clade (38% sampling) and less on the Cyperus clade (5% sampling), yet Cyperus clade exhibits wide morphological variation. With more intensive molecular phylogenetic studies and more extensive sampling to include the complete diversity of growth form and morphological types, we expect a better understanding of character homology, which will allow better-informed decisions about generic limits.

Acknowledgements AMM acknowledges a visiting postdoctoral fellowship from the Belgian Fund for Scientific Research-Flanders (FWO-Vlaanderen, G.0104.01N) and a grant from the K.U. Leuven (grant F/ 02/052) during the period this paper was prepared.

Open Access This article is distributed under the terms of the Creative Commons Attribution Noncommercial License which permits any noncommercial use, distribution, and reproduction in any medium, provided the original author(s) and source are credited.

Published online: 6 December 2008

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A. Muthama Muasya (1,2,6) * Alexander Vrijdaghs (2) * David A. Simpson (3) * Mark W. Chase (3) * Paul Goetghebeur (4) * Erik Smets (2,5)

(1) Botany Department, University of Cape Town, Rondebosch 7700, South Africa

(2) Laboratory of Plant Systematics, K.U. Leuven, Kasteelpark Arenberg 31, BE-3001 Leuven, Belgium

(3) Royal Botanic Gardens Kew, Richmond Surrey TW9 3DS, UK

(4) Department of Biology, Ghent University, K.L. Ledegancksraat 35, B-9000 Ghent, Belgium

(5) National Herbarium of the Netherlands, R O. Box 9514, NL-2300 RA Leiden, Netherlands

(6) Author for Correspondence; e-mail:
Table 1 List of Taxa Sampled with Vouchers and Genbank Accession

Taxon Voucher

Cyperoideae Suess.

 Abildgaardieae Lye

 Abildgaardia ovata Kenya: Muasya et al. 684
 (Burm. f.) Kral (EA, K)

 Fimbristylis dichotoma (L.) Kenya: Muasya 1006
 Vahl (EA, K)

 Cypereae Dumort.

 Alinula lipocarphoides Kenya: Muasya: 2592 (EA)
 (Kuk.) J. Raynal

 Alinula paradoxa Goetgh. & Tanzania: Faden et al.
 Vorster 96/29 (K)

 Androtrichum giganteum Argentina: Tressens et al.
 (Kunth) H. Pfeiff. 4292 (K)

 Androtrichum trigynum Argentina: Goetghebeur
 (Spreng.) H. Pfeiff. 4764 (GENT)

 Ascolepis capensis (Kunth) Kenya: Muasya 1009
 Ridl. (EA, K)

 Ascolepis protea Welw. Congo: Fay 2700 (K)

 Courtoisina assimilis Tanzania: Faden et al.
 (Steud.) Maquet 96/119 (K)

 Cyperus compressus L. Thailand: Muasya 1375 (K)

 Cyperus cuspidatus Kunth Thailand: Muasya 1374 (K)

 Cyperus involucratus Rottb. Madagascar: Kew Acc.

 Cyperus laevigatus L. Kenya: Muasya 1041 (EA)

 Cyperus longus L. Europe: Chase 2276 (K)

 Cyperus papyrus L. Chad: Hepper 4213 (K)

 Cyperus pulchellus R. Br. Thailand: Muasya 1377 (K)

 Cyperus pygmaeus Rottb. Kenya: Muasya 1133 (K)

 Desmoschoenus spiralis New Zealand: Ford 44/94
 Hook. f. (NU)

 Ficinia bergiana Kunth S. Africa: Muasya 2337 (BOL)

 Ficinia distans C. B. Clarke S. Africa: Muasya 2283 (BOL)

 Ficinia esterhuyseniae S. Africa: Muasya 2312 (BOL)

 Ficinia gracilis Schrad. Tanzania: Faden et al.
 96/433 (K)

 Ficinia nodosa (Rottb.) Australia: Stind 21216 (K)
 Goetgh., Muasya & D. A.

 Ficinia rigida Levyns S. Africa: Muasya 2319 (K)

 Ficinia trichodes (Schrad.) S. Africa: Muasya 2328 (K)
 Benth. & Hook. f.

 Ficinia radiata (L. f.) Kunth S. Africa: Muasya 2310 (K)

 Hellmuthia membranacea S. Africa: Weerderman et al.
 (Thunb.) R. W. Haines & 269 (K); Muasya 1145 (K)

 Isolepis cernua (Vahl) BRITAIN: Muasya 1058 (K)
 Roem. & Schult. var.

 Isolepis fluitans (L.) R. Br. Kenya: Muasya 1057 (K)

 Isolepis hystrix (Thunb.) S. Africa: Muasya 1150 (K)

 Isolepis levynsiana Muasya S. Africa: Muasya 1151 (K)
 & D. A. Simpson

 Isolepis marginata (Thunb.) Australia: Coveny et al.
 A. Dietr. 17452 (K)

 Isolepis setacea (L.) R. Br. Kenya: Muasya 1059 (K)

 Isolepis tenuissima (Nees) S. Africa: Muasya 2369 (K)

 Isolepis venustula Kunth S. Africa: Muasya 1189 (K)

 Kyllinga appendiculata K. Kenya: Muasya 1050 (EA, K)

 Kyllinga brevifolia Rottb. Australia: Coveny et al.
 17459 (K)

 Kyllinga bulbosa P. Beauv. Kenya: Muasya 1020 (EA, K)

 Kyllingiella microcephala Zimbabwe: Muasya et al.
 (Steud.) R. W. Haines & 1118 (K)

 Kyllingiella polyphylla (A. Tanzania: Wingfield 497 (K)
 Rich.) Lye

 Lipocarpha hemisphaerica Thailand: Muasya 1217 (K)
 (Roth.) Goetgh.

 Lipocarpha nana (A. Rich.) Kenya: Muasya 972
 J.Raynal (EA, K)

 Oxycaryum cubense (Poepp. ZAMBIA: Richards
 & Kunth) E.Palla 13318 (K)

 Pycreus flavescens (L.) Kenya: Muasya 1022 (EA, K)

 Pycreus nuerensis (Boeck.) Tanzania: Muasya 940
 S.S.Hooper (EA, K)

 Queenslandiella hyalina Kenya: Mwachala 296 (EA)
 (Vahl) Ballard

 Remirea maritima Aubl. Tanzania: Faden et al.
 96/48 (K)

 Scirpoides holoschoenus S. Africa: Acocks s.n. (K)
 (L.) Sojak

 Scirpoides thunbergii S. Africa: Muasya 1205 (K)
 (Schrad.) Sojak

 Scirpus falsus C. B. Clarke S. Africa: Hilliard 13609

 Scirpus ficinioides Kunth S. Africa: Hilliard 16095

 Sphaerocyperus erinaceus Tanzania: Faden et al. 96/338
 (Ridl.) Lye (K)

 Volkiella disticha Merxm. & Namibia: Muller et al.
 Czech 4245 (K)

 Eleocharideae Goetgh.

 Eleocharis marginulata Kenya: Muasya 1039 (EA, K)

 Fuireneae Reichenb. ex Fenzl

 Actinoscirpus grossus (L. f.) Malaysia: Simpson 2660 (K)
 Goetgh. & D. A. Simpson

 Bolboschoenus maritimus Botswana: Smith 2452 (K)
 (L.) Palla

 Bolboschoenus nobilis S. Africa: Leistner 144 (K)
 (Ridl.) Goetgh. & D. A.

 Fuirena sp. Brazil: Thomas et al. 10404

 Isolepis humillima (Benth.) Australia: Thomas et al. 622
 K. L. Wilson (BRI)

 Schoenoplectiella articulata Tanzania: Muasya 947
 (L.) Lye (EA, K)

 Schoenoplectus corymbosats Kenya: Muasya 1004 (EA)
 (Roth ex Roem. & Schult.)
 J. Raynal

 Schoenoplectus lacustris (L.) Britain: Muasya 1043 (K)

 Schoenoplectus litoralis Hong Kong: Shaw 883 (K)
 (Schrad.) Palla

 Scirpeae Kunth ex Dumort.

 Eriophorum vaginatum L. Poland: Beyer et al. 2 (K)

 Eriophorum viridicarinatum USA: Boufford 23053 (WS)
 (Engl.) Fern.

 Scirpus ancistrochaetus USA: Nacsi 7544 (DOV)

 Scirpus sylvaticus L. HBUG/86-0541 (GENT)

Mapanioideae C. B. Clarke

 Hypolytreae Presl ex Fenzl

 Hypolytrum nemorum (Vahl) Malaysia: Simpson 1379 (K)

 Mapania cuspidata (Miq.) Brunei: Marsh 4 (K)

Taxon Genbank accession numbers

 rbcL rps16 trnL-F OR
Cyperoideae Suess.

 Abildgaardieae Lye

 Abildgaardia ovata Y12985 AJ295754
 (Burm. f.) Kral

 Fimbristylis dichotoma (L.) Y13008 AJ295755

 Cypereae Dumort.

 Alinula lipocarphoides -- -- EF178608
 (Kuk.) J. Raynal

 Alinula paradoxa Goetgh. & AJ278290 -- AJ295756

 Androtrichum giganteum EF179546 --
 (Kunth) H. Pfeiff.

 Androtrichum trigynum EF178547 --
 (Spreng.) H. Pfeiff.

 Ascolepis capensis (Kunth) Y13003 AF449518 AJ295757

 Ascolepis protea Welw. Y13002 -- --

 Courtoisina assimilis AY40590 AY449519 AY040595
 (Steud.) Maquet

 Cyperus compressus L. AF449506 AF449521 AF449555/-

 Cyperus cuspidatus Kunth AF449508 AF449523 AF449557/

 Cyperus involucratus Rottb. Y12967 AF445920 AJ295758

 Cyperus laevigatus L. Y13017 AF449527 AY040596

 Cyperus longus L. Y13015 AF449528 AY040598

 Cyperus papyrus L. Y12966 AF449531 AJ295759

 Cyperus pulchellus R. Br. AY40591 -- AY040599

 Cyperus pygmaeus Rottb. AJ404698 AF449534 AJ295760

 Desmoschoenus spiralis AJ404701 -- AJ295753
 Hook. f.

 Ficinia bergiana Kunth EF200588 EF078974 EF179593

 Ficinia distans C. B. Clarke EF178548 EF178594

 Ficinia esterhuyseniae EF178549 EF078975 EF178590

 Ficinia gracilis Schrad. EF178550 EF178534

 Ficinia nodosa (Rottb.) Y12984 EF174386 AJ295793
 Goetgh., Muasya & D. A.

 Ficinia rigida Levyns EF178557 EF174387 EF178602

 Ficinia trichodes (Schrad.) EF178558 EF174388 EF178603
 Benth. & Hook. f.

 Ficinia radiata (L. f.) Kunth EF200589 EF078976 --

 Hellmuthia membranacea Y13000 EF174389 AJ295815
 (Thunb.) R. W. Haines &

 Isolepis cernua (Vahl) Y13014 AF449538 AJ295775
 Roem. & Schult. var.

 Isolepis fluitans (L.) R. Br. Y12961 EF174390 AJ295780

 Isolepis hystrix (Thunb.) AJ404711 -- AJ295785

 Isolepis levynsiana Muasya AF449514 AF449514 AF449563/
 & D. A. Simpson AF449575

 Isolepis marginata (Thunb.) AJ404714 EF174391 AJ295790
 A. Dietr.

 Isolepis setacea (L.) R. Br. Y12962 EF174392 AJ295799

 Isolepis tenuissima (Nees) AY725947 --

 Isolepis venustula Kunth AJ404724 -- AJ295804

 Kyllinga appendiculata K. Y13007 AF449542 AJ295761

 Kyllinga brevifolia Rottb. AF449515 AF449543 AF449564/

 Kyllinga bulbosa P. Beauv. Y12979 AF449544 AY040601

 Kyllingiella microcephala AY040592 AF449540 AJ295807
 (Steud.) R. W. Haines &

 Kyllingiella polyphylla (A. Y13013 AF449541 AJ295515
 Rich.) Lye

 Lipocarpha hemisphaerica AF449516 AF449546 AF449565/
 (Roth.) Goetgh. AF449577

 Lipocarpha nana (A. Rich.) Y12990 AF449545 AJ295762

 Oxycaryum cubense (Poepp. Y13006 -- AY040602
 & Kunth) E.Palla

 Pycreus flavescens (L.) Y13005 AF449547 AJ295763

 Pycreus nuerensis (Boeck.) Y13004 AF449549 AY040603

 Queenslandiella hyalina AY725953 -- --
 (Vahl) Ballard

 Remirea maritima Aubl. AY040593 AF449550 AY040604

 Scirpoides holoschoenus Y12994 AY344153 AJ295811
 (L.) Sojak

 Scirpoides thunbergii AJ404727 AF449551 AJ295812
 (Schrad.) Sojak

 Scirpus falsus C. B. Clarke EF178559 EF174393 --

 Scirpus ficinioides Kunth EF178560 EF174394 --

 Sphaerocyperus erinaceus AJ404699 AF449552 AJ295764
 (Ridl.) Lye

 Volkiella disticha Merxm. & EF178561 -- --

 Eleocharideae Goetgh.

 Eleocharis marginulata Y13011 -- AJ295768

 Fuireneae Reichenb. ex Fenzl

 Actinoscirpus grossus (L. f.) Y12953 -- AJ295765
 Goetgh. & D. A. Simpson

 Bolboschoenus maritimus Y12996 -- AJ295767
 (L.) Palla

 Bolboschoenus nobilis Y12995 -- --
 (Ridl.) Goetgh. & D. A.

 Fuirena sp. Y12970 -- --

 Isolepis humillima (Benth.) AJ404728 AF449539 AJ295784
 K. L. Wilson

 Schoenoplectiella articulata Y12987 -- --
 (L.) Lye

 Schoenoplectus corymbosats EF178570 -- EF178607
 (Roth ex Roem. & Schult.)
 J. Raynal

 Schoenoplectus lacustris (L.) Y12943 AF449554 AJ295809

 Schoenoplectus litoralis EF178571 -- --
 (Schrad.) Palla

 Scirpeae Kunth ex Dumort.

 Eriophorum vaginatum L.
 Y12951 AF449553 AJ295769
 Eriophorum viridicarinatum U49230 -- --
 (Engl.) Fern.

 Scirpus ancistrochaetus EF178578 EF174395 --

 Scirpus sylvaticus L.
 EF178586 EF174396 --
Mapanioideae C. B. Clarke

 Hypolytreae Presl ex Fenzl

 Hypolytrum nemorum (Vahl) Y12958 AY344142 AJ295816

 Mapania cuspidata (Miq.) Y12955 DQ058318 AJ295817

Classification following interpretation of current data and
Goetghebeur (1998)

Table 2 Summary of Some of the Diagnostic Characters of the
Genera in Cypereae

Genus (total/ Habit Floret Glume
studied species) no. arrangement

Alinula (4/2) Annual One Distichous

Androtrichum (2/2) Perennial Many Distichous

Ascolepis (20/2) Annual/ One Distichous

Ascopholis (1/0) Perennial One Distichous

Courtoisina (2/1) Annual Many Distichous

Cyperus (550/7) Annual/ 1-Many Distichous/
 perennial spiral

Desmoschoenus (1/1) Perennial Many Spiral

Ficinia (60/8) Perennial Many Distichous/

Hellmuthia (1/1) Perennial Many Spiral

Isolepis (70/9) Annual Many Distichous/
 (perennial) spiral

Kyllinga (60/2) Perennial Many Distichous

Kyllingiella (4/2) Perennial Many Spiral

Lipocarpha (35/2) Annual/ One Distichous

Oxycaryum (1/1) Annual Many Spiral

Pycreus (100/2) Annual/ Many Distichous

Queenslandiella (1/1) Annual Many Distichous

Remirea (1/1) Perennial One Distichous

Scirpoides (5/2) Perennial Many Spiral

Scirpus spp. (3/2; Perennial Many Spiral
 Southern Africa)

Sphaerocyperus (1/1) Perennial One Distichous

Volkiella (1/1) Annual One Distichous

Genus (total/ Dispersal Nutlet Photosynthetic
studied species) unit orientation type

Alinula (4/2) Nutlet Dorsiventral [C.sub.4]

Androtrichum (2/2) Nutlet & Dorsiventral [C.sub.3]

Ascolepis (20/2) Spikelet/ Dorsiventral [C.sub.4]

Ascopholis (1/0) Spikelet Dorsiventral [C.sub.4]

Courtoisina (2/1) Spikelet Dorsiventral [C.sub.3]

Cyperus (550/7) Spikelet/ Dorsiventral [C.sub.3] &
 nutlet [C.sub.4]

Desmoschoenus (1/1) Nutlet Dorsiventral [C.sub.3]

Ficinia (60/8) Nutlet Dorsiventral [C.sub.3]

Hellmuthia (1/1) Nutlet Dorsiventral [C.sub.3]

Isolepis (70/9) Nutlet Dorsiventral [C.sub.3]

Kyllinga (60/2) Spikelet Lateral [C.sub.4]

Kyllingiella (4/2) Nutlet Dorsiventral [C.sub.3]

Lipocarpha (35/2) Spikelet Dorsiventral [C.sub.4]

Oxycaryum (1/1) Nutlet Dorsiventral [C.sub.3]

Pycreus (100/2) Nutlet Lateral [C.sub.4]

Queenslandiella (1/1) Spikelet Lateral [C.sub.4]

Remirea (1/1) Spikelet Dorsiventral [C.sub.4]

Scirpoides (5/2) Nutlet Dorsiventral [C.sub.3]

Scirpus spp. (3/2; Nutlet Dorsiventral ?

Southern Africa)

Sphaerocyperus (1/1) Spikelet Dorsiventral [C.sub.4]

Volkiella (1/1) Spikelet Dorsiventral [C.sub.4]

Classification following interpretation of current data and
Goetghebeur (1998).
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Author:Muasya, A. Muthama; Vrijdaghs, Alexander; Simpson, David A.; Chase, Mark W.; Goetghebeur, Paul; Smet
Publication:The Botanical Review
Article Type:Report
Geographic Code:1USA
Date:Mar 1, 2009
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