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Phylogeny of cyperaceae based on DNA sequence data: current progress and future prospects.


Cyperaceae comprise 109 genera and approximately 5,500 species and have an almost cosmopolitan distribution (Govaerts et al., 2007). About 35% of the genera are monotypic, 26% have two to five species, and there are a seven (6%) genera with over 200 species, the largest being Cyperus (686 species) and Carex (1,757 species; Goetghebeur, 1998). The family shows extreme reduction in floral morphology, and the majority of the smaller genera are carved out of the larger ones on the basis on one or few distinguishing features.

Family level phylogenetic studies in the last ten years have used morphological (e.g. Simpson, 1995; Bruhl, 1995; Goetghebeur, 1998), molecular (e.g. Muasya et al., 1998; Simpson et al., in press), and combined morphological and molecular data (Muasya et al., 2000b). The two most recent classifications based on morphological data (including gross morphology, anatomy and embryology) differ in suprageneric groupings of tribes and subfamilies. Bruhl (1995) recognised two subfamilies, Cyperoideae and Caricoideae, whereas Goetghebeur (1998) recognised two additional subfamilies, Sclerioideae and Mapanioideae, both of which were included in Caricoideae by Bruhl (1995). The treatments also differed in tribal circumscription, with Bruhl (1995) recognising 12 tribes and treating Scirpeae broadly to include taxa classified in tribes Dulicheae, Fuireneae, Eleocharideae and Cypereae sensu Goetghebeur (1998).

Molecular DNA sequence data are increasingly used in angiosperm classification. In Cyperaceae, broad suprageneric studies have so far sampled all subfamilies and tribes, but sampling effort is not evenly distributed among all tribes. Family-level studies have been based mainly on rbcL sequence data (e.g. Muasya et al., 1998; Simpson et al., 2007), whereas at tribal or subfamilial levels other plastid and nuclear regions have been used. The plastid regions rps16 intron, trnL intron and trnL-F intergenic spacer have been used in studies of subfamily Mapanioideae (e.g. Simpson et al., 2003) and a number of studies at tribal and generic level.

This study uses three of the most commonly used plastid regions (the rbcL gene, the trnL intron, and the trnL-F spacer) to reconstruct relationships of the family and presents an overview of the current status of suprageneric phylogenetic studies. The rbeL gene has been sequenced for over 60% genera of Cyperaceae (e.g. Simpson et al., 2007)and can be aligned unambiguously, whereas trnL-F (both the trnL intron and the trnl-F intergenic spacer) has been used to a greater extent in generic studies and is more difficult to align at the family level.

Analysis of rbcL and trnl-F Data

The analysis includes a total 262 taxa (258 species) of Cyperaceae in 93 genera from the 15 tribes and four subfamilies recognised by Goetghebeur (1998). Sequences from previous studies (Bremer, 2002; Dhooge et al., 2003; Muasya et al., 1998, 2000a, 2000b, 2001, 2002; Simpson et al., 2003, 2007; Verboom, 2006; Zhang et al., 2004) were analysed together with 41 newly sequenced taxa representing 22 genera, nine of which had not been previously sequenced. Total DNA was extracted from vegetative material (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., 2002); the resulting sequences were aligned manually and are lodged with GenBank (Table 1).

Heuristic analyses were carried out using PAUP* (Swofford, 2002). Searches were conducted under Fitch (1971) parsimony, TBR (tree-bissection-reconnection) branch swapping, and random taxon addition (5,000) with the MulTrees option in effect and retaining only ten trees per replicate. Internal support was estimated using 1000 bootstrap replicates (Felsenstein, 1985), with the following search parameters: simple taxon addition, TBR branch-swapping, and MulTrees option in effect with only ten trees held per step.

The aligned matrix has 3,573 characters comprising 1,428 from rbcL and 2145 from trnl-F (intron and intergenic spacer) region. Some portions of trnl-F could not be unambiguously aligned, and 865 characters were excluded from the analysis, leaving 2,708 characters, of which 913 are potentially parsimony informative.

Fifty equally parsimonious trees were recovered of length=5,467 steps, consistency index (CI)=0.45 and retention index (RI)=0.79. The strict consensus tree (Figs. 1, 2 and 3) is presented and discussed below.

Subfamily Relationships

Cyperaceae are resolved as monophyletic and sister to Juncaceae, with Mapanioideae sister to all the other Cyperaceae (Fig. 1; tribes and subfamilies sensu Goetghebeur, 1998). Within the last clade, Coleochloa-Microdracoides (Trilepideae) form a clade sister to the rest of Cyperaceae. Trilepideae are not sister to Scleria-Diplacrum (Sclerieae) as the latter are embedded in the Schoeneae, and therefore the Sclerioideae are not monophyletic. Also Caricoideae are sister to Scirpeae and embedded within Cyperoideae (Fig. 2).

Similar studies in which Mapanioideae are resolved as sister to rest of Cyperaceae have been reported by Bruhl (1995) based on morphological studies and in previous family-level DNA studies (e.g. Muasya et al., 1998, 2000b; Simpson et al., 2007). Mapanioideae have a unique floral morphology compared with the rest of Cyperaceae, with floral units each comprising two to ten or more scales (the lower ones being keeled), two to ten stamens and a single gynoecium. The floral units have been variously interpreted as bisexual flowers in which the arrangement of the structures has been disturbed (Goetghebeur, 1998) or as reduced partial inflorescences termed spicoids (Simpson et al., 2003; Richards et al., 2006).


Recognition of Caricoideae and Sclerioideae as subfamilies separate from Cyperoideae (e.g. Goetghebeur, 1998), based on unique morphological characters is not supported by current analysis. Typical Cyperoideae are mostly diagnosed by having at least one (sometimes all) bisexual flower, whereas in Sclerioideae (and some unusual Cyperoideae) they are all unisexual and in Caricoideae they are all unisexual and enclosed by a utricle. This study and other analyses of DNA data support the recognition of two subfamilies in Cyperaceae, Mapanioideae and Cyperoideae, as proposed by Simpson et al. (2003, 2007).


Tribes of the Cyperaceae

A number of tribal groups recognised in the recent classification of Cyperaceae by Goetghebeur (1998) are supported by the current study. Within Mapanioideae, some Hypolytreae (Mapania, Hypolytrum and Scirpodendron) and Chrysitricheae (Lepironia, Chrysitrix and Chorizandra) form clades separate from a polytomy comprising other mapanioids (Fig. 1). Although the polytomy observed may be caused by insufficient data for some of the taxa, Capitularina and Exocarya (both of which traditionally have been placed in Hypolytreae) were resolved together in Chrysitricheae in a combined pollen and DNA data study (Simpson et al., 2003).

The inselberg taxa in Trilepideae (Coleochloa to Microdracoides; Fig. 1) form a strongly supported clade. This clade is sister to the rest of Cyperoideae and not to other tribes of Sclerioideae (sensu Goetghebeur 1998), namely Cryptangieae, Sclerieae and Bisboeckelereae. These other tribes are embedded among Schoeneae (Fig. 1). Notable is the position of Exochogyne, a genus unplaced in any tribe of Sclerioideae by Goetghebeur (1998) due to unclear morphological homologies, and which is resolved here among Cryptangieae.

Schoeneae are one of the most heterogeneous tribes in the family, having 29 genera of which Rhynchospora is among the largest; over 50% of the genera have fewer than 10 species (Goetghebeur, 1998). This analysis resolves four clades within Schoeneae: (1) Cladium, (2) Gymnoschoenus, (3) Caustis to Didymiandrum, and (4) Rhynchospora (Figs. 1 and 2). The moderately supported Rhynchospora clade has been previously classified in a separate tribe Rhynchosporeae (e.g. Goetghebeur, 1986; Bruhl, 1995) on the basis of, inter alia, distinct style base. Members of the former Sclerioideae (Cryptangieae, Bisboeckelereae and Sclerieae) are resolved among clade (3), an observation reported in previous studies (e.g. Simpson et al., 2007). Schoeneae have an essentially Gondwanan distribution, and several widely distributed genera (e.g. Costularia, Tetraria; Zhang et al., 2004, Verboom, 2006) are polyphyletic. Morelotia is resolved in a clade which includes Costularia, Tricostularia and reticulate-sheathed Tetraria (Tricostularia clade in Verboom, 2006), and not together with Ghania. A close relationship between Morelotia and Ghania has been suggested by several authors (e.g. Goetghebeur, 1986), while Bruhl (1995) argued against this relationship after recovering Morelotia distant from Ghania. The monotypic Schoenoides (Seberg, 1988) is embedded in Oreobolus here and in other studies (Mandrinan et al., 2004), further supporting the inclusion of Schoenoides in Oreobolus (e.g. Curtis & Morris, 1994; Govaerts et al., 2007). There have been limited phylogenetic studies in Schoeneae (e.g. Zhang et al., 2004; Verboom, 2006), which lack bootstrap support for the basal nodes, and more data are needed to resolve relationships among the taxa. Further studies of Schoeneae are in progress (Bruhl et al. and Verboom et al., unpublished data).


The moderately supported clade (Khaosokia to Dulichium) includes members of Cariceae, Scirpeae and Dulicheae (Fig. 2). Khaosokia is resolved sister to the rest of the members of this clade, a position suggested by Simpson et al. (2005) from observations of gross morphology and DNA studies. Scirpeae are not monophyletic, as Dulicheae are embedded between Scirpeae l and Scirpeae II. In Scirpeae I, the generic boundaries between Trichophorum and Oreobolopsis are unclear, and further attention is needed to resolve the polyphyly of Trichophorum. Phylogenetic studies involving Andean species of Scirpus have recently led to description of a new segregate genus, Zameioscirpus (e.g. Dhooge et al., 2003). Carex is polyphyletic and includes other genera of Cariceae, a similar pattern has been observed in previous studies (e.g. Yen & Olmstead, 2000; Start et al., 2004)

Fuireneae are split into four clades (Fig. 2) in our analysis. Fuireneae I (Fuirena) is sister to Eleocharideae, Fuireneae II (Bolboschoenus) is sister to Abildgaardieae, whereas Fuireneae III (Schoenoplectus and Actinoschoenus) and Fuireneae IV (Schoenoplectiella group) form a polytomy with Cypereae. Relationships among these groups based on DNA data remain unstable (cf. Simpson et al., 2007). Schoenoplectus is paraphyletic with several tropical African perennial taxa (e.g. S. mucronatus) being resolved together with Schoenoplectiella. Schoenoplectiella, recently segregated to include annual amphicarpous taxa of Schoenoplectus (Lye, 2003), is resolved into a strongly supported clade that includes perennial tropical Schoenoplectus species sharing a lateral spikelet morphology. Further studies are in progress (Muasya et al., unpublished data) evaluating relationships in the group.

Abildgaardieae are resolved to include Arthrostylis aphylla, Trachystylis strandbrokensis and Actinoschoenus repens (Fig. 2), taxa which have been previously placed in Schoeneae (Goetghebeur, 1998). Arthrostylis and Actinoschoenus have been shown to be closer to Abildgaardieae based on plastic and nuclear ribosomal (ITS) data (Ghamkhar et al., 2007). Both Arthrostylis and Trachystylis are monotypic Australian taxa with bisexual flowers that lack perianth segments, but share gross morphological similarity with Schoeneae (e.g. one- to few-flowered spikelet and wide glume wings enclosing the next flower). On the other hand, Actinoschoenus repens is a Zambian endemic, with morphological similarity to both Abildgaardieae and Schoeneae. Although these three taxa had been placed in Schoeneae even with decisive anatomical and embryological data lacking, the DNA data resolve them in Abildgaardieae, and similar results were obtained independently by Ghamkhar et al. (2007). We therefore propose their formal inclusion in this tribe.

Cypereae form a strongly supported clade (Fig. 3) that has received intensive DNA phylogenetic study, both at generic (e.g. Muasya et al., 2001, 2002) and tribal levels (Muasya et al., 2008). Cypereae are characterised by the presence of Cyperus-type embryo and here include Hellmuthia, a genus previously considered to belong in Chrysitricheae (e.g. Haines & Lye, 1976; Goetghebeur, 1998; cf. Vrijdaghs et al., 2006, 2008). Scirpus falsus and S. ficinioides, taxa from the Drakensberg Mountians in South Africa and previously placed in Scirpeae, are resolved here among Cypereae in a clade including Ficinia, Isolepis, Hellmuthia and Scirpoides. More studies are in progress to describe a new genus including these taxa (authors, unpublished data).

Revised Suprageneric Classification of Cyperaceae

Based on the available data, we support the revised classification of Cyperaceae into two subfamilies, Mapanioideae and Cyperoideae (Figs. 1, 2 and 3). We also broadly accept the tribal circumscriptions of Goetghebeur (1998) but with modification to tribes Cypereae (to include Hellmuthia and the perianth-bearing Drakensberg Scirpus, S. falsus and S. ficinioides); Abildgaardieae (to include Arthrostylis, Trachystylis and Actinoschoenus); Schoeneae (recognising Rhynchosporeae, Rhynchospora and Pleurostachys); and Cryptangieae (to include Didymiandrum and Exochogyne). We refrain from recognising Cladieae (Cladium) pending more studies.

Future Research

Choice of marker and uneven sampling limit the scope for analysing different data sets in combination. The current study and a number of other ongoing studies have focused on more slowly evolving plastid regions, which have less resolution but can be aligned across the family. Among research groups in different institutes, there is need to study the same DNA regions (e.g. rbcL, trnl-F, rps16) for similar taxa to enable different data sets to be aligned in combination.

The intensity of sampling varies among tribes. Although Chrysitricheae, Cypereae, Hypolytreae, Scirpeae and Cariceae are among the better studied tribes, more effort is needed to elucidate phylogenetic relationships within Cryptangineae, Bisboeckelerieae, Fuireneae, Schoeneae, and Sclerieae.

Acknowledgements A number of the contributors acknowledge funding from various sources. AMM acknowledges funding from the Belgian Fund for Scientific Research-Flanders (FWO-Vlaanderen, G.0104.01N); K.U.Leuven (grant F/02/052); University of Cape Town (Smuts Fellowship); and the Norwegian Council of Universities' Committee for Development Research and Education (NUFU project 53/03). GAV acknowledges funding from the National Research Council, South Africa.

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: 5 December 2008

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A. Muthama Muasya (1,6,7) * David A. Simpson (2) * G. Anthony Verboom (1) * Paul Goetghebeur (3) * Robert F. C. Naczi (4). Mark W. Chase (2) * Erik Smets (5,6)

(1) Botany Department, University of Cape Town, Rondebosch 7701, Cape Town, South Africa

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

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

(4) Department of Agriculture & Natural Resources, Delaware State University, Dover, DE 19901, USA

(5) National Herbarium of the Netherlands, PO Box 9514, NL-2300 RA Leiden, The Netherlands

(6) Laboratory of Plant Systematics, K.U. Leuven, Kasteelpark Arenberg 31, BE-3001 Leuven, Belgium; e-mail:

(7) Author for correspondence; e-mail:
Table 1 List of Taxa Sampled with vouchers and Genbank Accession
Numbers. Classification Following Interpretation of Current
Data and Goetghebeur (1998)

Taxon Voucher

Cyperoideae Suess.

Abildgaardieae Lye

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

 Actinoschoenus repens Raynal Zambia: Robinson 3643 (K)

 Arthrostylis aphylla R. Br. Australia: Wilson 8249 (NSW)

 Bulbostylis atrosanguinea Kenya: Muasya 1037 (EA, K)
 (Boeck.) C. B. Clarke

 Bulbostylis hispidula Kenya: Muasya 1025 (EA, K)
 (Vahl) R. W. Haines

 Crosslandia setifolia W. Fitzg. Australia: Wilson 10147 (K)

 Fimbristylis complanata Kenya: Muasya 1029 (EA, K)
 (Retz.) Link

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

 Nemum spadiceum (Lam.) WEST AFRICA: Baldwin
 Desv. ex Ham. 9766 (K)

 Trachystylis stradbrokensis Australia: Wilson 8175 (K)
 (Domin) Kuk.

Bisboeckelereae Pax ex
 L.T. Eiten

 Becquerelia cymosa Brongn. Brazil: Thomas et al.
 10284 (NY)

 Calyptrocarya bicolor RBGKEW DNA 10389
 (H. Pfeiff.) Koyama

 Diplacrum africanum Tanzania: Vollensen 3967 (K)
 C. B. Clarke

Cariceae Kunth ex Dumort.

 Carex cephalophora Kress et al. (2005)
 Muhl. ex Willd.

 Carex conferta A. Rich. Kenya: Muasya 1055 (K)

 Carex echinochloe Kunze Kenya: Muasya 1051 (K)

 Carex hostiana DC. Chase et al. (1993)

 Carex monostachya A. Rich. Kenya: Muasya 1052 (K)

 Carex sylvatica Huds. Simpson et al. (2003)

 Kobresia simpliciuscula Plunkett et al. (1995);
 (Wahlenb.) Mackenzie Yen et al. (2000)

 Uncinia nemoralis Australia: Wilson et al.
 K. L. Wilson 9533 (K)

 Schoenoxiphium ecklonii Nees S. Africa: Williams 968 (K)

 Schoenoxiphium lehmannii Tanzania: J M G 94/94 (K)
 (Nees) Kunth ex Steud.

 Schoenoxiphium sparteum Kenya: Muasya 2566 (EA)
 (Wahlenb.) C.B.Clarke

Uncertain tribe aff. Cariceae

 Khaosokia caricoides Thailand: Simpson et al.
 D.A.Simpson, Chayam. & 1886 (K)

Cryptangieae Benth.

 Didyndandrum stellatum Venezuela: Liesner 23562 (GENT)
 (Boeck.) Gilly

 Exochogyne amazonica Brazil: Aparecida da Silva
 C. B. Clarke 1986 (GENT)

 Lagenoearpus alboniger Brazil: Thomas 11111 (NY)
 (A. St.-Hil.) C. B. Clarke

Cypereae Dumort.

 Alinula paradoxa Tanzania: Faden et al.
 Goetgh. & 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 Kenya: Muasya 1009 (EA, K)
 (Kunth) Ridl.

 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 congestus Vahl Australia: Coveny et al.
 17492 (K)

 Cyperus cuspidatus Kunth Thailand: Muasya 1374 (K)

 Cyperus cyperoides (L.) Kuntze Thailand: Muasya 1277 (K)

 Cyperus dichroostachyus Kenya: Muasya 976 (EA, K)
 A. Rich.

 Cyperus endlichii Kuk. Kenya: Muasya 695 (K)

 Cyperus involucratus Rottb. Madagascar: Kew Ace. 6136603

 Cyperus kerstenii Boeck. Kenya: Muasya 984 (EA, K)

 Cyperus laevigatus L. Kenya: Muasya 1041 (EA)

 Cyperus longus L. Europe: Chase 2276 (K)

 Cyperus meeboldii Kuk. Kenya: Muasya 1255 (EA, K)

 Cyperus papyrus L. Chad: Hepper 4213 (K)

 Cyperus plateilema Kenya: Muasya 969 (EA, K)
 (Steud.) Kuk.

 Cyperus pseudovestitus Kenya: Muasya 1268 (K)
 (C. B. Clarke) Kuk.

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

 Cyperus pygmaeus Rottb. Kenya: Muasya 1133 (K)

 Cyperus rigidifolius Steud. Kenya: Muasya 1031 (K)

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

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

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

 Ficinia esterhuyseniae Muasya S. Africa: Muasya 2312 (BOL)

 Ficinia gracilis Schrad. S. Africa: Muasya 2355

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

 Ficinia gydomontana T. H. S. Africa: Muasya 2333 (BOL)
 Arnold & K. D. Gordon-Gray

 Ficinia indica S. Africa: Muasya 2318 (BOL)
 (Lam.) H. Pfeiff.

 Ficinia laciniata (Thunb.) Nees S. Africa: Muasya 2340 (BOL)

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

 Ficinia paradoxa (Schrad.) Nees S. Africa: Verboom 534 (BOL)

 Ficinia pinguior C. B. Clarke S. Africa: Muasya 1183 (K)

 Ficinia polystachya Levyns S. Africa: Muasya 2330 (K)

 Ficinia ramosissima Kunth S. Africa: Muasya 2288 (K)

 Ficinia repens Kunth S. Africa: Muasya 2347 (K)

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

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

 Ficinia tristachya S. Africa: Muasya 1233 (K)
 (Rottb.) Nees

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

 Isolepis aucklandica Hook. f Australia: Wilson et al.
 9462 (K)

 Isolepis bicolor Carmich. Tristan Da Cunha: Richardson
 105 (K)

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

 Isolepis cernua var. meruensis Tanzania: Muasya 1061 (K)
 (Lye) Muasya

 Isolepis cernua var. platycarpa Australia: Coveny et al.
 (S. T. Blake) Muasya 17465 (K)

 Isolepis cernua var. setiformis S. Africa: Muasya 1194 (K)
 (Benth.) Muasya

 Isolepis costata A. Rich. Kenya: Muasya 1049 (EA, K)

 Isolepis crassiuscula Hook. f Australia: Coveny et al.
 17478 (K)

 Isolepis diabolica S. Africa: Muasya 1163 (K)
 (Stead.) Schrad.

 Isolepis digitata S. Africa: Muasya 1230 (K)
 Nees ex Schrad.

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

 Isolepis gaudichaudiana Australia: Coveny et al.
 Knuth 17476 (K)

 Isolepis graminoides Kenya: Muasya 986 (EA, K)
 (R. W. Haines & Lye) Lye

 Isolepis habra (Edgar) Sojak Australia: Coveny et al.
 17477 (NSW)

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

 Isolepis inundata R. Br. Australia: Wilson et al.
 9461 (NSW)

 Isolepis inyangensis Zimbabwe: Muasya et al.
 Muasya & Goetgh. 1125 (K)

 Isolepis keniaensis Lye Kenya: Cabolt plant 'A' (K)

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

 Isolepis ludwigii (Stead.) S. Africa: Muasya 1191 (K)

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

 Isolepis montivaga Australia: Wilson et al.
 (S. T. Blake) K.L. Wilson 9480 (K)

 Isolepis pellocolea Lesotho: Gordon-Gray
 B. L. Burtt 49694 (NU)

 Isolepis producta Australia: Wilson et al.
 (C. B. Clarke) K. L. Wilson 9510 (K)

 Isolepis prolifera (Rottb.) Australia: Coveny et al.
 R. Br. 17487 (K)

 Isolepis rubicunda (Nees) Kunth S. Africa: Muasya 1221 (K)

 Isolepis sepulcralis Steud. Australia: Coveny et al.
 17456 (K)

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

 Isolepis striata (Nees) Knuth S. Africa: Muasya 1141 (K)

 Isolepis subtillisima Boeck. Australia: Coveny et al.
 17474 (K)

 Isolepis sulcata Tristan Da Cunha:
 (Thouars) Carmich. Richardson 80 (K)

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

 Isolepis varians Steud. Chile: Pisano 259 (K)

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

 Isolepis wakefieldiana Australia: Neish et al. 110 (K)
 (S. T. Blake) K. L. Wilson

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

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

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

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

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

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

 Lipocarpha microcephala Australia: Wilson et al.
 (R. Br.) Knuth 3383 (K)

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

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

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

 Pycreus mundtii Nees Thailand: Muasya 1464 (K)

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

 Pycreus sanguinolentus Australia: Coveny et al.
 (Vahl) Nees 17461 (K)

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

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

 Scirpoides burkei S. Africa: Hargreaves 3361 (K)
 (C. B. Clarke) Goetgh.,
 Muasya & D. A. Simpson

 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 (GENT)

 Scirpus ficinioides Knuth S. Africa: Hilliard 16095 (GENT)

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

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

Dulicheae Rchb. ex
 J. Schultze-Motel

 Blysmus compressus Panz. Afghanistan: Dobson 221 (K)

 Dulichium arundinaceum USA: Goetghebeur 9914 (GENT);
 (L.) Britton Roalson et al. (2001)

Eleocharideae Goetgh.

 Chillania pusilla Roiv. Chile: Grau 1433 (GENT)

 Eleocharis atropurpurea Kenya: Muasya et al.
 (Retz.) Presl 752 (EA, K)

 Eleocharis gracilis R.Br. Australia: Wilson et al.
 9462 (K)

 Eleocharis marginulata Steud. Kenya: Muasya 1039 (EA, K)

 Eleocharis pauciflora USA: Mastrogiuseppe 7461 (WS)
 (Lightf.) Link.

Fuireneae Reichenb. ex Fenzl

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

 Bolboschoenus caldwelli Australia: Wilson et al.
 (V. Cook) Sojak 9530 (K)

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

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

 Fuirena abnormalis Tanzania: Faden et al.
 C. B. Clarke 96/118 (K)

 Fuirena ciliaris (L.) Roxb. Tanzania: Muasya 951 (EA, K)

 Fuirena coerulescens Steud. S. Africa: Muasya 2322 (K)

 Fuirena hirsuta (Berger) S. Africa: Muasya 2324 (K)
 P. L. Forbes

 Fuirena welwitschii Ridl. Kenya: Muasya 1024 (EA, K)

 Fuirena sp. Brazil: Thomas et al. 10404 (NY)

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

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

 Schoenoplectiella juncea Kenya: Muasya et al. 775 (K)
 (Willd.) Lye

 Schoenoplectiella senegalensis Kenya: Muasya et at. 2440 (EA)
 (Hochst. ex A. Rich.) Lye

 Schoenoplectus confusus Kenya: Muasya et al. 2438 (EA)
 (N. E. Br.) Lye

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

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

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

 Schoenoplectus mucronatus Thailand: Muasya et al. 1290 (K)
 (L.) Palla

 Scirpus varius Boeck. ex Botswana: Smith 5376 (NU)


 Pleurostachys sp. Brazil: Kallunki et al. 513 (NY)

 Rhynchospora alba (L.) Vahl Simpson et al. (2003)

 Rhynchospora brownii S. Africa: Verboom 616 (BOL)
 Roem. et Schult.

 Rhynchospora fascicularis Plunkett et al. (1995)
 (Michx.) Vahl

 Rhynchospora nervosa Brazil: Kallunki et al.
 (Vahl.) Boeck. 512 (NY)

Schoeneae Dumort.

 Baumea rubiginosa Australia: Wilson et al.
 (Spreng.) Boeck. 9471 (K)

 Capeobolus brevicaulis S. Africa: Verboom 646, BOL
 (C. B. Clarke) J. Browning

 Carpha alpina R. Br. Wardle et al. (2001);
 Zhang et al. (2004)

 Carpha sp. Australia: Wilson et al.
 9456 (K)

 Carpha glomerata S. Africa: Muasya 1176 (K)
 (Thunb.) Nees.

 Caustis dioica R. Br. Australia: Chase 2225 (K)

 Cladium jamaicensis Crantz Brazil: Thomas et al.
 10403 (NY)

 Cladium sp. Brazil: Mayo 259 (K)

 Cladium mariscus (L.) R. Br. Locality unknown: MJC 292 (K)

 Costularia arundinacea Zhang et al. (2004)
 (Sol. ex Vahl) Kuk.

 Costularia fragilis New Calendonia McKee
 (Daniker) Kuk. NSW41617 (K)

 Costularia nervosa Raynal Zhang et al. (2004)

 Costularia pubescens Raynal Zhang et al. (2004)

 Cyathochaeta driandra Zhang et al. (2004)
 (R. Br.) Nees

 Cyathocoma bachmannii S. Africa: Browning 835 (GENT)
 (Kuk.) C. Archer

 Cyathocoma hexandra S. Africa: Verboom 648, BOL)
 (Nees) J. Browning

 Epischoenus quadrangularis S. Africa: Verboom 636 (BOL)
 (Boeck.) C. B. Clarke

 Evandra aristata R. Br. Australia: Wilson et al. 8974

 Gahnia baniensis Benl. Malaysia: Simpson 2737 (K)

 Gahnia deusta (R. Br.) Benth. Australia: Alcock 11198 (WS)

 Gymnoschoemis Australia: Wilson et al.
sphaerocephalus 9463 (K); Zhang et al. (2004)
 (R. Br.) Hook. f.

 Lepidosperma tortuosum Australia: Coveny et al.
 F. Muell. 17470 (K); Roalson
 et al. (2001)
 Machaerina mariscoides Tahiti: Sachet 2636 (GENT)
 (Gaudich.) Kern

 Machaerina sp. New Guinea: Johns 9195 (K)

 Mesomelaena pseudostygia Australia: Chase 2226 (K)
 (Kuk.) K. L. Wilson

 Mesomelaena tetragona Australia: Chase 2227 (K)
 (R. Br.) Benth.

 Morelotia gahniiformis Hawaii: Herbst 1167 (GENT)

 Neesenbeckia punctoria S. Africa: Muasya 1214 (K)
 (Vahl) Levyns

 Oreobolus kukenthalii Steenis Malaysia: Simpson 2659 (K)

 Oreobolus obtusangulus Wardle et al. (2001)

 Oreobolus oligocephalus Zhang et al., (2004)
 W. M. Curtis

 Oreobolus pectinatus Hook. f. Wasdle et al. (2001)

 Ptilothrix deusta (R. Br.) Zhang et al. (2004)
 K. L. Wilson

 Schoenus nigricans L. Saudi Arabia: Edmondson
 3382 (K)

 Tetraria bolusii C. B. Clarke S. Africa: Verboom 606 (BOL)

 Tetraria capillaris (F. Muell.) Australia: Wilson et al.
 J. M. Black 9464 (K)

 Tetraria compacta Levyns S. Africa: Verboom 614 (BOL)

 Tetraria compar (L.) Lestib. S. Africa: Verboom 549, (BOL)

 Tetraria crassa Levyns S. Africa: Verboom 507 (BOL)

 Tetraria crinifolia (Nees) S. Africa: Verboom 638 (BOL)
 C. B. Clarke

 Tetraria microstachys S. Africa: Verboom 640 (BOL)
 (Vahl) Pfeiffer

 Tetraria thermalis (L.) S. Africa: Verboom 643 (BOL)
 C. B. Clarke

 Trianoptiles solitaria Zhang et al. (2004)
 (C. B. Clarke) Levyns

 Tricostularia pauciflora Australia: Coveny et al.
 (R. Br.) Benth. 17484 (K); Zhang
 et al. (2004)
Scirpeae Kunth ex Dumort.

 Amphiscirpus nevadensis Argentina: Charpin et al. 20575
 (S. Watson) Oteng-Yeboa (GENT)

 Eriophorum angustifolium Simpson et al. (2003)

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

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

 Oreoholopsis clementis Dhooge (2005)
 (M. E. Jones) Dhooge &

 Oreoholopsis inversa Ecuador: Laegaard 21492 (GENT)
 Dhooge & Goetgh.

 Oreoholopsis tepalifera Dhooge et al. (2003)
 T. Koyama & Guagl.
 Phylloscripus acaulis (Phil.) Dhooge et al. 2003
 Goetgh. & D. A. Simpson

 Phylloscirpus boliviensis Ecuador: Laegaard 102805 (GENT)
 (Barros) Dhooge & Goetgh.

 Phylloscirpus deserticola Ecuador: Laegaard et al.
 (Phil.) Dhooge & Goetgh. 21478 (GENT)

 Scirpus ancistrochaetus USA: Naczi 7544 (DOV)

 Scirpus atrocinctus Fernald USA: Naczi 10456 (DOV)

 Scirpus cyperinus (L.) Kunth USA: Naczi (DOV)

 Scirpus expansus Fernald USA: Naczi 10050 (DOV)

 Scirpus flaccidifolius USA: Naczi 9774 (DOV)
 (Fernald) Schuyler

 Scirpus georgianus Harper USA: Naczi 10458 (DOV)

 Scirpus hattorianus Makino USA: Naczi 10369 (DOV)

 Scirpus pendulus Muhl. USA: Naczi 10394 (DOV)

 Scirpus polystachyus F. Muell. Australia: Pullen 4091 (K)

 Scirpus radicans Schkuhr Czechia: Goetghebeur
 9882 (GENT)

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

 Scirpus ternatanus Hong Kong: Shaw 917 (K)
 Reinw. ex Miq.

 Trichophorum alpinum CANADA: Waterway
 (L.) Pers. 2002.95 (GENT)

 Trychophorum caespitosum British Isles: Nelmes 954 (K)
 (L.) Hartm.
 Trichophorum clintonii Gray Canada: Baldwin 4856 (K)

 Trichophorum planifolium USA: Dhooge 24 (GENT)
 (Spreng.) Palla

 Trichophorum pumilum Uncertain locality: Morse &
 (Vahl) Schinz & Thellung Jordon 2272 (GENT)

 Trichophorum rigidum Argentina: Renvoize et al.
 (Steud.) Goetgh., Muasya & 5021 (K)
 D. A. Simpson subsp. rigidum

 Trichophorum rigidum Ecuador: Laegaard et al.
 subsp. ecuadoriensis 21574 (GENT)
 Dhooge & Goetgh.

 Trichophorum subcapitatum Papua New Guinea: Goetghebeur
 (Thwaites & Hook.) et al. 6581 (GENT)
 D. A. Simpson

 Zameioscirpus atacamensis Bolivia: Ruthsatz & Budde
 (Phil.) Dhooge & Goetgh. 10328 (Trier)

 Zameioscirpus gaimardioides Argentina: Ruthsatz 9212 (gent);
 (E. Desv.) Dhooge & Goetgh. Dhooge et al. (2003)

 Zameioscirpus muticus Dhooge et al. (2003)
 Dhooge & Goetgh.

Sclerieae Kunth ex Fenzl

 Scleria distans Poir. Kenya: Muasya 1023 (EA, K)

 Scleria foliosa A. Rich. Tanzania: Muasya 939 (EA, K)

 Scleria terrestris (L.) Fassett Malaysia: Simpson 2658 (K)

Trilepideae Goetgh.

 Coleochloa abyssinica Ethiopia: Vollesen 80/2 (K)
 (A. Rich.) Gilly

 Microdracoides squamosus Hua Bonn Ace. 150

 Trilepis lhotzkiana Nees Bonn Ace. s.n.

Mapanioideae C. B. Clarke

Chrysitricheae Lestib. ex Fenzl

 Capitularia, foliata Uitt. Indonesia: Johns 8725 (K)

 Capitularina involucrata Simpson et al. (2003)
 (J. V. Suringar) Kern

 Chorizandra cymbaria R. Br. Bremer (2002)

 Chorizandra enodis Nees Bremer (2002)

 Chorizandra sphaerocephala Simpson et al. (2003)
 R. Br.

 Chrysitrix capensis L. S. Africa: Muasya 1242 (K)

 Exocarya sclerioides Simpson et al. (2003)
 (F. Muell.) Benth.

 Lepironia articulata Malaysia: Simpson 1236 (K)
 (Retz.) Domin.

Hypolytreae Presl ex Fenzl

 Diplasia karatifolia Simpson et al. (2003)
 Rich. ex Pers.

 Hypolytrum bullatum Brazil: Thomas et al.
 C. B. Clarke 10318 (NY)

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

 Hypolytrum testui Cherm. Simpson et al. (2003)

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

 Mapania lorea Uitt. Simpson et al. (2003)

 Mapania meditensis Brunei: Simpson et al. 2515 (K)
 D. A. Simpson

 Mapania tenuiscapa Simpson et al. (2003)
 C. B. Clarke

 Scirpodendron bogneri Malaysia: Simpson
 S.S. Hooper 2650 (K)

 Scirpodendron ghaeri Simpson et al. (2003)
 (Gaertn.) Merrill


 Juncus effusus L. Simpson et al. (2003);
 Chase et al., 1993

 Juncus gerardii Loisel. Simpson et al. (2003);
 Drabbkova et al.

 Luzula multiflora (Retz.) Lej. Bremer (2002); Simpson
 et al. (2003)

 Luzula sylvatica (Huds.) Gaud. Simpson et al. (2003)

 Prionium serratum Drege S. Africa: Gettliffe Norris,
 s.n. (NBG)

Taxon Genbank accession

 rbcL trnL-F OR
Cyperoideae Suess.

Abildgaardieae Lye

 Abildgaardia ovata Y12985 AJ295754
 (Burm. f.) Kral

 Actinoschoenus repens Raynal EF178537

 Arthrostylis aphylla R. Br. AY725939

 Bulbostylis atrosanguinea Y12992
 (Boeck.) C. B. Clarke

 Bulbostylis hispidula Y12944
 (Vahl) R. W. Haines

 Crosslandia setifolia W. Fitzg. EF178538 EF178592

 Fimbristylis complanata Y13009
 (Retz.) Link

 Fimbristylis dichotoma Y13008 AJ295755
 (L.) Vahl

 Nemum spadiceum (Lam.) Y12945
 Desv. ex Ham.

 Trachystylis stradbrokensis EF178539 EF178591
 (Domin) Kuk.

Bisboeckelereae Pax ex
 L.T. Eiten

 Becquerelia cymosa Brongn. Y12948

 Calyptrocarya bicolor EF178540
 (H. Pfeiff.) Koyama

 Diplacrum africanum AY725942
 C. B. Clarke

Cariceae Kunth ex Dumort.

 Carex cephalophora DQ006089
 Muhl. ex Willd.

 Carex conferta A. Rich. Y12999

 Carex echinochloe Kunze Y12997 AF191818

 Carex hostiana DC. L12672

 Carex monostachya A. Rich. Y12998

 Carex sylvatica Huds. AY344175

 Kobresia simpliciuscula U49232 AF164948
 (Wahlenb.) Mackenzie

 Uncinia nemoralis AY725956
 K. L. Wilson

 Schoenoxiphium ecklonii Nees EF178541

 Schoenoxiphium lehmannii EF178542
 (Nees) Kunth ex Steud.

 Schoenoxiphium sparteum EF178543
 (Wahlenb.) C.B.Clarke

Uncertain tribe aff. Cariceae

 Khaosokia caricoides AY725948 EF178535
 D.A.Simpson, Chayam. &

Cryptangieae Benth.

 Didyndandrum stellatum EF178544
 (Boeck.) Gilly

 Exochogyne amazonica EF178545
 C. B. Clarke

 Lagenoearpus alboniger aY725949
 (A. St.-Hil.) C. B. Clarke

Cypereae Dumort.

 Alinula paradoxa aJ278290 aJ295756
 Goetgh. & Vorster

 Androtrichum giganteum EF178546
 (Kunth) H. Pfeiff.

 Androtrichum trigynum EF178547
 (Spreng.) H. Pfeiff.

 Ascolepis capensis Y13003 AJ295757
 (Kunth) Ridl.

 Ascolepis protea Welw. Y13002

 Courtoisina assimilis AY40590 AY040595
 (Steud.) Maquet

 Cyperus compressus L. AF449506 AF449555/

 Cyperus congestus Vahl AF449507 AF449556/

 Cyperus cuspidatus Kunth AF449508 AF449557/

 Cyperus cyperoides (L.) Kuntze AF449509 AF449558/

 Cyperus dichroostachyus Y12965 /AF449571
 A. Rich.

 Cyperus endlichii Kuk. AF449510 AF449559/

 Cyperus involucratus Rottb. Y12967 AJ295758

 Cyperus kerstenii Boeck. Y13018 AY040597

 Cyperus laevigatus L. Y13017 AY040596

 Cyperus longus L. Y13015 AY040598

 Cyperus meeboldii Kuk. AF449511 AF449560/

 Cyperus papyrus L. Y12966 AJ295759

 Cyperus plateilema AF449512 AF449561/
 (Steud.) Kuk. AF449573

 Cyperus pseudovestitus AF449513 AF449562/
 (C. B. Clarke) Kuk. AF449574

 Cyperus pulchellus R. Br. AY40591 AY040599

 Cyperus pygmaeus Rottb. AJ404698 AJ295760

 Cyperus rigidifolius Steud. Y13016 AY040600

 Desmoschoenus spiralis AJ404701 AJ295753
 Hook. f.

 Ficinia bergiana Kunth EF200588 EF178593

 Ficinia distans C. B. Clarke EF178548 EF179594

 Ficinia esterhuyseniae Muasya EF178549 EF178590

 Ficinia gracilis Schrad. EF178599 EF178595

 Ficinia gracilis Schrad. EF178550 EF178534

 Ficinia gydomontana T. H. EF178551 EF178596
 Arnold & K. D. Gordon-Gray

 Ficinia indica EF178552 EF178597
 (Lam.) H. Pfeiff.

 Ficinia laciniata (Thunb.) Nees EF178553 EF178598

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

 Ficinia paradoxa (Schrad.) Nees DQ058354 DQ058317

 Ficinia pinguior C. B. Clarke AJ404703 AJ295772

 Ficinia polystachya Levyns EF178554 EF178599

 Ficinia ramosissima Kunth EF178555 EF178600

 Ficinia repens Kunth EF178556 EF178601

 Ficinia rigida Levyns EF178557 EF178602

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

 Ficinia tristachya AJ404702 AJ295771
 (Rottb.) Nees

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

 Isolepis aucklandica Hook. f AJ404704 AJ295773

 Isolepis bicolor Carmich. AJ404705 AJ295774

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

 Isolepis cernua var. meruensis AJ404715 AJ295791
 (Lye) Muasya

 Isolepis cernua var. platycarpa AJ404716 AJ295794
 (S. T. Blake) Muasya

 Isolepis cernua var. setiformis AJ404725 AJ295805
 (Benth.) Muasya

 Isolepis costata A. Rich. Y12981 AJ295776

 Isolepis crassiuscula Hook. f AJ404706 AJ295777

 Isolepis diabolica AJ404707 AJ295778
 (Stead.) Schrad.

 Isolepis digitata AJ404708 AJ295779
 Nees ex Schrad.

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

 Isolepis gaudichaudiana AJ404709 AJ295781

 Isolepis graminoides Y12960 AJ295782
 (R. W. Haines & Lye) Lye

 Isolepis habra (Edgar) Sojak AJ404710 AJ295783

 Isolepis hystrix (Thunb.) Nees AJ404711 AJ295785

 Isolepis inundata R. Br. AJ404712 AJ295786

 Isolepis inyangensis AJ297506 AJ295787
 Muasya & Goetgh.

 Isolepis keniaensis Lye Y12980 AJ295788

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

 Isolepis ludwigii (Stead.) AJ404713 A3295799

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

 Isolepis montivaga AJ297507 AJ295792
 (S. T. Blake) K.L. Wilson

 Isolepis pellocolea AJ404729 AJ297514
 B. L. Burtt

 Isolepis producta AJ404717 AJ295795
 (C. B. Clarke) K. L. Wilson

 Isolepis prolifera (Rottb.) AJ404718 AJ295796
 R. Br.

 Isolepis rubicunda (Nees) Kunth AJ404719 AJ295797

 Isolepis sepulcralis Steud. AJ404720 AJ295798

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

 Isolepis striata (Nees) Knuth AJ404721 AJ295801

 Isolepis subtillisima Boeck. AJ297509 AJ295800

 Isolepis sulcata AJ404722 AJ295802
 (Thouars) Carmich.

 Isolepis tenuissima AY725947
 (Nees) Kunth

 Isolepis varians Steud. AJ404723 AJ295803

 Isolepis venustula Kunth AJ404724 AJ295804

 Isolepis wakefieldiana AJ404726 AJ295806
 (S. T. Blake) K. L. Wilson

 Kyllinga appendiculata Y13007 AJ295761
 K. Schum.

 Kyllinga brevifolia Rottb. AF449515 AF449564/

 Kyllinga bulbosa P. Beauv. Y12979 AY040601

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

 Kyllingiella polyphylla Y13013 AJ295515
 (A. Rich.) Lye

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

 Lipocarpha microcephala Y12991
 (R. Br.) Knuth

 Lipocarpha nana (A.Rich.) Y12990 AJ295762
 J. Raynal

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

 Pycreus flavescens (L.) Rchb. Y13005 AJ295763

 Pycreus mundtii Nees AF449517 AF449566/

 Pycreus nuerensis Y13004 AY040603
 (Boeck.) S. S. Hooper

 Pycreus sanguinolentus AF449567/
 (Vahl) Nees AF449579

 Queenslandiella hyalina AY725953
 (Vahl) Ballard

 Remirea maritima Aubl. AY040593 AY040604

 Scirpoides burkei Y13001 AJ295810
 (C. B. Clarke) Goetgh.,
 Muasya & D. A. Simpson

 Scirpoides holoschoenus Y12994 AJ295811
 (L.) Sojak

 Scirpoides thunbergii AJ404727 AJ295812
 (Schrad.) Sojak

 Scirpus falsus C. B. Clarke EF178559

 Scirpus ficinioides Knuth EF178560

 Sphaerocyperus erinaceus AJ404699 AJ295764
 (Ridl.) Lye

 Volkiella disticha EF178561
 Merxm. & Czech

Dulicheae Rchb. ex
 J. Schultze-Motel

 Blysmus compressus Panz. AJ404700 AJ295766

 Dulichium arundinaceum AY725943 AF285067
 (L.) Britton

Eleocharideae Goetgh.

 Chillania pusilla Roiv. EF178562

 Eleocharis atropurpurea Y13012
 (Retz.) Presl

 Eleocharis gracilis R.Br. EF178563

 Eleocharis marginulata Steud. Y13011 AJ295768

 Eleocharis pauciflora U49229
 (Lightf.) Link.

Fuireneae Reichenb. ex Fenzl

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

 Bolboschoenus caldwelli EF178564
 (V. Cook) Sojak

 Bolboschoenus maritimus Y12996 AJ295767
 (L.) Palla

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

 Fuirena abnormalis EF179565
 C. B. Clarke

 Fuirena ciliaris (L.) Roxb. Y12971

 Fuirena coerulescens Steud. EF178566

 Fuirena hirsuta (Berger) EF178567
 P. L. Forbes

 Fuirena welwitschii Ridl. Y12993 EF178605

 Fuirena sp. Y12970

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

 Schoenoplectiella articulata Y12987
 (L.) Lye

 Schoenoplectiella juncea Y12952
 (Willd.) Lye

 Schoenoplectiella senegalensis EF178568 EF178606
 (Hochst. ex A. Rich.) Lye

 Schoenoplectus confusus EF178569
 (N. E. Br.) Lye

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

 Schoenoplectus lacustris Y12943 AJ295809
 (L.) Palla

 Schoenoplectus litoralis EF178571
 (Schrad.) Palla

 Schoenoplectus mucronatus EF178572
 (L.) Palla

 Scirpus varius Boeck. ex EF178573


 Pleurostachys sp. Y12989

 Rhynchospora alba (L.) Vahl AY344174

 Rhynchospora brownii DQ058353 DQ058316
 Roem. et Schult.

 Rhynchospora fascicularis U49233
 (Michx.) Vahl

 Rhynchospora nervosa Y12977
 (Vahl.) Boeck.

Schoeneae Dumort.

 Baumea rubiginosa AY725940
 (Spreng.) Boeck.

 Capeobolus brevicaulis DQ058343 DQ058303
 (C. B. Clarke) J. Browning

 Carpha alpina R. Br. AF307909 AY230012

 Carpha sp. EF178574

 Carpha glomerata AY725941
 (Thunb.) Nees.

 Caustis dioica R. Br. Y12976

 Cladium jamaicensis Crantz Y12988

 Cladium sp. Y12950

 Cladium mariscus (L.) R. Br. DQ058338 DQ058298

 Costularia arundinacea -- AY230036
 (Sol. ex Vahl) Kuk.

 Costularia fragilis EU828589
 (Daniker) Kuk.

 Costularia nervosa Raynal -- AY230032

 Costularia pubescens Raynal -- AY230037

 Cyathochaeta driandra -- AY230042
 (R. Br.) Nees

 Cyathocoma bachmannii EF200590 EF178604
 (Kuk.) C. Archer

 Cyathocoma hexandra DQ058344 DQ058304
 (Nees) J. Browning

 Epischoenus quadrangularis DQ058349 DQ058311
 (Boeck.) C. B. Clarke

 Evandra aristata R. Br. AY725944

 Gahnia baniensis Benl. DQ058342 DQ058302

 Gahnia deusta (R. Br.) Benth. U49231

 Gymnoschoemis AY725945 AY230033
 (R. Br.) Hook. f.

 Lepidosperma tortuosum AY725950 AF285074
 F. Muell.

 Machaerina mariscoides EF178575
 (Gaudich.) Kern

 Machaerina sp. DQ058340 DQ058300

 Mesomelaena pseudostygia Y12959 DQ058301
 (Kuk.) K. L. Wilson

 Mesomelaena tetragona Y12949
 (R. Br.) Benth.

 Morelotia gahniiformis EF178576

 Neesenbeckia punctoria AY725952 DQ058306
 (Vahl) Levyns

 Oreobolus kukenthalii Steenis Y12972 EF178536

 Oreobolus obtusangulus AF307926

 Oreobolus oligocephalus -- AY230031
 W. M. Curtis

 Oreobolus pectinatus Hook. f. AF307927

 Ptilothrix deusta (R. Br.) -- AY230041
 K. L. Wilson

 Schoenus nigricans L. Y12983 AJ295814

 Tetraria bolusii C. B. Clarke -- DQ058315

 Tetraria capillaris (F. Muell.) EF178577
 J. M. Black

 Tetraria compacta Levyns DQ058351 DQ058313

 Tetraria compar (L.) Lestib. DQ058350 DQ058312

 Tetraria crassa Levyns DQ058352 DQ058314

 Tetraria crinifolia (Nees) DQ058348 DQ058309
 C. B. Clarke

 Tetraria microstachys DQ058347 DQ058307
 (Vahl) Pfeiffer

 Tetraria thermalis (L.) -- DQ058308
 C. B. Clarke

 Trianoptiles solitaria -- AY230028
 (C. B. Clarke) Levyns

 Tricostularia pauciflora AY725954 AY230038
 (R. Br.) Benth.

Scirpeae Kunth ex Dumort.

 Amphiscirpus nevadensis DQ317926 DQ317925
 (S. Watson) Oteng-Yeboa

 Eriophorum angustifolium AY344177

 Eriophorum vaginatum L. Y12951 AJ295769

 Eriophorum viridicarinatum U49230
 (Engl.) Fern.

 Oreoholopsis clementis AJ811011 Dhooge
 (M. E. Jones) Dhooge & (2005)

 Oreoholopsis inversa AJ811009 DQ317923
 Dhooge & Goetgh.

 Oreoholopsis tepalifera AJ575932 AJ576035
 T. Koyama & Guagl.
 Phylloscripus acaulis (Phil.) AJ575926 AJ576029
 Goetgh. & D. A. Simpson

 Phylloscirpus boliviensis AJ566081 AJ566082
 (Barros) Dhooge & Goetgh.

 Phylloscirpus deserticola AJ704785 AJ704786
 (Phil.) Dhooge & Goetgh.

 Scirpus ancistrochaetus EF178578

 Scirpus atrocinctus Fernald EF178579

 Scirpus cyperinus (L.) Kunth EF178580

 Scirpus expansus Fernald EF178581

 Scirpus flaccidifolius EF178582
 (Fernald) Schuyler

 Scirpus georgianus Harper EF178583

 Scirpus hattorianus Makino EF178584

 Scirpus pendulus Muhl. EF178585

 Scirpus polystachyus F. Muell. Y12974 AJ295813

 Scirpus radicans Schkuhr AJ811012 Dhooge

 Scirpus sylvatictis L. EF178586

 Scirpus ternatanus EF178587
 Reinw. ex Miq.

 Trichophorum alpinum AJ810999 DQ317924
 (L.) Pers.

 Trychophorum caespitosum Y12969 Dhooge
 (L.) Hartm. (2005)
 Trichophorum clintonii Gray Y12982 Dhooge
 Trichophorum planifolium AJ811001 Dhooge
 (Spreng.) Palla (2005)

 Trichophorum pumilum AJ811000 Dhooge
 (Vahl) Schinz & Thellung (2005)

 Trichophorum rigidum AJ297509 AJ295908
 (Steud.) Goetgh., Muasya &
 D. A. Simpson subsp. rigidum

 Trichophorum rigidum AJ811008 Dhooge
 subsp. ecuadoriensis (2005)
 Dhooge & Goetgh.

 Trichophorum subcapitatum AJ811006 Dhooge
 (Thwaites & Hook.) (2005)
 D. A. Simpson

 Zameioscirpus atacamensis AJ575929 AJ576032
 (Phil.) Dhooge & Goetgh.

 Zameioscirpus gaimardioides AJ575938 AJ576031
 (E. Desv.) Dhooge & Goetgh.

 Zameioscirpus muticus AJ575927 AJ576030
 Dhooge & Goetgh.

Sclerieae Kunth ex Fenzl

 Scleria distans Poir. Y12968 DQ058299

 Scleria foliosa A. Rich. Y12986

 Scleria terrestris (L.) Fassett Y12947

Trilepideae Goetgh.

 Coleochloa abyssinica Y12975
 (A. Rich.) Gilly

 Microdracoides squamosus Hua AY725951

 Trilepis lhotzkiana Nees AY725955

Mapanioideae C. B. Clarke

Chrysitricheae Lestib. ex Fenzl

 Capitularia, foliata Uitt. EF178588

 Capitularina involucrata AY344168
 (J. V. Suringar) Kern

 Chorizandra cymbaria R. Br. AJ419940

 Chorizandra enodis Nees AJ419939

 Chorizandra sphaerocephala AY344170
 R. Br.

 Chrysitrix capensis L. AJ419938 AY344171

 Exocarya sclerioides AY344167
 (F. Muell.) Benth.

 Lepironia articulata Y12957 AY344169
 (Retz.) Domin.

Hypolytreae Presl ex Fenzl

 Diplasia karatifolia AY344166
 Rich. ex Pers.

 Hypolytrum bullatum Y12956
 C. B. Clarke

 Hypolytrum nemorum Y12958 AJ295816
 (Vahl) Spreng.

 Hypolytrum testui Cherm. AY344163

 Mapania cuspidata Y12955 AJ295817
 (Miq.) Uittien

 Mapania lorea Uitt. AY344161

 Mapania meditensis Y12954 AY344160
 D. A. Simpson

 Mapania tenuiscapa AY344162
 C. B. Clarke

 Scirpodendron bogneri Y12946 AY344164
 S.S. Hooper

 Scirpodendron ghaeri AY344165
 (Gaertn.) Merrill


 Juncus effusus L. L12681 AY344156

 Juncus gerardii Loisel. AY216613 AY344157

 Luzula multiflora (Retz.) Lej. AJ419945 AY344158

 Luzula sylvatica (Huds.) Gaud. AY216637 AY344159

 Prionium serratum Drege U49223 AY344155
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Author:Muasya, A. Muthama; Simpson, David A.; Verboom, G. Anthony; Goetghebeur, Paul; Naczi, Robert F.C.; C
Publication:The Botanical Review
Article Type:Report
Geographic Code:1USA
Date:Mar 1, 2009
Previous Article:Foreword.
Next Article:A preliminary molecular phylogeny of the Rhynchosporeae (Cyperaceae).

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