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


Cyperaceae comprise 109 genera genera, in taxonomy: see classification.  and approximately 5,500 species and have an almost cosmopolitan distribution In biogeography, a biological category of living things is said to have cosmopolitan distribution if this category can be found almost anywhere around the world. See "cosmopolitan" for etymology.

An example of a cosmopolitan species is the Painted Lady butterfly.
 (Govaerts et al., 2007). About 35% of the genera are monotypic monotypic

said of a genus with only one species.
, 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 flo·ral  
Of, relating to, or suggestive of a flower: a fabric with a floral pattern.

 morphology morphology

In biology, the study of the size, shape, and structure of organisms in relation to some principle or generalization. Whereas anatomy describes the structure of organisms, morphology explains the shapes and arrangement of parts of organisms in terms of such
, and the majority of the smaller genera are carved carve  
v. carved, carv·ing, carves
a. To divide into pieces by cutting; slice: carved a roast.

 out of the larger ones on the basis on one or few distinguishing features.

Family level phylogenetic phy·lo·ge·net·ic
1. Of or relating to phylogeny or phylogenetics.

2. Relating to or based on evolutionary development or history.
 studies in the last ten years have used morphological mor·phol·o·gy  
n. pl. mor·phol·o·gies
a. The branch of biology that deals with the form and structure of organisms without consideration of function.

 (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 anatomy (ənăt`əmē), branch of biology concerned with the study of body structure of various organisms, including humans. Comparative anatomy is concerned with the structural differences of plant and animal forms.  and embryology embryology

Study of the formation and development of an embryo and fetus. Before widespread use of the microscope and the advent of cellular biology in the 19th century, embryology was based on descriptive and comparative studies.
) 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 cir·cum·scrip·tion  
1. The act of circumscribing or the state of being circumscribed.

2. Something, such as a limit or restriction, that circumscribes.

3. A circumscribed space or area.

, 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 DNA sequence Genetics The precise order of bases–A,T,G,C–in a segment of DNA, gene, chromosome, or an entire genome. See Base pair, Base sequence analysis, Chromosome, Gene, Genome.  data are increasingly used in angiosperm angiosperm (ăn`jēəspûrm'), term denoting seed plants in which the ovules, or young seeds, are enclosed within the ovary (that part of the pistil specialized for seed production), in contrast to the gymnosperms, in which the seeds  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 plas·tid
1. Any of several pigmented cytoplasmic organelles found in plant cells and other organisms, having various physiological functions, such as the synthesis and storage of food. Also called trophoplast.

 and nuclear regions have been used. The plastid regions rps16 intron Intron

In split genes, a portion that is included in ribonucleic acid (RNA) transcripts but is removed from within a transcript during RNA processing and is rapidly degraded.
, trnL intron and trnL-F intergenic spacer have been used in studies of subfamily subfamily /sub·fam·i·ly/ (sub´fam-i-le) a taxonomic division between a family and a tribe.

A taxonomic category ranking between a family and a genus.
 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 re·con·struct  
tr.v. re·con·struct·ed, re·con·struct·ing, re·con·structs
1. To construct again; rebuild.

 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 align (līn),
v to move the teeth into their proper positions to conform to the line of occlusion.
 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 DNA: see nucleic acid.
 or deoxyribonucleic acid

One of two types of nucleic acid (the other is RNA); a complex organic compound found in all living cells and many viruses. It is the chemical substance of genes.
 was extracted from vegetative vegetative /veg·e·ta·tive/ (vej?e-ta?tiv)
1. of, pertaining to, or characteristic of plants.

2. concerned with growth and nutrition, as opposed to reproduction.

 material (leaves or culms culms

the shoots and roots of sprouted grains in the brewing process. These are removed during brewing and are salvaged as a food supplement. Called also malt culms.
) collected in the field or from herbarium herbarium, collection of dried and mounted plant specimens used in systematic botany. To preserve their form and color, plants collected in the field are spread flat in sheets of newsprint and dried, usually in a plant press, between blotters or absorbent paper.  specimens (Table 1). DNA extraction DNA extraction is a routine procedure to collect DNA for subsequent molecular or forensic analysis. Outline of a DNA extraction
There are three basic steps in a DNA extraction, the details of which may vary depending on the type of sample and any substances that may
, amplification amplification /am·pli·fi·ca·tion/ (33000) (am?pli-fi-ka´shun) the process of making larger, such as the increase of an auditory stimulus, as a means of improving its perception.  and sequencing were performed according to according to
1. As stated or indicated by; on the authority of: according to historians.

2. In keeping with: according to instructions.

 published procedures (e.g. Muasya et al., 2002); the resulting sequences were aligned manually and are lodged with GenBank (Table 1).

Heuristic A method of problem solving using exploration and trial and error methods. Heuristic program design provides a framework for solving the problem in contrast with a fixed set of rules (algorithmic) that cannot vary.

 analyses were carried out using PAUP PAUP Phylogenetic Analysis Using Parsimony * (Swofford, 2002). Searches were conducted under Fitch fitch: see polecat.  (1971) parsimony par·si·mo·ny  
1. Unusual or excessive frugality; extreme economy or stinginess.

2. Adoption of the simplest assumption in the formulation of a theory or in the interpretation of data, especially in accordance with the rule of
, TBR TBR Tennessee Board of Regents
TBR Technology Business Research
TBR To Be Read
TBR Travel Business Roundtable
TBR To Be Resolved
TBR To Be Reviewed
TBR Technical Basis for Regulation
TBR To Be Recorded
TBR Total Business Return
TBR To Be Revised
 (tree-bissection-reconnection) branch swapping, and random taxon taxon (pl. taxa), in biology, a term used to denote any group or rank in the classification of organisms, e.g., class, order, family.  addition (5,000) with the MulTrees option in effect and retaining only ten trees per replicate rep·li·cate
1. To duplicate, copy, reproduce, or repeat.

2. To reproduce or make an exact copy or copies of genetic material, a cell, or an organism.

A repetition of an experiment or a procedure.
. Internal support was estimated using 1000 bootstrap See boot.

(operating system, compiler) bootstrap - To load and initialise the operating system on a computer. Normally abbreviated to "boot". From the curious expression "to pull oneself up by one's bootstraps", one of the legendary feats of Baron von Munchhausen.
 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 par·si·mo·ni·ous  
Excessively sparing or frugal.

 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 monophyletic /mono·phy·let·ic/ (mon?o-fi-let´ik) descended from a common ancestor or stem cell.

1. Descended or derived from one original stock or source.
 and sister to Juncaceae, with Mapanioideae sister to all the other Cyperaceae (Fig. 1; tribes and subfamilies sensu Goetghebeur, 1998). Within the last clade clade Cladus, subtype Genetics A branch of biological taxa or species that share features inherited from a common ancestor; a single phylogenetic group or line. See Inheritance, Species. , 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 Inserted into. See embedded system.  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 bisexual /bi·sex·u·al/ (-sek´shoo-al)
1. pertaining to or characterized by bisexuality.

2. an individual exhibiting bisexuality.

3. pertaining to or characterized by hermaphroditism.

 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 en·close   also in·close
tr.v. en·closed, en·clos·ing, en·clos·es
1. To surround on all sides; close in.

2. To fence in so as to prevent common use: enclosed the pasture.
 by a utricle utricle /utri·cle/ (u´tri-k'l)
1. any small sac.

2. the larger of the two divisions of the membranous labyrinth of the internal ear.
. 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 in·sel·berg  
See monadnock.

[German : Insel, island (from Middle High German insule, from alteration of Old High German isila, probably from Vulgar Latin
 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 genus, in taxonomy: see classification.

Biological classification. It ranks below family and above species, consisting of structurally or phylogenetically (see
 unplaced in any tribe tribe [Lat., tribus: the tripartite division of Romans into Latins, Sabines, and Etruscans], a social group bound by common ancestry and ties of consanguinity and affinity; a common language and territory; and characterized by a political and economic  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 [Latin, Among other things.] A phrase used in Pleading to designate that a particular statute set out therein is only a part of the statute that is relevant to the facts of the lawsuit and not the entire statute. , 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 Adj. 1. widely distributed - growing or occurring in many parts of the world; "a cosmopolitan herb"; "cosmopolitan in distribution"

bionomics, environmental science, ecology - the branch of biology concerned with the relations between organisms
 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 seg·re·gate  
v. seg·re·gat·ed, seg·re·gat·ing, seg·re·gates
1. To separate or isolate from others or from a main body or group. See Synonyms at isolate.

 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 paraphyletic  

Relating to a taxonomic group that includes some but not all of the descendants of a common ancestor. In the traditional taxonomy of vertebrates, where fish are a separate class from the classes of terrestrial vertebrates, the class of fish is
 with several tropical African perennial perennial, any plant that under natural conditions lives for several to many growing seasons, as contrasted to an annual or a biennial. Botanically, the term 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 Similarity is some degree of symmetry in either analogy and resemblance between two or more concepts or objects. The notion of similarity rests either on exact or approximate repetitions of patterns in the compared items.  with Schoeneae (e.g. one- to few-flowered spikelet and wide glume wings enclosing en·close   also in·close
tr.v. en·closed, en·clos·ing, en·clos·es
1. To surround on all sides; close in.

2. To fence in so as to prevent common use: enclosed the pasture.
 the next flower). On the other hand, Actinoschoenus repens is a Zambian endemic endemic /en·dem·ic/ (en-dem´ik) present or usually prevalent in a population at all times.

, with morphological similarity to both Abildgaardieae and Schoeneae. Although these three taxa had been placed in Schoeneae even with decisive anatomical anatomical /ana·tom·i·cal/ (an?ah-tom´i-kal) pertaining to anatomy, or to the structure of an organism.

an·a·tom·i·cal or an·a·tom·ic
1. Concerned with anatomy.

 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 embryo (ĕm`brēō), name for the developing young of an animal or plant. In its widest definition, the embryo is the young from the moment of fertilization until it has become structurally complete and able to survive as a separate organism.  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 South Africa, Afrikaans Suid-Afrika, officially Republic of South Africa, republic (2005 est. pop. 44,344,000), 471,442 sq mi (1,221,037 sq km), S 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 e·lu·ci·date  
v. e·lu·ci·dat·ed, e·lu·ci·dat·ing, e·lu·ci·dates
To make clear or plain, especially by explanation; clarify.

To give an explanation that serves to clarify.
 phylogenetic relationships within Cryptangineae, Bisboeckelerieae, Fuireneae, Schoeneae, and Sclerieae.

Acknowledgements A number of the contributors acknowledge funding from various sources. AMM AMM Autorisation de Mise sur le Marche (French)
AMM Autorisation de Mise sur le Marché (French: Commission of Marketing Authorization)
AMM ASEAN Ministerial Meeting
AMM American Metal Market
 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 Coordinates:
“UCT” redirects here. For other uses, see UCT (disambiguation).
 (Smuts Fellowship); and the Norwegian Council of Universities' Committee for Development Research and Education (NUFU NUFU National Urban Forestry Unit (UK)  project 53/03). GAV GAV Gateway Anti-Virus (Sonicwall)
GAV Gross Asset Value
GAV Great American Volleyball
GAV Giubbotto Assetto Variabile (Italian: life jacket)
GAv Gatha-Avestan (linguistics) 
 acknowledges funding from the National Research Council, South Africa.

Open Access This article is distributed under the terms of the Creative Commons An organization that has defined an alternative to copyrights by filling in the gap between full copyright, in which no use is permitted without permission, and public domain, where permission is not required at all.  Attribution at·tri·bu·tion  
1. The act of attributing, especially the act of establishing a particular person as the creator of a work of art.

 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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gem, commonly, a mineral or organic substance, cut and polished and used as an ornament. Gems also are used as seals (items of assurance) and as talismans (good-luck charms). For birthstones, see month.
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A synthetic discipline that describes the distributions of living and fossil species of plants and animals across the Earth's surface as consequences of ecological and evolutionary processes.
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n. Scots
1. A waterfall.

2. A steep ravine.

[Scottish Gaelic linne, pool, waterfall.]
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n. pl. sub·gen·e·ra
An occasionally used taxonomic category ranking between a genus and a species.

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1. Of, relating to, or characteristic of a particular district.

2. Composed of or divided into component sections.

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In biology, the classification of organisms into a hierarchy of groupings, from the general to the particular, that reflect evolutionary and usually morphological relationships: kingdom, phylum, class, order,
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This group of flowering plants (angiosperms), with one seed leaf, was previously thought to be one of the two major categories of flowering plants (the other group is dicotyledons).
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1. Of or relating to plants or plant life.

2. Of or relating to the science of botany.

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GMT - Universal Time 1

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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 Cape Town or Capetown, city (1991 pop. 854,616), legislative capital of South Africa and capital of Western Cape, a port on the Atlantic Ocean. It was the capital of Cape Province before that province's subdivision in 1994. , South Africa

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

(3) Department of Biology, Ghent University It is a relatively young university, founded 9 October 1817. The year before, king William I of the Netherlands had proclaimed the establishment of three universities in the Southern Netherlands. , K.L. Ledegancksraat 35, BE-9000 Ghent, Belgium

(4) Department of Agriculture & Natural Resources, Delaware State University Delaware State University (DSU), the second-largest university in the state of Delaware, is a historically black university. Over the last 116 years, it has evolved into a fully accredited, comprehensive university with a main campus located in Dover, Delaware and two satellite , 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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