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CYPERACEAE<br />

507<br />

become split by growth <strong>of</strong> culm), with or without a scaly ring or fringe <strong>of</strong> hair (ligule) at junction<br />

<strong>of</strong> sheath and blade on upper (inner) side, and a usually elongate blade (leaves all reduced<br />

<strong>to</strong> inconspicuous sheaths in Eleocharis and some Cyperus and Scirpus); inflorescences various<br />

(umbellate in Cyperus and Fimbristylis and less distinctly so in some o<strong>the</strong>r genera); flowers<br />

(<strong>of</strong>ten referred <strong>to</strong> as florets) perfect or unisexual (in Carex and Scleria), each subtended by a<br />

single (rarely 2) scale-like bract (<strong>the</strong>se bracts <strong>of</strong>ten referred <strong>to</strong> as floral scales or in this treatment<br />

as scales <strong>of</strong> spikelets or simply scales), without perianth or perianth reduced <strong>to</strong> bristles or<br />

small perianth scales, solitary or in spikelets (= basic unit <strong>of</strong> inflorescence, consisting <strong>of</strong> a shortened<br />

axis and 1–numerous scale-like bracts, <strong>the</strong> lowermost <strong>of</strong> which are <strong>of</strong>ten empty); stamens<br />

1–3, with an<strong>the</strong>r attached by one end; pistil 1; fruit an achene, usually trigonous or biconvex.<br />

AA large, cosmopolitan (greatest diversity in <strong>the</strong> tropics but <strong>of</strong>ten dominant in cold regions),<br />

taxonomically difficult family <strong>of</strong> herbs, with over 5,000 species in 104 genera (Goetghebeur<br />

1998); ca. 2,000 <strong>of</strong> <strong>the</strong> species are in <strong>the</strong> huge genus Carex. As such, it is <strong>the</strong> third largest monocot<br />

family, following <strong>the</strong> Orchidaceae and Poaceae. The Cyperaceae, with 248 species (7.3% <strong>of</strong><br />

<strong>the</strong> <strong>to</strong>tal number <strong>of</strong> species), is <strong>the</strong> third largest family in <strong>the</strong> East TX flora (after Poaceae and<br />

Asteraceae). Floral structures are quite reduced in association with wind pollination, resulting<br />

in a lack <strong>of</strong> useful taxonomic characters (Yen & Olmstead 2000). Because <strong>of</strong> <strong>the</strong> <strong>of</strong>ten similar<br />

vegetative parts and reduced reproductive structures, technical characters requiring at l<strong>east</strong> a<br />

hand lens frequently have <strong>to</strong> be used <strong>to</strong> distinguish species. Cyperaceae species superficially<br />

resemble both grasses and rushes, and recent evidence suggests a close relationship between<br />

Cyperaceae and Juncaceae (e.g., Simpson 1995; Goetghebeur 1998; Chase et al. 2000; Muasya<br />

2000a; Ball et al. 2002). Phylogenetic analysis indicates that <strong>the</strong> family is monophyletic<br />

(Muasya et al. 2000a). Some molecular data suggest that Juncaceae and Cyperaceae are sister<br />

taxa (Chase et al. 1995b; Linder & Kellogg 1995) or even that Cyperaceae may be derived from<br />

within Juncaceae, possibly making Juncaceae paraphyletic (Plunkett et al. 1995; Munro &<br />

Linder 1998). According <strong>to</strong> Plunkett et al. (1995), <strong>the</strong> “progeni<strong>to</strong>r-derivative relationship <strong>of</strong><br />

Juncaceae and Cyperaceae … reveals an additional example <strong>of</strong> paraphyletic families which presents<br />

a series <strong>of</strong> taxonomic dilemmas.” A number <strong>of</strong> similar situations exist (e.g., Brassicaceae<br />

and Capparaceae), and if paraphyletic families are disallowed (as favored by many cladists),<br />

taxonomists are thus faced with wholesale rearrangement <strong>of</strong> many long established and easily<br />

recognized families—see Appendix 6 for fur<strong>the</strong>r discussion <strong>of</strong> <strong>the</strong>se issues. The family is <strong>of</strong> economic<br />

importance as wildlife food, for woodland grazing, or for erosion control; in n temperate<br />

parts <strong>of</strong> <strong>the</strong> world <strong>the</strong> <strong>plants</strong> sometimes replace grasses as forage; in TX in <strong>the</strong> Hill Country and<br />

w part <strong>of</strong> <strong>the</strong> state, Carex emoryi L. becomes important for lives<strong>to</strong>ck during summer months<br />

(S.D. Jones, pers. comm.); also some are problematic weeds. The sedge family (with ca. 32% <strong>of</strong> its<br />

species having C4 pho<strong>to</strong>syn<strong>the</strong>sis) is one <strong>of</strong> only two monocot families (<strong>the</strong> o<strong>the</strong>r is <strong>the</strong><br />

Poaceae) with <strong>the</strong> typical C4 pho<strong>to</strong>syn<strong>the</strong>tic pathway (Soros & Bruhl 2000); this pathway is an<br />

adaptation <strong>to</strong> prevent water loss and is an advantage in arid environments. Phylogenetic analysis<br />

indicates that <strong>the</strong> C4 pathway has arisen independently four times in <strong>the</strong> Cyperaceae (Soros<br />

& Bruhl 2000). (subclass Commelinidae—Cronquist; order Poales—APG II)<br />

FAMILY RECOGNITION IN THE FIELD: grass-like or rush-like herbs with solid internodes, round or<br />

<strong>of</strong>ten 3-angled culms (“sedges have edges”), and <strong>of</strong>ten 3-ranked leaves; many (but not all) species<br />

grow in wet habitats; flowers small, inconspicuous, without perianth or perianth reduced<br />

<strong>to</strong> bristles or small scales, subtended by 1 scale-like bract each (or in Carex <strong>the</strong> female flower<br />

enclosed in a sac-like perigynium), and arranged in very reduced spikes/spikelets. The ± similar<br />

Poaceae (GRASSES) have hollow or solid internodes, round culms, 2-ranked leaves usually<br />

with a ligule, and flowers subtended by 2 scale-like bracts each (lemma and palea); <strong>the</strong> ± similar<br />

Juncaceae (RUSHES) have flowers with a small 6-parted perianth.<br />

REFERENCES: Svenson 1957; Dahlgren et al. 1985; Goetghebeur 1987, 1998; Tucker 1987; Bruhl et al.<br />

1992; Bruhl 1995; Rolfsmeier 1995; Muasya et al. 2000a, 2000b; Simpson & Inglis 2001; Ball et al. 2002.

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