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558 CYPERACEAE/CYPERUS<br />

sexual (seed producing) and asexual (reproducing by rhizome propagation or formation <strong>of</strong> vegetative<br />

plantlets in <strong>the</strong> spikelets) reproduction is known in <strong>the</strong> species (Miao et al. 1998). This<br />

species has <strong>of</strong>ten been treated as a subspecies <strong>of</strong> <strong>the</strong> Old World C. mariscus (e.g., Kartesz 1999).<br />

However, we are following Hatch et al. (1990), Jones et al. (1997), and Tucker (2002c) in treating<br />

it as a distinct species. Tucker (2002c) recognized it as such “with some trepidation.”<br />

Cladium mariscoides (Muhl.) Torr., (resembling Mariscus, a segregate now included in Cyperus),<br />

SMOOTH SAW-GRASS, TWIG-RUSH, SWAMP SAW-GRASS. Plant 0.3–1 m tall; rhizomes ca. 2 mm in<br />

diam.; leaf blades involute, with scaberulous margins; inflorescences 5–30 cm long, slender, ca.<br />

2–5 cm wide, usually <strong>of</strong> relatively few cymes, <strong>the</strong> inflorescences much smaller than in C.<br />

jamaicense, ca. 15–25 mm long, with spikelets in groups <strong>of</strong> 3–10 at <strong>the</strong> ends <strong>of</strong> short, erect<br />

branches; spikelets 3–6 mm long; achenes smooth, short-cylindric, apiculate-pointed, truncate<br />

basally, 2.5–3.5 mm long. Seepage-fed peat bogs, wet areas; Anderson (McRoberts & McRoberts<br />

3990—BRIT), Smith (Bridges & Orzell 1989a), and Henderson (Turner et al. 2003) cos. in <strong>the</strong> nc<br />

part <strong>of</strong> East TX; very rare in <strong>the</strong> state; mainly se Canada and ne U.S., disjunct s <strong>to</strong> FL and TX.<br />

Jul–Sep. [Mariscus mariscoides (Muhl.) Kuntze] First reported for TX in 1989 (Bridges & Orzell<br />

1989a) from bogs associated with Queen City Sand (Eocene) and an adjacent terrace. While not<br />

<strong>of</strong>ficially designated as such (e.g., TOES 1993; Carr 2002d; Poole et al. 2002), given its limited<br />

distribution in <strong>the</strong> state, we consider this species <strong>to</strong> be <strong>of</strong> conservation concern in TX. It is possibly<br />

a glacial age relict—surviving in a specialized microhabitat, long after <strong>the</strong> general climate<br />

<strong>of</strong> <strong>the</strong> area is no longer appropriate (Kral 1966; MacRoberts & MacRoberts 1997; Diggs 2002;<br />

Diggs & O’Kennon 2003). �<br />

CYPERUS L. FLAT SEDGE, UMBRELLA SEDGE, SEDGE-GRASS<br />

Annuals or usually perennials; <strong>plants</strong> largely glabrous except for scabrous-margined leaves;<br />

culms (= stems) triangular in cross section; leaves basal or nearly so; inflorescences terminal,<br />

head-like or umbel-like, <strong>of</strong>ten conspicuously branched, leafy-bracted at base; scales <strong>of</strong> spikelets<br />

usually 4–many, usually visibly 2-ranked (= in two distinct rows), <strong>the</strong> spikelets � flattened or<br />

square in cross section, usually with 3–many bisexual flowers; perianth bristles absent; style<br />

branches 2 or 3; achenes � 1 per spikelet, trigonous or lenticular, without a tubercle.<br />

AA genus <strong>of</strong> ca. 600 species <strong>of</strong> annual or perennial herbs <strong>of</strong> pantemperate and tropical distribution<br />

(Tucker et al. 2002); as such it is <strong>the</strong> second largest genus in <strong>the</strong> Cyperaceae following<br />

Carex (Tucker 1987). The spikelet scales in two vertical rows or ranks (not spirally arranged)<br />

makes Cyperus species easily recognized. Recent phylogenetic analyses suggest that <strong>the</strong> genus<br />

is paraphyletic with a number <strong>of</strong> genera embedded within it, including Kyllinga and Lipocarpha<br />

(Muasya et al. 2000a, 2000b, 2002). Cyperus is a taxonomically difficult genus with a number<br />

<strong>of</strong> taxa apparently hybridizing and intergrading morphologically. Intermediates between C.<br />

croceus, C. echinatus, C. retr<strong>of</strong>lexus, and C. retrorsus are frequently seen. Similar problems occur<br />

within o<strong>the</strong>r species complexes. Both C3 and C4 pho<strong>to</strong>syn<strong>the</strong>sis are known in <strong>the</strong> genus<br />

(Tucker 1987; Li et al. 1999). Some species are problematic weeds while o<strong>the</strong>rs are cultivated as<br />

ornamentals. The pith from <strong>the</strong> culms <strong>of</strong> <strong>the</strong> commonly cultivated Old World Cyperus papyrus<br />

L. (PAPYRUS, PAPER-REED) was used by <strong>the</strong> Egyptians <strong>to</strong> make paper at l<strong>east</strong> 5,500 years ago<br />

and later by <strong>the</strong> Greeks and Romans (Tucker et al. 2002); <strong>the</strong> Greek word for <strong>the</strong> plant was<br />

papyros from which our word paper is derived (Hepper 1992). This species was also used <strong>to</strong><br />

make sandals, ropes, and boats (e.g., Moses in <strong>the</strong> bulrushes). The Greek word byblos was <strong>the</strong><br />

name for <strong>the</strong> white pith <strong>of</strong> PAPYRUS used in making paper (<strong>the</strong> pith was cut in<strong>to</strong> strips, glued<br />

<strong>to</strong>ge<strong>the</strong>r, and <strong>the</strong>n pressed and dried—Zohary 1982); <strong>the</strong> word byblos became modified in<strong>to</strong><br />

biblion and was applied <strong>to</strong> all scrolls or books and eventually <strong>to</strong> <strong>the</strong> Bible (Hepper 1992). (Greek,<br />

kupeiros or cypeiros, ancient name for <strong>the</strong> Eurasian species, Cyperus longus L.—Tucker et al. 2002)<br />

REFERENCES: Kükenthal 1935–1936; McGivney 1938, 1941a, 1941b; Corcoran 1941; Marcks 1972,

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