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1116 RUPPIACEAE/RUPPIA<br />

waters (Hellquist 1980), possibly explaining its distribution only on <strong>the</strong> extreme w margin <strong>of</strong><br />

East TX where soils are alkaline. Guo and Cook (1989) found that at l<strong>east</strong> some pollination can<br />

occur underwater in this species.<br />

RUPPIACEAE Horan. ex Hutch.<br />

DITCH-GRASS FAMILY<br />

AA very small (ca. 10 species in a single genus—Haynes 2000c), nearly cosmopolitan family <strong>of</strong><br />

submersed, glabrous, aquatic herbs <strong>of</strong> brackish or saline conditions or <strong>of</strong> fresh waters with high<br />

calcium or sulfur ion concentrations. The reduced flowers (perianth absent) and morphological<br />

variation have made it difficult <strong>to</strong> delimit species within <strong>the</strong> group (e.g., some workers have recognized<br />

only a single variable species). Ruppiaceae are unusual in lacking vessel elements<br />

(Mabberley 1997). The family is similar <strong>to</strong> and has sometimes been included in <strong>the</strong><br />

Potamoge<strong>to</strong>naceae (e.g., Haynes 1978; Jacobs & Brock 1982; Dahlgren et al. 1985; Thorne 1993b),<br />

and it has been suggested that <strong>the</strong> Ruppiaceae evolved from <strong>the</strong> Potamoge<strong>to</strong>naceae by reduction<br />

(Heywood 1993). Molecular studies have suggested a closer relationship <strong>to</strong> <strong>the</strong> marine families<br />

Cymodoceaceae and Posidoniaceae (Les et al. 1993; Les & Haynes 1995; Haynes et al. 1998b),<br />

and we are following <strong>the</strong> recent treatment by Haynes (2000c) in segregating Ruppia in<strong>to</strong> its<br />

own family. (subclass Alismatidae—Cronquist; order Alismatales—APG II)<br />

FAMILY RECOGNITION IN THE FIELD: submersed aquatic herbs <strong>of</strong> brackish, saline, or mineral waters,<br />

with thread-like leaves 0.5 mm or less wide, flowers without perianth, and a terminal spike<br />

that in fruit has an elongate, usually coiled peduncle.<br />

REFERENCES: Haynes 1978; 2000c; Rico-Gray 1991; Haynes et al. 1998b.<br />

RUPPIA L. DITCH-GRASS, WIDGEON-GRASS, RUPPIA<br />

AOnly two species occur in North America, with both known from TX (one known only <strong>to</strong><br />

<strong>the</strong> s <strong>of</strong> East TX). The two are very similar and have been treated as variants <strong>of</strong> a single species<br />

by some authorities (e.g., Fernald & Wiegand 1914). However, we are following Thorne (1993b)<br />

and Haynes (2000c) in recognizing <strong>the</strong>m at <strong>the</strong> specific level. Pollination takes place on <strong>the</strong> surface<br />

<strong>of</strong> <strong>the</strong> water. After release, <strong>the</strong> boomerang-shaped pollen grains aggregate by adhering end<br />

<strong>to</strong> end in chains or in snowflake-like groups and are rafted <strong>to</strong> <strong>the</strong> stigma, which is held at <strong>the</strong><br />

water surface due <strong>to</strong> <strong>the</strong> elongated peduncle and buoyancy provided by s<strong>to</strong>mata and intercellular<br />

spaces in <strong>the</strong> spongy tissue <strong>of</strong> <strong>the</strong> inflorescence (Haynes 1978; Dahlgren et al. 1985; Cox &<br />

Knox 1989; Lacroix & Kemp 1997). This type <strong>of</strong> hydrophily (= water-mediated pollination) occurring<br />

at <strong>the</strong> water surface is known as epihydrophily (in contrast <strong>to</strong> hypohydrophily or underwater<br />

pollination as seen in Najas—Hydrocharitaceae) (Philbrick 1993). Some workers (e.g.,<br />

Cox 1988) fur<strong>the</strong>r divide epihydrophily in<strong>to</strong> a category in which pollen is transported just<br />

above <strong>the</strong> surface <strong>of</strong> <strong>the</strong> water (e.g., Hydrilla, Vallisneria) and a category in which pollen is transported<br />

directly on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> water (e.g., Elodea, Ruppia). Cox and Knox (1989) have indicated<br />

that a variety <strong>of</strong> <strong>plants</strong> with pollen transported directly on <strong>the</strong> water surface (a two dimensional<br />

system) show striking convergent evolution that can be defined as a “surface-pollination<br />

syndrome”—floating pollen grains that join <strong>to</strong>ge<strong>the</strong>r in long floating rafts or “search vehicles”<br />

whose large size greatly increases <strong>the</strong> probability <strong>of</strong> encountering a stigma, as well as surface<br />

borne stigmas that are ei<strong>the</strong>r elongate or that create a slight depression in <strong>the</strong> water surface (thus<br />

maximizing <strong>the</strong> probability <strong>of</strong> being encountered). Ruppia species are considered <strong>to</strong> be among<br />

<strong>the</strong> most valuable submersed aquatics as sources <strong>of</strong> food for waterfowl and o<strong>the</strong>r wildlife. They<br />

provide food and cover for fish, all parts <strong>of</strong> <strong>the</strong> <strong>plants</strong> are eaten by many species <strong>of</strong> waterfowl, and<br />

marsh and shorebirds eat <strong>the</strong> fruits and foliage (Correll & Correll 1972; Haynes 1978; Godfrey &<br />

Wooten 1979). (Named for Heinrich Bernhard Ruppius, 1688–1719, German botanist)<br />

REFERENCES: Fernald & Wiegand 1914; Gamerro 1968; Richardson 1980; Jacobs & Brock 1982;<br />

Lacroix & Kemp 1997; Morgan & Holmes 2004.

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