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HYDRILLIA/HYDROCHARITACEAE 673<br />

margined and lacking a fringe <strong>of</strong> orange-brown hairs; internodes mostly much shorter than<br />

<strong>the</strong> leaves and up <strong>to</strong> 10 mm long 1 dm below <strong>the</strong> growing tip; flowers with round petals and<br />

with nectaries _____________________________________________________________________ E. najas<br />

2. Leaves linear <strong>to</strong> attenuate, straight <strong>to</strong> slightly concave between <strong>the</strong> serrations, <strong>the</strong>ir midvein<br />

sometimes with prickles on <strong>the</strong> abaxial leaf surface; scales in <strong>the</strong> leaf axis mostly ca. 0.5 mm<br />

long, mostly with a fringe <strong>of</strong> orange-brown hairs; internodes mostly as long as <strong>the</strong> leaves and<br />

up <strong>to</strong> 50 mm long 1 dm below <strong>the</strong> growing tip; flowers with linear petals, lacking nectaries<br />

________________________________________________________________________ Hydrilla verticillata<br />

HYDRILLA Rich. WATER-THYME<br />

AA monotypic Old World genus (Haynes & Holm-Nielsen 2001) introduced in<strong>to</strong> <strong>the</strong> U.S. and<br />

Central America. The male flowers are released underwater, rise unopened <strong>to</strong> <strong>the</strong> surface, and<br />

float <strong>to</strong> <strong>the</strong> female flowers (an example <strong>of</strong> epihydrophily—see discussion in Vallisneria generic<br />

synopsis). Cook and Lüönd (1982) indicated that during flower opening <strong>the</strong> stamens spring<br />

upward, <strong>the</strong> an<strong>the</strong>rs burst, and pollen is scattered in <strong>the</strong> air. Some <strong>of</strong> this lands on <strong>the</strong> female<br />

flowers and some on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> water; apparently <strong>the</strong> pollen grains landing on <strong>the</strong> water<br />

are lost in terms <strong>of</strong> reproduction and only those actually landing on <strong>the</strong> female flower are involved<br />

in pollination. Even though <strong>the</strong> effective pollen grains never contact <strong>the</strong> water, <strong>the</strong> male<br />

flowers float and <strong>the</strong> system can thus be described as epihydrophily. (Greek: hydr-, water, in<br />

reference <strong>to</strong> <strong>the</strong> habitat, plus <strong>the</strong> diminutive suffix—Haynes & Holm-Nielsen 2001)<br />

REFERENCES: Lazor 1975; Godfrey & Wooten 1979; Cook & Lüönd 1982; Pieterse 1983; Steward et<br />

al. 1984; Langeland 1996.<br />

Hydrilla verticillata (L.f.) Royle, (whorled), WATER-THYME. Perennial, submersed, dioecious or<br />

monoecious aquatic; horizontal stems in <strong>the</strong> substrate sometimes forming turions (= swollen<br />

structure or <strong>of</strong>fshoot, <strong>of</strong>ten serving <strong>to</strong> overwinter); erect stems branching, capable <strong>of</strong> ascending<br />

as much as 8.5 m (Godfrey & Wooten 1979) and growing horizontally near <strong>the</strong> water surface;<br />

leaves in whorls <strong>of</strong> (2–)3–8, sessile, mostly 1.5(–2) cm or less long, 1.5–2 mm wide, narrowly oblong,<br />

visibly serrate marginally, with a single vein which on <strong>the</strong> lower surface bears conical<br />

protrusions tipped with teeth (fresh leaves thus noticeably rough <strong>to</strong> <strong>the</strong> <strong>to</strong>uch); staminate flowers<br />

detaching and floating <strong>to</strong> <strong>the</strong> surface; pistillate flowers reaching <strong>the</strong> surface at <strong>the</strong> end <strong>of</strong> an<br />

elongate, thread-like floral tube 4–5 cm long; perianth segments 6, colorless, very inconspicuous,<br />

3–5 mm long; fruits 5–6 mm long, � fusiform. Lakes and o<strong>the</strong>r aquatic habitats; rapidly<br />

spreading at present in East TX (M. Smart, pers. comm); probably spread vegetatively from lake<br />

<strong>to</strong> lake by boats or boat trailers and also intentionally by fishermen (L. Hartman, pers. comm.)<br />

<strong>to</strong> “improve” <strong>the</strong> habitat (this is both illegal and ill-advised, since it ultimately degrades <strong>the</strong><br />

fishery); scattered in e 1/3 <strong>of</strong> TX. Summer–fall. Native <strong>of</strong> <strong>the</strong> Old World, probably <strong>the</strong> warmer<br />

areas <strong>of</strong> Asia (Cook & Lüönd 1982). Austin (1978) indicated that <strong>the</strong> species apparently first<br />

entered <strong>the</strong> U.S. at Miami, FL in 1959, while Blackburn et al. (in Steward et al. 1984) noted that<br />

female <strong>plants</strong> were first reported in <strong>the</strong> U.S. in 1960 from South Florida (misidentified as Elodea<br />

canadensis). Monoecious <strong>plants</strong> were first observed in <strong>the</strong> U.S. in Washing<strong>to</strong>n, D.C. in 1982. Female<br />

(dioecious) individuals are triploid, while monoecious <strong>plants</strong> are diploid (Steward et al.<br />

1984). Since its introduction in FL, this problematic species has spread across <strong>the</strong> sou<strong>the</strong>rn U.S.<br />

<strong>to</strong> TX, AZ, CA, and WA, and up <strong>the</strong> <strong>east</strong> coast <strong>to</strong> DE, MD, and CT. The earliest TX collection we<br />

are aware <strong>of</strong> is from 1974 (Amerson 2097, BRIT), while <strong>the</strong> species was first collected in LA in<br />

1973 (Solymosy 1974). Hydrilla verticillata is a serious invasive pest which can completely<br />

dominate aquatic habitats, eliminating native species, clogging waterways, and severely curtailing<br />

recreational use (Steward et al. 1984; Flack & Furlow 1996; Langeland 1996; Kartesz<br />

1999). It is considered by many wetland biologists as “potentially <strong>the</strong> most dreaded species <strong>of</strong><br />

aquatic weed in warm temperate areas” (Crow & Hellquist 2000a). In TX, H. verticillata is considered<br />

a “harmful or potentially harmful exotic plant,” and it is illegal <strong>to</strong> release, import, sell,

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