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<br />tions. Chironomids, baetid mayflies, amphipods, planar- <br />ians, oligochaetes, and snails tend to be predominant in <br />tailwaters below deep-release dams. Common benthic <br />invertebrates in potamon reaches in the upper basin and <br />lower river are oligochaetes, chironomids, gastropods, <br />leeches, turbellarians, sphaeriid clams, odonates, beetles, <br />simuliids, net-spinning caddisflies, and baetid mayflies. <br />Introduced freshwater shrimp and crayfishes are locally <br />abundant in the lower river, and the introduced Asiatic <br />clam, Corbicula fluminea, occurs as far upstream as Lake <br />Mead. The Colorado River System appears unique in that <br />unionacean clams are virtually absent from its waters and <br />isopods are usually absent from Iotic sites. <br />The fish fauna of the Colorado River Basin bears little <br />resemblance to its original state. Approximately <br />100 species are now present; some 67 non-native fish spe- <br />cies in 16 families have been introduced since the turn of <br />the century and are now predominant in most fish commu- <br />nities (Miller and Lowe 1964; Minckley 1973, 1979; Moyle <br />1976; Carothers and Minckley 1981; Tyus et al. 1982). In <br />terms of numbers of species, cyprinids, centrarchids, <br />salmonids, catostomids, and ictalurids head the list of <br />introduced fishes. Of the 54 natives listed in Table 2, 17 are <br />either threatened, endangered, or extinct, and most have <br />experienced drastic abundance and range reductions (Miller <br />1972; Minckley 1973, 1979; Joseph et al. 1977; Behnke <br />and Benson 1983). Two species (Pahranagat spinedace and <br />Las Vegas dace) are extinct, and the woundfin is almost <br />gone. The cutthroat trout is threatened in the upper basin <br />(Behnke 1979), and most stream- and spring-inhabiting <br />fishes of the middle and lowermost Colorado River drainage <br />are legally protected or of special concern (Johnson 1987). <br />All of the "big-river" fishes are in jeopardy (Minckley <br />1973, 1983; Carothers and Minckley 1981; Tyus et al. <br />1982; Behnke and Benson 1983; Hickman 1983). Wild <br />Colorado squawfish are gone from the lower basin, and the <br />flannelmouth sucker is extirpated from the Gila River drain- <br />age (Williams et al. 1985). Tyus (1987) considered razor- <br />back sucker one of the rarest fishes in the Colorado River <br />Basin. The humpback chub persists tenuously in the Little <br />Colorado River and Grand Canyon (Kaeding and Zimmer- <br />man 1983) and occupies a few scattered canyon areas in the <br />upper basin (Behnke and Benson 1983). The bony tail, origi- <br />nally widespread and abundant in the basin, is functionally <br />extinct; a few scattered individuals exist in the Green and <br />Upper Mainstem Colorado rivers and in Lake Mohave in the <br />lower basin (Behnke and Benson 1983). Behnke and Benson <br />(1983), said of the bony tail's demise that "If it were not for <br />the stark example provided by the passenger pigeon, such <br />rapid disappearance of a species once so abundant would be <br />almost beyond belief". Several authors (Minckley 1979; <br />Behnke 1980; Hubbard 1980; Molles 1980; Behnke and <br />Benson 1983; Williams et al. 1985; Stanford and Ward <br />1986c) have attributed decline of native fishes to (1) modifi- <br />cation and loss of habitat and (2) introduction of non-native <br />specIes. <br />Construction and regulation of dams have had severe <br />impacts on the fish fauna of the Colorado River, and little <br />unaltered habitat remains (Tyus 1984). Coats (1984) <br />described general lack of regard for minimum flow needs <br />of fishes in operating Colorado River dams. Extreme fluctu- <br />ations and alteration of seasonal flow regimes have been <br />implicated in alleged loss of 1983 and 1984 year classes of <br /> <br />232 <br /> <br />the Colorado squawfish in its most productive remaining <br />nursery habitat (Jones and Tyus 1985). <br />Williams et al. (1985) discussed adverse impacts of <br />introduction of non-native species on native fishes in most <br />of the 15 aquatic ecosystems in North American deserts that <br />they considered. Schoenherr (1981) described behavioral <br />interactions between introduced redbelly tilapia, Tilapia zil- <br />iii, and sailfin molly, Poecilia latipinna, leading to replace- <br />ment of desert pupfish. Schoenherr (1981), MinckIey et al. <br />(1977); Meffe et al. (1983), and Meffe (1984, 1985) studied <br />predation by mosquitofish, Gambusia affinis, resulting in <br />endangerment of Sonoran topminnow. Behnke and Benson <br />(1983) discussed possible redside shiner, Richardsonius <br />balteatus, competition with Colorado squawfish in the <br />upper basin. Colorado squawfish interactions with other <br />non-natives, e.g., "choking" on channel catfish, Ictalurus <br />punctatus, (McAda 1983; Pimentel et aI. 1985) and compe- <br />tition with northern pike, Esox lucius, (Wick et al. 1985) <br />need further research. <br />Surveys by Moffett (1942, 1943), Dill (1944), and Wallis <br />(1951), stimulated stocking of game fishes, and threadfin <br />shad (Dorosoma petenense), and various invertebrates were <br />stocked as forage. Trouts (Salmo gairdneri, S. clarki, and <br />Salvelinus fontinalis) and Pacific salmon (Oncorhynchus <br />kisutch and O. nerka) were stocked in reservoirs, and col- <br />dwater fisheries developed. Striped bass, Morone saxatilis, <br />were introduced in Lake Mead in 1969, and a successful <br />fishery developed in the 1970's. Rainbow trout and thread- <br />fin shad populations declined as a result of predation by <br />striped bass (Baker and Paulson 1983). <br />A number of attempts were made to remove "coarse" <br />fishes to make room for introduced species. In 1962, <br />700-800 km of the Green River and its tributaries were <br />treated with rotenone to allow Flaming Gorge Reservoir and <br />the streams to realize their full potentials as trout fisheries <br />(Miller 1963; Dexter 1965; Pearson et al. 1968). Down- <br />stream detoxification failed, and rare endemic fishes were <br />killed in Dinosaur National Monument. Binns (1967) <br />reported that Colorado squawfish, razorback sucker, and <br />rare mayflies had not reestablished populations in the treated <br />area after 2 yr. Rotenone was also applied in the San Juan <br />River prior to closure of Navaho Reservoir and on the Gila <br />River upstream from San Carlos Reservoir. Impacts of these <br />incidents have never been fully assessed. <br />Non-native trout fisheries downstream from Colorado <br />River dams (Mullan et al. 1976) have become valuable <br />assets. Flaming Gorge Dam has been modified to improve <br />such a fishery through increase in tail water temperatures <br />(Holden 1979). <br />Other fisheries in the Colorado River Basin are dependent <br />on non-native centrarchids and striped bass, but fishing for <br />channel and flathead catfish (Pylodictis olivaris), walleye <br />(Stizostedion vitreum), and northern pike is popular locally <br />(Behnke et aI. 1982; Stanford and Ward 1986c). Threadfin <br />shad are important as food for piscivores in Lake Powell and <br />lower-basin reservoirs (Johnson 1970, 1971; Stanford and <br />Ward 1986c). Martin et al. (1982) estimated the annual eco- <br />nomic value of Lake Mead fisheries at $69 million. Mullan <br />et al. (1976), Carothers and Dolan (1982), Persons and <br />Bulkley (1982), Morgenson (1983), and Baker and Paulson <br />(1983) considered fishery management and cited similar <br />studies on the Colorado system. <br />Studies of fish production in the system are rare. Scarnec- <br />