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<br />STATUS OF ENDANGERED COLORADO SQUAWFISH <br /> <br />967 <br /> <br />of larvae drifted to the lower reach, and (2) re- <br />production or hatching success in the upper reach <br />was formerly much greater than that today, and <br />substantial numbers of larvae were retained in the <br />upper reach even though proportions drifting to <br />the lower reach might have been similar to those <br />in recent years. <br /> <br />Status and Future Prospects <br /> <br />This work and that of others strongly suggest <br />that the population of Colorado squawfish in the <br />Colorado River is small but actively recruiting new <br />individuals to the adult population. Whether pop- <br />ulation size is relatively constant over time is dif- <br />ficult to ascertain, but abundance is apparently <br />much less than that reported earlier in this century. <br />Strong year-classes augment the population <br />enough to cause observable changes in population <br />structure and abundance. The significance of these <br />large year-classes in maintaining the adult popu- <br />lation is also difficult to ascertain, but the apparent <br />increase in numbers of adults observed during this <br />study suggests that the infrequency of recruitment <br />pulses currently limits the size of the adult pop- <br />ulation. <br />Metapopulation theory argues that stochastic <br />fluctuations cause 10cal extinctions, and the prob- <br />ability of extinction decreases with increased pop- <br />ulation size. Thus, to minimize the probability of <br />being driven to extinction, the goal is to achieve <br />a population size large enough to withstand sto- <br />chastic fluctuations and also to maintain sufficient <br />genetic diversity (see Soule 1986; Simberloff <br />1988). Crossing over from the Green River does <br />occur at some unknown rate: the first documented <br />case was an individual caught at Green River rkm <br />52.5 in 1994 and recaptured in the lower Gunnison <br />River (rkm 3.5) in 1996 (B. Burdick, USFWS, per- <br />sonal communication; T. Chart, Utah Division of <br />Wildlife Resources, personal communication). <br />Such movement, even at a low rate (I-tO/gener- <br />ation), should alleviate genetic problems associ- <br />ated with small population size (see Franklin 1980; <br />Gilpin 1993; Mills and Allendorf 1996); therefore, <br />stochastic demographic fluctuations are probably <br />a greater threat to this small population. <br />The abundance of age-O fish and subsequent re- <br />cruitment is probably the greatest source of de- <br />mographic fluctuation in the population of Colo- <br />rado squawfish in the Colorado River. If popula- <br />tion size and viability is limited by this recruit- <br />ment, then strong year-classes are needed more <br />frequently. Presently, variables controlling the rel- I <br />ative success of annual reproduction and first-year <br /> <br />survival are not clearly understood. However, if <br />spawning and hatching success are linked to con- <br />ditions created by spring runoff, changes in runoff <br />patterns in the Colorado River during the past half <br />century resulting from water development may in <br />part explain the decline of this population. <br />A greatly reduced frequency of high spring run- <br />off flows (Osmundson and Kaeding 1991; Van <br />Steeter 1996) influences four factors that might <br />have negatively affected this population. First, <br />high flows during spring may be required to create <br />fresh cobble bars for spawning (Harvey et al. <br />1993) and adequately cleanse fines from existing <br />bars (Haynes et al. 1984; Reiser et al. 1989), con- <br />ditions apparently necessary for spawning site se- <br />lection by both the northern squawfish P. orego- <br />nensis (now northern pikeminnow; Nelson et al. <br />1998) and Colorado squawfish (Beamesderfer and <br />Congleton 1981; Lamarra et al. 1985). The exis- <br />tence of interstitial voids for protection of depos- <br />ited eggs and creation of microcurrents among <br />voids for successful egg incubation may contribute <br />to high egg-hatching success. Second, high flows <br />may serve to dilute waterborne contaminants from <br />agricultural and urban sources that may interfere <br />with reproductive behavior, reduce egg viability, <br />or reduce larval survival (Woodward et al. 1985; <br />Hamilton and Waddell 1994). Third, river bottom- <br />lands require periodic high flows to maintain chan- <br />nel (and thus habitat) diversity and biological pro- <br />ductivity (Junk et al. 1989; Bayley 1991; Ras- <br />mussen 1996) important to young Colorado <br />squawfish. Finally, high, sustained spring flows <br />serve to reduce numbers of nonnative minnows <br />that now dominate backwater nursery habitats <br />(McAda and Kaeding 1989; Osmundson and Kaed- <br />ing 1991; Muth and Nesler 1993; Gido et al. 1997). <br />Although a food source for subadult Colorado <br />squawfish, the prolific red shiner Cyprinella lu- <br />trensis preys on Colorado squawfish larvae (Rup- <br />pert et al. 1993), and larvae of fathead minnow <br />Pimephales promelas compete with Colorado <br />squawfish larvae for food (Beyers et al. 1994). <br />Low adult numbers and infrequent, pulsed re- <br />cruitment make this population vulnerable to ex- <br />tirpation over time. Natural variation in demo- <br />graphics alone makes viability of this population <br />tenuous, even if stability of habitat conditions is <br />assumed. However, for many imperiled species, <br />the threat of deterministic habitat change is prob- <br />ably greater than stochastic demographic variation <br />(Caughley 1994; Harcourt 1995). This may be es- <br />pecially true for this population: more water de- <br />pletions are planned, more nonnative species may <br />