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<br />1738 <br /> <br />Based on simulation results, this scenario suggested <br />that when large red shiners are present in early to <br />midsummer, wild Colorado pikeminnow must grow <br />quickly to survive to autumn. This hypothesis was <br />supported because autumn juveniles averaged 12-73% <br />faster growth than summer juveniles. A potential <br />explanation for this pattern is size-dependent predation <br />mortality from a gape-limited predator. Under this <br />hypothesis, slow-growing larvae should be susceptible <br />to predators for a longer period of time than fast- <br />growing fish, so only the fastest-growing fish in a <br />cohort survive (Houde 1987; Miller et al. 1988). <br />Colorado pikeminnow hatched later in summer sur- <br />vived to autumn, even though they were slower <br />growing, probably because large red shiners that <br />caused differential mortality were not present. <br />Even though relatively small and slow-growing <br />Colorado pikeminnow can survive to autumn, small- <br />bodied juveniles may have relatively low overwinter <br />survival. In a laboratory study, Thompson et al. (1991) <br />found that both fed and starved Colorado pikeminnow <br />that were relatively small (mean, 30 or 36 mm TL) had <br />lower survivallhan larger fish (44 mm) over simulated <br />winter conditions. In a Green River field study, Haines <br />et al. (1998) found that small Colorado pikeminnow <br />(mode, 28 mm TL) had only 6% overwinter survival <br />durillg a high-flow winter period, whereas in a different <br />year, when Colorado pikeminnow were larger (>38 <br />mm), they had higher overwinter survival (56-65%), <br />albeit winter flows were lower. Poor overwinter <br />survival of small Colorado pikeminnow in the Green <br />River places added importance on the few larger fish <br />from early cohorts that survive, because those fish may <br />have higher overwinter survival. <br />Intra-armual variation in survival of cohorts of larvae <br />with different hatching dates has been observed for <br />other freshwater fishes (Crecco and Savoy 1985; Rice <br />et al. 1987; Limburg 1996; Cargnelli and Gross 1996; <br />Mion et al. 1998; Garvey et al. 2002). Most of those <br />studies invoked low temperatures or high flows as <br />mechanisms for reduced survival of portions of the <br />year-class. Poor survival of early spawned bloater <br />Coregonus hoyi was thought a function of high <br />predation rates on slow-growing larvae when water <br />was relatively cold (Rice et al. 1987; Luecke et al. <br />1990), which similar to our study, suggested Ihat <br />bloater recruitment may be structured by interacting <br />physical and biotic factors. <br /> <br />IBM Simulations <br /> <br />The ffiM simulations confirmed patterns observed in <br />field studies and provided support that predation <br />interacting with environmental variables was a main <br />mechanism structuring recruitment patterns. Simula- <br /> <br />BESTGEN ET AL. <br /> <br />tions were also useful to explore "what if' scenarios <br />such as the potential effect of reducing predator <br />abundance or exploring effects of different thermal <br />regimes on Colorado pikeminnow growth and recruit- <br />ment. <br />Additional data are needed to make ffiM simulations <br />more representative of the processes in the Green <br />River. For example, understanding seasonal size and <br />density shifts of red shiners and other potentially <br />predaceous fishes would better reflect Ihe predator <br />assemblage in backwaters. Information on the poten- <br />tially important effects of discharge level and flow <br />fluctuations on food availability and fish growth would <br />also be useful, particularly if ffiM simulations were <br />linked to bioenergetics modeling (Burke and Rice <br />2002). This is important because discharge fluctua- <br />tions, such as those produced by releases from <br />hydroelectric dams or natural flow increases, reduce <br />the abundance of nearshore chironomid assemblages <br />(Blinn et al. 1995), which are important in the diets of <br />most young fishes in the Green River (Muth and <br />Snyder 1995). <br /> <br />Alternative Recruitment Hypotheses <br /> <br />Starvation is another possible mechanism affecting <br />recruitment of early life stages of Colorado pike- <br />minnow in Green River backwaters, but we found no <br />support for that hypothesis. Starvation is primarily a <br />function of body size for fish larvae (Miller et al. <br />1988), and Colorado pikeminnow larvae are only of <br />moderate size at hatching (5.5 mm TL). However, 87% <br />of laboratory-reared larvae starved for 15 d after they <br />could eat (21 d posthatch) survived when offered food <br />(Bestgen 1996). Further, survival predictions from <br />Miller et al. (1988) for larvae that hatch at the same size <br />as Colorado pikeminnow suggested that pikeminnow <br />have higher than average resistance to starvation. Thus, <br />high starvation resistance makes it unlikely that direct <br />starvation mortality is an important mode of population <br />regulation. <br />We did find evidence that intra-annual flow <br />fluctuations affected the recruitment of cohorts of <br />Colorado pikeminnow larvae, perhaps through interac- <br />tions with predation. This was apparent in the lower <br />Green River in 1991 and 1992, when increased <br />discharge and very high turbidity from thunderstorm <br />runoff were associated with reduced growth and <br />recruitment of juvenile Colorado pikeminnow. At <br />discharge levels above 80 m3fs, which was in the <br />range observed in 1992, much of the available <br />backwater habitat was eliminated in that mostly <br />canyon-constrained reach (Haines and Tyus 1990; <br />K.R.B., personal observation) and habitat loss may <br />have reduced foraging efficiency and Colorado pike- <br />