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<br />174 <br /> <br />D. J. ORTH <br /> <br />Orth and Maughan, 1982, 1986). Temporal variations in stream flow can limit populations of stream fish <br />by affecting emigration, growth, survival, or recruitment. Consequently, population size at a given time is <br />determined by past habitat limitations for any life stage and instantaneous measures of microhabitat <br />availability will not be related to instantaneous population size. It is, therefore, not surprising that many <br />attempts to correlate instantaneous WUA and instantaneous fish standing crop have been unsuccessful <br />(Scott and Shirvell, in press). <br />For stream fishes, survival during the early life stages is critical in determining adult population size, <br />and for many species, these stages can tolerate only near-zero water velocities (Larimore, 1975; Ottaway <br />and Clark, 1981; Ottaway and Forrest, 1983). Therefore, fish densities may be strongly related to habitat <br />conditions during the critical early life stages. Anderson and Nehring (1985) found strong negative <br />relationships between juvenile trout (Salmo gairdneri and S. trulla) numbers and peak flows during the <br />time when fry have emerged from the gravel, presumably due to excessive water velocities. Loar el al. <br />(1985) found that rainbow trout densities were correlated with minimum incubation habitat (minimum <br />WUA during incubation period) in southern Appalachian streams without brown trout. However, the <br />lack of a significant relationship for sites where rainbow trout occurred with brown trout suggests a <br />potential competitive interaction. In populations where recruitment is highly responsive to flow <br />conditions, which may be unpredictable, population densities may exhibit stochastic annual variability. <br />Consequently, adult population densities cannot be predicted on the basis of present WUA measures <br />alone; knowledge of past limiting habitat events and past population densities would be needed to predict <br />population density. <br />It is crucial that those involved in negotiating stream flow regimes in regulated rivers recognize that <br />microhabitat availability is not the only regulatory factor and does not operate continuously to limit <br />stream fish populations. The prevailing limiting factors vary widely among streams and may vary <br />temporally within the same stream. Moyle and.Vondracek' {1985-l_fourid that despite variable flow <br />conditions in a small stream, stream fish ass;mblage 'structure -Wa~Ppersistent, suggesting that biotic <br />interactions play an important role in fish population regulation. However,in other streams, stochastic <br />factors (e.g. variable flow) may be equally important~ resulting in more variable fish assemblage structure <br />(Grossman el al., 1985; Schlosser, 1985).-. . '._ ._ <br /> <br />ENERGETICS AND-FOOD LIMlTATION <br /> <br />The usual omission of aquatic invertebrates from instream flow assessments presumes that either food <br />availability does not normally limit fish production in_~treams or adequate flows for fish result in suitable <br />flows for invertebrates. At the present time, there is insufficient evidence to support either of these <br />assumptions. Furthermore, evidence indicates that some aquatic insects are quite sensitive to small <br />changes in flow and shoulci be included in instream flow asseSSS1ents (Gore, in press). Stream hydraulics <br />has been proposed as a major determinant affecting stream invertebrate assemblages (Statzner and <br />Higler, 1986). In recent years several studies have described the microhabi!at requirements for several <br />stream insects for application in instream flow assessments (Gor~ and Judy, -1981; Orth and Maughan, <br />1983; Teague el al., 1985). Morin el al. (i986) has critically ~valuated methods for developing habitat <br />suitability criteria for stream insects and Newbury (1984) described the measurement of important <br />hydraulic characteristics. <br />One line of evidence that supports a regulatory influence of prey availability on biomass of stream fish <br />populations is the indication that fish predators depress the density of their invertebrate prey, <br />subsequently resulting in reduced growth of individual fish (Brocksen et al., 1968; F1e~ker, 1984; <br />Anderson, 1985; Angermeier, 1985). In laboratory streams, Brocksen et al. (1968) found that reticulate <br />sculpins (COllus perplexus) and cutthroat trout (Salmo clarki) could overexploit their invertebrate prey. <br />Increased fish biomass reduced the density of food organisms, which directly reduced growth rate of trout <br />and sculpins. Anderson (1985) also observed that growth rates in wild populations of sculpins were <br />inversely related to population density. In exclosure experiments, Angermeier (1985) demonstrated that <br />- predation by stream fish depressed invertebrate densities in silt-sand habitats but not in gravel-cobble <br />