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food sources) in which streams provide the environmental (abiotic and biotic) conditions <br />that each life history stage needs for survival, growth, and reproduction. This requires a <br />knowledge of habitat use, incorporating information such as the path and timing of <br />migrations, location and time of spawning, location of nursery areas, time of occupation, <br />habitats occupied by juveniles and adults at different-times of the year, and other <br />pertinent information. The amount of such information varies greatly among the four <br />endangered fishes, -from excellent information on Colorado pikeminnow to mostly ' <br />anecdotal accounts for the bonytail (e.g., see reviews in recovery plans: USFWS <br />1990ab, 1991, 1998; surveys: Quartarone 1993; and other information: Osmundson et <br />al. 1997,1998; Osmundson and Burnham 1998; and Muth et al. 2000). <br />In their natural riverine habitat, the timing of most life history events for the four <br />endangered fishes is due to interactions among the environmental variables that <br />provide cues, and of these, flow is an important component (Nester et al. 1988;Tyus <br />and Karp 1989, 1991; Tyus 1990; Figure 2). Seasonal changes in temperature and <br />photoperiod also are likely involved in the timing of life history events, but are difficult to <br />separate from flow events such as spring runoff. Flow also plays a significant role in <br />the availability of certain types of habitat (e.g., habitat in the floodplain will only be <br />inundated during peak flows), and in the physical dimensions of habitat (higher flow <br />usually means deeper, wider habitat). However, relatively simplistic attempts to provide <br />habitat utilization profiles as a way of describing usable area of habitat for the <br />endangered fishes as a function of flow has been frustrated due to fish use of different <br />habitat parameters (e.g., water depth or velocity) depending on: location (e.g:,'river), <br />time of year or life stage, and by use of habitats such as eddies and backwaters that <br />are not amenable to flow models (Valdez et al. 1986, 1990; Tyus 1992). <br />Superimposed on the spatial and temporal map of physical habitat are biological habitat <br />components, which are defined largely by predator-prey or competitive interactions. <br />The endangered fishes must have access to an abundance of suitable food species, <br />but not be exposed excessively to predation. In some locations, high predation by <br />introduced nonnative fishes may be the greatest impediment to expanding or <br />reestablishing populations of the endangered fishes (e.g., Marsh and Brooks 1989, <br />Marsh and Douglas 1997). <br />A less obvious biological aspect that influences habitat selection and use is learned <br />andlor instinctive (genetic) behavior. Major behavioral attributes tend to have a <br />phylogenetic basis and are commonly shared among related taxa. Examples include a <br />propensity for selecting certain habitats, prey selection, extent and direction of <br />migrations, and orientations to flow, temperature or substrate. Learned responses such <br />as imprinting, are essential to some migratory species (e.g., acipenserids, clupeids, <br />salmonids, and catostomids), which may rely on subtle environmental cues, such as <br />chemical composition of the water, to guide them back to the spawning areas from <br />which they emerged several years earlier (e.g., see reviews by Hasler and Scholz 1983; <br />McKeown 1984, Smith 1985). Concern about the role of these cues is raised whenever <br />the natural habitat, or access to it, is altered. <br />11 <br />