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<br />homing behavior, and the fish possess the biological "equipment" necessary for <br />responding to those cues. The chemical cues could govern the direction of fish <br />movements by eliciting positive (upstream) or negative (downstream) responses, for <br />~ example. Chemical inputs, such as natural organic matter, from tributaries, seeps, or <br />flooded lands may provide gross cues for locating a spawning reach, and more subtle <br />cues, such as reproductive byproducts from previously hatched young, may guide the <br />fish to more specific locations within the spawning reach (reviewed by Tyus 1990). <br />Spawning probably occurs within a very small part of the range inhabited by each <br />population of the Colorado pikeminnow. The location of each spawning site could be <br />established unequivocally if spawning activity were observed directly. However, direct <br />observation is unlikely because mature adults are rare and the rivers they inhabit are <br />usually so turbid that the substrate is obscured. Consequently, most evidence <br />supporting the case for spawning at a particular site will be circumstantial. Deductions <br />~ can be based on the movements and reproductive status of adults, as well as the <br />geographic distribution of larvae. Because newly-hatched larvae have virtually no <br />ability to swim against river currents, their presence establishes a downstream limit on <br />the area within which spawning must have occurred. Captures of larvae can be used to <br />delineate a "suspected" spawning area, according to USFWS criteria (USFVNS 1987). <br />~ The boundary is imprecise insofar as the larvae can drift far from the spawning area in <br />just a few hours after hatching. <br />The movements of mature adults at the time of spawning also can provide <br />circumstantial evidence for the location of spawning sites. Radiotracking of adults has <br />~ shown convergence of individuals on a particular river reach (e.g., Tyus 1985, 1990). <br />Tracking data can define a "suspected" spawning area, but the boundaries of the river <br />reach are likely to be overly broad because it may be very difficult to distinguish staging <br />areas from the actual sites of egg deposition. <br />. Collection of adults in "ripe" condition indicates that a spawning area is in close <br />proximity, although adults can still move some distance when in ripe condition (cf. Tyus <br />et al. 1987). The presence of ripe males is sufficient for defining a "suspected" <br />spawning area (USFWS 1987). Females in "running ripe" condition are close enough <br />to the time of egg laying that they are unlikely to move far. Thus, the USFWS (1987) <br />has set the presence of running ripe females as the only acceptable criterion for <br />~ "confirming" the location of a spawning site. <br />The best documentation of spawning sites has come from the Green River system. <br />The reproductive ecology of Colorado pikeminnow in the Green River basin was studied <br />intensively over a 10-year period (1980-90), in which studies encompassed many facets <br />~ of reproduction including the timing of pre- and past-spawning movements, the extent <br />and duration of spawning, and habitat use (Haynes et al. 1984, Nester et al. 1988, Tyus <br />1990, Tyus and Haines 1991). Thousands of larvae and juveniles were captured, the <br />movements of 150 radio-tagged adults were monkored, and 233 fish were captured in <br />breeding condition. In the Yampa River, evidence supporting a suspected spawning <br /> <br />13 <br /> <br />