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increase is real, as we suspect, it suggests an increase in food availability as Colorado <br />squawfish reach this size. Increased gape size, or more effective foraging and handling <br />abilities may allow utilization of larger and more abundant prey. <br />While there is much variation in growth among individual fish, it appears that in successive <br />years most fish do not grow consistently faster or slower than the average and average growth <br />rates do not differ greatly among years. Nevertheless, there can be large differences among <br />individuals in time it takes to reach a given length: there was a 20-year range of potential ages <br />for a given length for fish > 800 mm TL. Although length is therefore a rather imprecise <br />indicator of age, we can conclude that the largest fish present in the population are quite old. <br />For individuals 900 mm long, 34 years might be a minimum age, but an average age would <br />likely be 47-55 years (Fig. 3 a and b). These fish are very rare: of approximately 1,080 <br />subadult and adult Colorado squawfish captured from the Colorado River during 1979-1995 <br />only three have been > 900 mm TL (0.28%) and two of these were < 905 mm TL. The <br />largest, captured in 1990 by UDWR biologists, was 960 mm TL, considerably larger than any <br />other reported from upper basin rivers (McAda et al. 1994). <br />There are numerous anecdotal accounts of Colorado squawfish historically reaching lengths of <br />1,200-1,800 mm TL (e.g., Quarterone 1993), far larger than any found during the past 20 <br />years. Two explanations for the disappearance of such large individuals have been proposed: <br />1) growth rate has declined (Behnke and Benson 1983), and 2) survival rate has declined <br />(Gilpin 1993). Behnke and Benson (1983) suggested that extirpation of the bonytail (Gila <br />elegans) impacted the food supply of Colorado squawfish. Gilpin (1993) showed, through <br />simulations, that average and maximum fish size would significantly increase if adult survival <br />rate was increased to 0.95, and suggested that rates may have declined due to angling <br />mortality. However, neither hypothesis adequately explains the disappearance of very large <br />individuals. Kaeding and Osmundson (1988) concluded that slow growth in the upper basin <br />was the historic norm because temperature regimes of mainstem rivers have, with few <br />exceptions, remain unchanged and potential foods, Gila spp. and native suckers, remain <br />plentiful. Also, the size structure of Gilpin's simulated population with a survival rate of 0.95 <br />indicated that the largest individuals would still be < 1,000 mm long. <br />Though the preceding two explanations may explain in part the reduction in average and <br />maximum size, we offer a third hypothesis: very large fish may have attained their size while <br />residing in the Lower Colorado River Basin (downstream of Lee's Ferry) and moved <br />upstream later in life where they were eventually captured. Kaeding and Osmundson (1988) <br />demonstrated that longer growing seasons and warmer temperatures in the lower basin during <br />historic times would have provided 1.5-2.3 times the annual thermal units for growth than that <br />in upper basin reaches. Additionally, Colorado squawfish are capable of making long-distance <br />movements; radio-tagged adults have been documented traversing the entire length of their <br />current range in the upper Colorado River (313 km) in less than 3 months (McAda and <br />A-17