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291 <br />offspring of one mature female. Our initial assump- <br />tion was that, in fish of the slow-growing stock, <br />growth rate, length at maturity, and age at maturity <br />resembled these characteristics in upper-basin Col- <br />orado squawfish, whereas in the fast-growing <br />stock, squawfish growth resembled that in Os- <br />mundson's pond (Fig. 2). First maturity of Col- <br />orado squawfish in the upper basin occurs at a <br />length of about 428 mm and an age of about 6 years <br />(Seethaler 1978) - a size that pond-raised fish might <br />conceivably reach in about half that time (Fig. 2). <br />For computational convenience, we assumed that <br />maturity in both simulated stocks occurred at a <br />length of 410 mm and the age of 6 years in slow- <br />growing fish and 3 years for fast-growing fish. <br />There are no definitive data on the survival of <br />Colorado squawfish in its natural environment. In <br />the simulations, we therefore assumed annual mor- <br />talities of 80, 90, 95 or 99% in the shortest length <br />class (10-110 mm TL), and 20% in each of the three <br />larger, 100 mm length classes. Although arbitrary, <br />these rates were chosen because such rapid, early- <br />life mortality and a reduced, constant rate for later <br />ages is typical of many freshwater fishes (e.g. <br />Weatherley 1972). Thus the important difference <br />between our simulated populations was the length <br />of time that fish remained in each length class, <br />which was determined by growth rate. The number <br />of fish that died in each length class was calculated <br />as the product of the initial number of fish, mortal- <br />ity rate, and duration of time spent within the <br />length class. For periods longer than 1 year, the <br />number of deaths during the first year was calculat- <br />ed as described above, and deaths during the re- <br />maining time period were similarly calculated for <br />fish that survived the first year. <br />Vital statistics for our simulated populations <br />were calculated using equations provided by Krebs <br />(1972): <br />r = (IogeR°)lG, and f = e', <br />where G = mean length of generation (the mean <br />period between the birth of the parent and that of <br />offspring); lx = age-specific survival; mX = age- <br />specific effective birth rate; x =age in years; R° _ <br />E lXmx =net reproductive rate (the number of ma- <br />ture female offspring produced in the lifetime of a <br />female parent); r = intrinsic rate of natural in- <br />crease; and f =finite rate of increase (the multipli- <br />cation factor by which the adult female stock will <br />annually grow if that particular value of R° is main- <br />tained). For these computations it was assumed <br />that, beginning in the first year of maturity and <br />continuing through age 10 (arbitrarily taken to be <br />the age of last reproduction), each slow- and fast- <br />growing female produces mature female offspring <br />at an annual rate equal to the age-specific effective <br />birth rate (mx) for our simulated stocks when early- <br />life mortality was 95 or 99%. <br />Development of the hypothesis <br />Temperature regime analysis <br />Although the temperature data used in our analys- <br />es do not reflect the precise temperatures that Col- <br />orado squawfish may experience throughout their <br />life history, they nonetheless allow demonstration <br />of the marked differences in suitability for Col- <br />orado squawfish growth among the temperature <br />regimes of present and historic habitats. If growth <br />of Colorado squawfish occurs only when water <br />temperatures exceed 13° C, growing seasons in the <br />upper-basin river reaches are less than 6 months, <br />whereas they were 7 to 9 months in historic, pre- <br />development, middle- and lower-basin reaches <br />(Fig. 3). Comparisons among these temperature <br />regimes are more useful, however, if their relative <br />suitability for Colorado squawfish growth is con- <br />sidered. Such suitability was estimated as 3.2, 2.1, <br />3.9, and 4.9 for the Green River and the upper, <br />middle, and lower Colorado River, respectively. <br />If the bias in the Green River data (Fig. 3) was as <br />large as our worst-case estimate of 1° C, and we <br />reduced these data by that amount, the Green Riv- <br />erwould continue to warm earlier and have warm- <br />er temperatures during the growing season than <br />does the upper Colorado. Suitability of the Green <br />River temperature regime would be 2.7, an aver- <br />age value 29% larger than that of the upper Col- <br />orado. <br />