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<br />6 Abundance Trends and the Status of the little Colorado River Population of Humpback Chub 1989-2006
<br />
<br />Sainsbury, 1980; Kirkwood and Somers, 1984; Francis, 1988).
<br />Typically, k will be negatively biased and L~ will be posi-
<br />tively biased using this technique. This, in turn, has prompted
<br />others to develop models to account and adjust for these biases
<br />(e.g., James, 1991; Wang and others, 1995; Laslett and others,
<br />2002). Van Bertalanffy growth functions were estimated with
<br />these most recent methods. In general, however, poor results
<br />were obtained primarily because of the inability of the models
<br />to predict growth increments exhibited by small fish and large
<br />fish simultaneously (U.S. Geological Survey, unpub. data,
<br />2007). These results suggest a "kink" in the growth curve, as
<br />would be found if fish grew along one curve when small and
<br />then switched to another when larger.
<br />Because water temperature is a dominant driver of
<br />metabolic rate, and hence the von Bertalanffy k parameter
<br />(Paloheimo and Dickie, 1966; Essington and others, 2001), the
<br />"kink" hypothesis is consistent with a fish that is demonstrat-
<br />ing an ontogenetic shift among habitats that have different
<br />water temperatures, as is the case with the LCR and mainstem
<br />Colorado River. The current reproductive ecology paradigm
<br />for the LCR population of HBC is that most successful recruit-
<br />ment occurs when fish are spu'N"Iled and reared within the LCR
<br />(Valdez and Ryel, 1995; Gorman and Stone, 1999). However,
<br />as fish approach some critical size, it is thought that they begin
<br />to engage in a potaclromous migration between the Colorado
<br />River and the LCR (Gorman and Stone, 1999). This ontoge-
<br />netic shift in primary occupancy between two thermal habitats,
<br />and therefore two basal metabolic rates, could induce a pattern
<br />of shifting growth rate.
<br />To account for this apparent pattern of shifting growth
<br />rate, methods proposed by Walters and Essington (University
<br />of British Columbia, University of Washington, respectively;
<br />written commun.; 20(7) (hereafter Walters and Essington,
<br />written commun., 20(7) to tit growth increment data to a gen-
<br />eral growth model (Paloheimo and Dickie, 1965) were used to
<br />describe the rate of change in weight as:
<br />
<br />d1V m. va .1.T..TYI
<br />"- - IlfY -111 'f'
<br />dt -. ,
<br />
<br />where the first term describes anabolic (i.e., mass-acquisition)
<br />processes and is governed by a teml representing the mass-
<br />normalized rate at which the animal acquires mass (ll), the
<br />mass of the animal (W), and a parameter (d) describing the
<br />scaling of the anabolic process with mass. The second term
<br />represents catabolic (Le., mass loss through basal metabolism,
<br />activity, and gonad production) processes where m is the mass-
<br />normalized rate at which the animal looses mass ane!ll is the
<br />scaling factor of catabolic processes with mass. Assuming a
<br />constant relationship between length and weight over time as:
<br />
<br />l'V:::::at',
<br />
<br />where L is length and a and b are constant, simple algebra
<br />provides an analogous relationship for the rate of change in
<br />length as:
<br />
<br />0)
<br />
<br />dL =a[," [f!
<br />_ - K._ .
<br />dt
<br />
<br />Constants in this relationship are related to those in equa-
<br />tions (5) and (6) as:
<br />
<br />a"-JH
<br />a:::::~
<br />b '
<br />
<br />(8)
<br />
<br />a"-'m
<br />K-
<br />--;-'
<br />
<br />(9)
<br />
<br />(10)
<br />
<br />o:::::bd-b+t,and
<br />
<br />(11)
<br />
<br />17 ::::: bn - b + t .
<br />
<br />Essington and others (2001) review these relationships
<br />and describe the derivation of the standard von Bertalanffy
<br />growth function as the integral of equation (7) when n = 1, b
<br />= 3. and d = 2/3. This is the situation where catabolism scales
<br />linearly with mass, the length-weight relationship is isometric,
<br />anabolism scales as 213 mass, and results in the standard von
<br />Bertalantfy growth model:
<br />
<br />(12)
<br />
<br />L(/)::::: Ljt_e-k(H"J),
<br />
<br />(5)
<br />
<br />where lo is the theoretical age where body length is equal to zero.
<br />Walters ane! Essington (written commun., 2007) then
<br />define a protocol to estimate the parameters of equation (5).
<br />To restate this definition, tlrst assume that measurement errors
<br />in the length offish are normal with variance a,:, and that all
<br />fish follow a standard von Bertalanffy growth curve (equa-
<br />tion (2) with shared k and individual L~.. The predicted length
<br />of fish at time of recapture can be found by rearranging the
<br />Fabens equation (4) as:
<br />
<br />(13)
<br />
<br />L(t + ~t)::::: L(t) + (Lx - L(t)XI- e-k<J.I).
<br />
<br />Assuming that individual L~ is nomlaUy distributed with vari-
<br />ance ai ' the variance of eachL(t + M) given L(t), mean L7.'
<br />(1 Z. and (1,~, will be:
<br />
<br />(6)
<br />
<br />(14)
<br />
<br />cr1(r+&)1 ::::: cr,;, (1 + e2kAr. )+ crt (1
<br />
<br />J.
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