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<br />Dizon et ,}, <br /> <br />differentiated. Presumably the genetic stock is the pop- <br />ulation that Sinclair (1988) called the "local popula- <br />tion" in his essay on population regulation and specia- <br />tion. Recently, Gauldie (1991) published a thorough <br />review of stock concepts and their deficiencies as ap- <br />plied to exploited fish populations and argued for a <br />model of populations defined on the basis of degree of <br />interchange. <br />This sense of biological uniqueness via isolation is an <br />evolutionary one, because it acknowledges the popula- <br />tion's adaptation to local conditions and indicates that it <br />possesses a reservoir of unique genetic variability, This <br />is the "evolutionarily significant unit" (ESU) (Ryder <br />1986; see Waples 1991 for a review of its application to <br />northwestern salmonid stocks); it is a biological popu- <br />lation that is distinguished by its presumed 'evolutionary <br />uniqueness and significance, It is a natutal unit and <br />should be a better management unit than an unnatural <br />unit (Sinclair 1988). <br />Few would argue with this reasoning. In practice, <br />however, it is difficult to define a stock based upon <br />principles of adaptive genetic uniqueness. Existing mea- <br />sures of fitness are of little use in defining specific ad- <br />aptation to a local environment; experimentally this is <br />an intractable problem for higher animals. As a result, <br />stocks are typically defined using a variety of proxies <br />that suggest this adapted genetic uniqueness. <br /> <br />Stock Criteria <br /> <br />To study stock structure, an investigator makes obser- <br />vations of allopatry or isolation that imply reproductive <br />isolation; differential life history responses, which also <br />imply reproductive isolation; morphological (i.e., pro- <br />tein structures) differentiation, indicating drift or evo- <br />lution under different selective regimes; or differentia- <br />tion of neutral genetic characters quantifying the degree <br />of isolation and the time since an ancestor was shared. <br />We have categorized these sources of information as (a) <br />distributional, (b) population response, ( c) phenotypic, <br />and (d) genotypic. <br /> <br />Distributional Data <br /> <br />Initially the most important items of knowledge are <br />those of distribution and abundance. These data pertain <br />to all aspects of abundance, migration, pollutant and <br />parasite loads, zones of fishery interaction and conflict, <br />etc., that provide information about the population <br />movements relative to geographical space and time. <br />These data largely define whether there are major geo- <br />graphical barriers between putative stocks and whether <br />they are allopatric, parapatric, or sympatric. <br />At the most basic level and without additional infor- <br />mation, disjunct populations frequently have been des- <br />ignated as separate stocks for management purposes <br />(e.g., Perrin et al. 1985). 1bis is certainly a conservative <br /> <br />, ,i <br /> <br />Rethinking the Stock Concept' '1.7 <br /> <br />approach. Even populations that overlap for portions of <br />their life cycle may be disjunct Slocks (lles & Sinclair <br />1982). Overlapping distribution does not necessarily <br />imply gene flow; natural processes favoring adaptation <br />to local conditions, or social selection favoring mainte- <br />nance of local social behaviors, especially in the case of <br />marine mammals, may preserve genetic differentiation <br />in the face of apparent overlap or movements between <br />populations (Ehrlich & Raven 1969; Slatkin 1987; Baker <br />et al. 1990). <br />Presence and absence data on distribution are the <br />most readily available sources of information on which <br />to base stock identification decisions, but they may be <br />misinterpreted when search effort is not continuous, <br />causing artificial discontinuities in the reported distri- <br />bution. Abundance information provides a better proxy <br />for exchange rates, Density "troughs': or large areas of <br />zero density indicate extrinsic barriers. Unfortunately, <br />the variability associated with most estimates of density <br />is such that the statistical power of using differences in <br />density gradients in asSigning stock status is often low. <br />Because of the importance of reproductive isolation <br />in speciation, information on the physical movement of <br />individuals (telemetry studies, mark-and-recapture, <br />etc.) within their ranges brings one closer to properly <br />assigning stock status to populations. Reeb and A vise <br />( 1990) and Avise and Ball ( 1990) described evidence of <br />concordance of genetic patterns with movements for a <br />variety of species in response to biogeographic barriers. <br />Thus, for most populations, research on movement pat- <br />terns should be given high priority. <br /> <br />Population Response Data <br /> <br />Although used frequently to distinguish putative popu- <br />lations, a population's life histories and behavioral traits <br />may be modified by the environment through density- <br />dependent control mechanisms. Data include all aspects <br />of demography (age at sexual maturity, fecundity, <br />growth rate, and mortality), other biological parame- <br />ters, social behavior, vocalization, and specific interac- <br />tions peculiar to a particular population. Behaviotal <br />characteristics are probably modified by exploitation <br />and intra- and interspecies competition, which affect <br />breeding site fidelity, nursing and care-giving to the <br />young, and feeding habits, which in turn affect the pol- <br />lutant and parasite loads. Still, an advantage to using <br />population response criteria over abundance criteria <br />should be recognized: Samples taken at one point in <br />time when pOpulation response criteria are used inte- <br />grate movement patterns over a much greater time scale <br />than do abundance criteria. <br />Differences in the timing of breeding provide a par- <br />ticularly valuable criterion, because they imply that a <br />barrier exists to, gene flow between populations. A stock <br />division of spotted dolphins (Stenella attenuata) in the <br /> <br />ConscrYatlon BJoIogy <br />Volume 6, No.1, March 1992 <br />