<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
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