multiple populations within groups. Only when armed with
<br />such knowledge can the manager make informed decisions
<br />with respect to preservation of genetic resources.
<br />Electrophoretic analysis of all brood stocks should be con-
<br />ducted to determine levels of genetic variation, from which
<br />all subsequent variation will be derived. Such analysis can
<br />be completed without sacrificing individuals: muscle, fin,
<br />blood, or even skin mucous (M. H. Smith, personal com-
<br />munication) for example, may be obtained for testing with
<br />little damage to the fish. Subsequent captive generations
<br />should also be sampled periodically (every few generations,
<br />if they are retained in captivity for that long) to determine
<br />changes in allele frequencies or detect loss of alleles. Such
<br />programs have revealed significant genetic deterioration of
<br />hatchery salmonid stocks (Ryman and Stahl 1980; Cross and
<br />King 1983; Stahl 1983), and could provide the basis for man-
<br />agement decisions with respect to genetic welfare of captive
<br />endangered fishes. It is only through informed genetic mon-
<br />itoring that we can reasonably expect to maintain rare fishes
<br />for extended periods of time.
<br />In all cases of intervention for management, the largest
<br />feasible Ne should be initiated and maintained in a captive
<br />breeding program (Table 1). "It is inevitable that some of
<br />the natural variability of a species is inadvertently lost when
<br />individuals are established as founders of a hatchery stock,
<br />and that more and more variability is inadvertently lost in
<br />each generation of intensive hatchery production" (Wilkins
<br />1981, p. 217). As a result, the "basic rule of conservation
<br />genetics" (Soule 1980) is that a genetically effective popu-
<br />lation size of 50 is the minimum requirement for short-term
<br />survival (on the order of several generations). This number
<br />will result in a 1% AF per generation, which is tolerable for
<br />several generations. For long-term genetic health, Franklin
<br />(1980) suggests a minimum Ne of at least 500, with an as-
<br />sociated inbreeding coefficient of 0.1% and a low rate of
<br />genetic drift. Of course, larger populations than this are
<br />desirable. Ne in any case may be maximized by maintaining
<br />equal sex ratios of breeding adults, insuring even progeny
<br />distribution by individual breeding and culling of excess
<br />offspring (Frankel and Soule 1981), and avoiding population
<br />crashes. Such action will minimize further erosion of quan-
<br />titative genetic variation through bottlenecks or drift, and
<br />will reduce loss of rare alleles and inbreeding effects. These
<br />problems can also be countered by periodic introduction of
<br />wild individuals (from the same locality as the original brood
<br />stock) into the captive population.
<br />If a large population cannot be acquired or maintained in
<br />captivity, inbreeding can still be avoided by a controlled
<br />selective breeding program that minimizes relatedness of
<br />mated individuals (Tave 1984). One example of such a breed-
<br />ing scheme is outlined in Fig. 5; many other schemes are
<br />possible (e.g., Flesness 1977; Senner 1980). An innovative
<br />program that actually utilizes inbreeding to an advantage in
<br />cases where few founders are available and others cannot
<br />be obtained was successfully applied to the rare Speke's
<br />Gazelle (Gazella spekel) by Templeton and Reed (1983).
<br />Stocks should be kept in hatchery environments only long
<br />enough to accomplish outlined breeding goals such as sci-
<br />entific investigation or building a reintroduction stock. Min-
<br />imizing time spent under artificial conditions reduces the
<br />probability of bottlenecks, drift, inbreeding, artificial selec-
<br />tion, domestication, catastrophic losses or disease. By "cir-
<br />culating" populations through a hatchery and quickly back
<br />P
<br />F?
<br />F2
<br />Figure 5. One potential scheme for reducing inbreeding in a small
<br />captive population, starting with a parental stock of only three
<br />males and three females. In each generation, offspring are out-
<br />crossed with other lines rather than allowed to freely interbreed
<br />as one common stock. The latter practice would result in sib coat-
<br />ings and an increase in inbreeding.
<br />into the wild, one can avoid most of the pitfalls associated
<br />with genetic changes in captivity, and thereby maintain
<br />healthier populations of endangered fishes. Smith and
<br />Chesser (1981, p. 18) emphasized this point: "To prevent
<br />the loss of genetic variability caused by drift and inbreeding,
<br />it is important that hatchery and restocking programmes do
<br />not maintain spawning stocks as independent units for long
<br />periods (Ryman and Stahl 1980). Periodic replacement of
<br />hatchery fish with those from natural populations is also
<br />advised to prevent possible adaptations of fish to hatchery
<br />conditions; these adaptations may not be advantageous in
<br />natural environments."
<br />Separate stocks of isolated populations (Table 1) will max-
<br />imize inter-population variance, maintaining potentially dis-
<br />tinct forms and thereby allowing greatest flexibility in man-
<br />agement. For example, if a particular field locality is to be
<br />stocked with an endangered fish, we have the option of
<br />selecting from among several stock populations, and can
<br />choose the one whose habitat of origin most closely matches
<br />the new site. By not mixing wild populations, we retain the
<br />option of experimentally combining portions of these denies
<br />in the future, should adverse effects of small populations
<br />appear; if initially combined, however, the populations can-
<br />not later be separated.
<br />A Final Caution
<br />I have herein outlined theoretical aspects of genetic perils
<br />faced by small populations, supported by appropriate data
<br />wherever possible, and have suggested actions that may be
<br />taken to minimize these problems. For many species, stated
<br />goals simply cannot be met or even closely approximated
<br />because facilities or budgets are not adequate, or the number
<br />of extant individuals is minimal (e.g., Gambusia gagei, Poe-
<br />
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<br />20 Fisheries, Vol. 11, No. i
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