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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 /> <br /> <br />20 Fisheries, Vol. 11, No. i <br />