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It Allendorf This approach should be modified in captive breeding programs in which individuals spend part of their life cycle outside of captivity. For example, anadromous Pa- cific salmon (Oncorhynchus spp) are raised in a fresh- water hatchery for approximately one year and then released so that they can migrate to the ocean. Sexually mature fish return to the hatchery and are used to pro- duce the following generation. In this case, family size should be equalized at the time of release, not at the time of return. This would reduce the selection for ad- aptation to captivity, but would still allow natural selec- tion to occur during the part of the life cycle spent in the wild. Equalizing family size is a potentially important tech- nique to retard adaptation to captivity in species with certain life-history characteristics. In contract, breeding programs based upon pedigree analysis are a much more powerful method to minimize genetic drift and adaptation to captivity in species with relatively low fecundity (such as birds and mammals; Haig et aI. 1990). However, rapid genetic change caused by adaptation to captivity is less likely to occur in such species because their fecundity liinits the potential intensity of selection. In species with high individual fecundity (such as many fish, amphibians, and insects), rapid adaptation to captivity is more likely because hundreds of progeny or more can be produced by individual matings, and it is also often not possible or practical to base matings on pedigrees. In such species, equalizing progeny number should be considered even if captive population sizes are large. Acknowledgments I thank Pedro Lesica, Penny Kukuk, Tom Mitchell-Olds, and Nils Ryman for their helpful comments; Robin Wa- ples for his instructive suggestions on anadromous fish; James F. Crow for his insightful review and for the Haldane and King references; and especially Dick Frankham for his helpful review and comments. Literature Cited Allendorf, F. W., and Ryman, N. 1987. Genetic management of hatchery stocks. Pages 141-159 in N. Ryman and F. M. Utter, editors. Population genetics and fishery management Univer- sity of Washington Press, Seattle, Washington. Chilcote, M. W., S. A. Leider, and J. J. Loch. 1.986. Differential reproductive success of hatchery and wild summer-rlln steel- head under natural conditions. Transactions of the American Fisheries Society 115:726-735. Crow, J. F., and C. Denniston. 1988. Inbreeding and variance effective population numbers. Evolution 42:482-495. Crow, J. F., and M. Kimura. 1970. An introduction to popula- tion genetics theory. Burgess, Minneapolis, Minnesota. Darwin, C. 1868. The variation of animals and plants under domestication. 2 volumes. John Murray, London, England. Adapf3tion to Captive Breeding 419 David, B. 1990. Apache trout culture: An aid to restoration. Endangered Species Update 8:76-78. Frankham, R, and D. A. Loebe!. Modeling conservation genet- ics problems using captive Drosophila populations. 4. Rapid adaptation to captivity. Zoo Biology. In Press. Frankham, R, H. Hemmer, O. A. Ryder, E. G. Cothran, M. E. Soule, N. D. Murray, and M. Snyder. 1986. Selection in captive populations. Zoo Biology 5:127-138. Fraser,]. M. 1981. Comparative survival and growth of planted wild, hybrid, and domestic strains of brook trout (Salvelinus fontinalis) in Ontario Lakes. Canadian Journal of Fisheries and Aquatic Sciences 38:1372-1384. Haig, S. M., J. D. Ballou, and S. R. Derrickson. 1990. Manage- ment options for preserving genetic diversity: Reintroduction of Guam rails to the wild. Conservation Biology 4:290-300. Haldane, J. B. S. 1924. A mathematical theory of natural and artificial selection. Transactions of the Cambridge Philosoph- ical Society 23: 19-41. ]ohnsson, J. I., and M. V. Abrahams. 1991. 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