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1 SS Conserntion Genetics of Desert Fishes <br />Introduction <br />Fishes in the Sonoran, Mohave, Great Basin, and Chihua- <br />huan deserts of North America aze declining at an alarttl- <br />ing rate. At least 20 taxa have gone extinct in the last <br />several decades (Meffe, in press), and many more are in <br />immediate danger of similaz fates (Williams et al. 1985). <br />This major conservation problem has prompted govern- <br />mental agencies and academic scientists to conduct re- <br />search on basic ecologies and life histories, develop <br />propagation programs for species perpetuation and re- <br />introduction, and restore degraded habitats. We address <br />here a different aspect of conservation of this highly <br />endangered fauna, the application of population genet- <br />ics to management and long-term perpetuation of desert <br />fishes. <br />Desert Aquatic Systems <br />North American deserts aze extremely arid, with high <br />geographic relief, heavily dissected drainages, and <br />widely dispersed surface waters. Natural aquatic sys- <br />tems in these regions aze fragmented on both broad <br />geographic and local scales. In addition to being iso- <br />lated, desert aquatic systems can experience enormous <br />physico-chemical fluctuations, as in streams and rivers <br />(Hinckley & Meffe 1987), or constant but hazsh condi- <br />tions, as in some springs (Deacon & Hinckley 1974). <br />The extreme isolation and severe abiotic conditions <br />have led to a high degree of phenotypic divergence of <br />fishes, often with small populations living in small hab- <br />itats (Miller 1958). Many taxa aze relicts from better- <br />watered times that have been trapped in isolated springs <br />and streams during the last 10,000- to 12,000-yeaz post- <br />pluvial period. This has resulted in a high degree of <br />endemism (Williams et al. 1985 ). <br />Several genetic and demographic consequences of <br />such an isolated, fragmented distribution are likely, in- <br />cluding (1) local divergence via natural selection or <br />genetic drift; (2) little or no gene IIow among isolated <br />demes that might otherwise moderate losses of genetic <br />variability after population crashes; and (3) little or no <br />recolonization of isolated habitats after local extinction. <br />Southwestern desert fishes aze thus a vulnerable, "~- <br />tinction-prone" goup due to the limited geographic <br />range and isolation of many species. <br />Added to this natural fragility aze major anthropo- <br />genic disturbances of two types: habitat destruction <br />(impoundment and diversion of rivers, groundwater <br />pumping, and drying of surface springs) and the intro- <br />duction of a huge diversity of exotic fishes, many of <br />whom prey on, compete with, or hybridize with native <br />species (Naiman & Soltz 1981). Management of such a <br />Mege and Viijenhoek <br />fauna over the long term must therefore consider the <br />genetic consequences both of natural fragmentation <br />with small population sizes and of artificial disruption by <br />man. General and philosophical aspects of genetic man- <br />agement of fishes have been outlined elsewhere (Meffe <br />1986, 1987; Ryman & Utter 1987). Here we consider <br />more specific approaches of genetic studies and exper- <br />imental manipulations of raze and endangered south- <br />western fishes. <br />Divergence, Migration, and Gene Flow <br />Two demographic factors, genetically effective popula- <br />tion size and migration rate, affect the degree of diver- <br />gence among local colonies of fishes. Colonies diverge <br />from one another as a consequence of local selection <br />pressures, mutation, and random genetic drift; drift <br />alone can cause considerable divergence among small <br />colonies. The rate of divergence due to drift in turn <br />depends on the genetically effective size of the local <br />populations (Ne). Ne represents the number of breeding <br />adults in each generation, and is affected by sex ratio, <br />pattern of mating, and variance in reproductive output <br />among individuals (Craw & Kimura 1970). <br />Gene flow via migration (m, or the proportion of in- <br />dividuals exchanged between colonies per generation) <br />maintains genetic variation within colonies and retazds <br />divergence among colonies. Divergence occurs as a <br />product of Ne and m. if Ne is small, colonies tend to <br />diverge rapidly, as a result of random processes. High <br />rates of gene IIow (m) aze needed to maintain genetic <br />homogeneity among them. As long as N~rn > 1, local <br />colonies will tend not to diverge significantly with re- <br />gazd to the types of alleles present (Allendorf 1983)• <br />Thus, a pair of large populations with mean Ne > 10,000 <br />that exchange individuals at a rate of m = 0.001 will not <br />diverge significantly by chance alone, since Nenn > 10. <br />However, a pair of small populations with Ne < 1000 <br />and with a similaz rate of gene exchange would diverge, <br />sitlce Ne < 1. <br />This scenario is based on the assumption that colonies <br />are dispersed geographically as if they were islands and <br />gene flow can occur equally among all the islands. As we <br />discuss below, this scenario may be an appropriate de- <br />scription for some aquatic systems (the Death Valley <br />model), but for most freshwater fishes, riverine habitats <br />impose a more complex population structure (the <br />Stream Hierarchy model). <br />Zoogeographic Models of Gene Flow <br />We envision two models of genetical population struc- <br />ture in desert fishes. The existence of these different <br />Conservation Biology <br />Volume 2, No. 2, Junc-1988 <br />