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
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