Pattern of neutral polymorphism in a geographically structured population
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Genet Res 18 (2): 125-131 (October 1971)

In a two-dimensional stepping-stone model of finite size, if a pair of alleles happen to segregate in the whole population, marked local differ- entiation of gene frequencies can occur only if migration between colonies is sufficiently rare so that Nm < 1, where N is the effective size of each colony and m is the rate at which each colony exchanges individuals with four surrounding colonies each generation. On the other hand, if Nm ^ 4, the whole population behaves as if it were panmictic and the allelic fre- quencies become uniform over the entire distribution range unless muta- tion is unusually high. Tendency toward local differentiation is much weaker in two-dimensional than in one-dimensional habitats.
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