No.7 wards equilibrium; progress that is ultimately terminated by local extinction. As the extinction rate increases, this equilibration is truncated more quickly, and the founding event comes to dominate. With local extinction and recolonization, the metapopulation will display an age-structure since subpopulations can vary in the time since each was formed. Any difference in the allele frequency variance among newly founded groups relative to that among older populations can cause the genetic variance among the whole array of populations to deviate from equilibrium conditions. It is the purpose of this paper to describe a general population genetic model offounding group formation and to illustrate the effects of a broad range of colonization patterns on the neutral genetic differentiation of populations. Wright (1940) was the first to discuss the Abstract. -Extinction and recolonization in an island model affects genetic differentiation among subpopulations through a combination ofsampling and mixing. We investigate the balance ofthese forces in a general model of population founding that predicts first the genetic variance among new groups and then the effect of these new groups on the total genetic variance among all populations. We allow for a broad range of types of mixing at the time of colonization and demonstrate the significant effects on differentiation from the probability of common origin of gametes (». We further demonstrate that kin-structured founding and inbreeding within populations can have a significant effect on the genetic variance among groups and use these results to make predictions about lineal fission and fusion ofpopulations. These results show that population structure is critically affected by non-equilibrium dynamics and that the properties of new populations, especially founding number, probability of common origin, and kin structure, are vital in our understanding of genetic variation.Received October 23, 1989. Accepted April 9, 1990 Much of theoretical population genetics is simplified by the assumption that the forces ofmigration, random drift, mutation, and selection have reached an equilibrium in their effects on the distribution of gene frequencies among populations. For example, at equilibrium between the forces of migration and drift, differentiation of local populations (FST) in an island model is ex-where N is the local population size and m is the migration rate (Wright, 1940). Nature is not always so patient. In many species, local populations come and go on a temporal scale that does not permit the attainment of that equilibrium. In this case, the distribution ofgene frequencies among populations reflects the interaction oftwo phases of a population's history: first, a perturbation from equilibrium caused by a founding event, followed by the progress to-1717