EVOLUTION OF ALTRUISM IN KIN-STRUCTURED AND RANDOM SUBDIVIDED POPULATIONS.
A. population structure favorable to the evolution of an altruistic trait is studied by Monte Carlo simulation. The model is based on a small-scale nonindustrial human society but seems generalizable to other highly social mammals. Three hierarchical levels are recognized: 1) the ecologically isolated local group (hamlet) which may be composed of kin and/or unrelated individuals; 2) the deme (settlement) comprising several such groups which interbreed; and 3) the set of demes (metapopulation) among which gene flow occurs. The first two levels of the model are based on D. S. Wilson's structured deme concept; the third allows for gene flow among demes in the metapopulation and for the structured diffusion of alleles across a wider area than might be included within the scope of a single deme. The simulation models genetic drift by a process of hamlet formation which may be random, or variously kin-structured. Hamlets may then become extinct based on a probability function of their gene frequencies. Individual selection within settlements is modeled deterministically, and gene flow among settlements is modeled as two-dimensional steppingstone migration of random or kin-structured groups. Results of the simulations show that, with realistic values for group sizes, moderate extinction rate, and high rates of migration (m > 27%), disadvantageous alleles (s = 10% and 25%) may increase markedly due to differential hamlet extinction over the course of 50 generations. The greater the degree of kin-structuring of founder groups, the higher the variance among hamlets and the faster the rate of increase of the allele for altruism. Nonetheless, even in some randomly founded groups, a clear increase in the altruism gene frequency occurred. It is also notable that kin-structured group selection by hamlet extinction may be effective when the initial frequency of altruism genes is very low (average of one per deme) and among a relatively small number of demes (25). Thus the process of group extinction in a hierarchically structured population allows rapid increase of an allele for altruism under plausible demographic conditions.