Group selection among alternative evolutionarily stable strategies

Many important models of the evolution of social behavior have more than one evolutionarily stable strategy (ESS). Examples include co-ordination games, contests, mutualism, reciprocity, and sexual selection. Here we show that when there are multiple evolutionarily stable strategies, selection among groups can cause the spread of the strategy that has the lowest extinction rate or highest probability of contributing to the colonization of empty habitats, and that this may occur even when groups are usually very large, migration rates are substantial, and "extinction" entails only the disruption of the group and the dispersal of its members. The main requirements are: (1) individuals drawn from a single surviving group make up a sufficiently large fraction newly formed groups, and (2) the processes increasing the frequency of successful strategies within groups are strong compared to rate of migration among groups. The latter condition suggests that this form of group selection will be particularly important when behavioral variation is culturally acquired.