Demography and the tragedy of the commons
  Individual success in group-structured populations has two components. First, an individual gains by outcompeting its neighbors for local resources. Second, an individual's share of group success must be weighted by the total productivity of the group. The essence of sociality arises from the tension between selfish gains against neighbors and the associated loss that selfishness imposes by degrading the efficiency of the group. Without some force to modulate selfishness, the natural tendencies of self interest typically degrade group performance to the detriment of all. This is the tragedy of the commons. Kin selection provides the most widely discussed way in which the tragedy is overcome in biology. Kin selection arises from behavioral associations within groups caused either by genetical kinship or by other processes that correlate the behaviors of group members. Here, I emphasize demography as a second factor that may also modulate the tragedy of the commons and favor cooperative integration of groups. Each act of selfishness or cooperation in a group often influences group survival and fecundity over many subsequent generations. For example, a cooperative act early in the growth cycle of a colony may enhance the future size and survival of the colony. This time-dependent benefit can greatly increase the degree of cooperation favored by natural selection, providing another way in which to overcome the tragedy of the commons and enhance the integration of group behavior. I conclude that analyses of sociality must account for both the behavioral associations of kin selection theory and the demographic consequences of life history theory. 
Steven A. Frank
q-bio.PE