The evolution of human cooperation.

Darwin viewed cooperation as a perplexing challenge to his theory of natural selection. Natural selection generally favors the evolution of behaviors that enhance the fitness of individuals. Cooperative behavior, which increases the fitness of a recipient at the expense of the donor, contradicts this logic. William D. Hamilton helped to solve the puzzle when he showed that cooperation can evolve if cooperators direct benefits selectively to other cooperators (i.e. assortment). Kinship, group selection and the previous behavior of social partners all provide mechanisms for assortment (Figure 1), and kin selection and reciprocal altruism are the foundation of the kinds of cooperative behavior observed in many animals. Humans also bias cooperation in favor of kin and reciprocating partners, but the scope, scale, and variability of human cooperation greatly exceed that of other animals. Here, we introduce derived features of human cooperation in the context in which they originally evolved, and discuss the processes that may have shaped the evolution of our remarkable capacity for cooperation. We argue that culturally-evolved norms that specify how people should behave provide an evolutionarily novel mechanism for assortment, and play an important role in sustaining derived properties of cooperation in human groups.