Cooperation in animals: An evolutionary overview

Evolutionary biologists have grappled with the question of the emergenceand maintenance of cooperation since Darwin first listed animal cooperation asapotential problem for his theory of natural selection. Here I review four pathsthat have been delineated in the study of intra-specific cooperation amonganimals. These paths – kinship, reciprocity, byproduct mutualism andgroupselection – serve as a starting point for behavioral ecologistsinterestedstudying the initiation and maintenance of cooperation. After reviewing theempirical and theoretical underpinnings of these paths to cooperation, I touchupon some recent work that has attempted to examine (or reexamine) the role ofphylogeny, punishment and morality in the light of cooperative behavior.