Animal cooperation among unrelated individuals

The evolution of cooperation has long been a topic near and dear to the hearts of behavioral and evolutionary ecologists. Cooperative behaviors run the gamut from fairly simple to very complicated and there are a myriad of ways to study cooperation. Here I shall focus on three paths that have been delineated in the study of intraspecific cooperation among unrelated individuals: reciprocity, byproduct mutualism, and group selection. In each case, I attempt to delineate the theory underlying each of these paths and then provide examples from the empirical literature. In addition, I shall briefly touch upon some recent work that has attempted to examine (or re-examine) the role of cognition and phylogeny in the study of cooperative behavior. While empirical and theoretical work has made significant strides in the name of better understanding the evolution and maintenance of cooperative behavior in animals, much work remains for the future.