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. "From the point of view of the moralist, the animal world is on about the same level as the gladiator's show. The creatures are fairly well treated, and set to fight; whereby the strongest, the swiftest and the cunningest live to fight another day. The spectator has no need to turn his thumb down, as no quarter is given em leader the weakest and the stupidest went to the wall, while the toughest and the shrewdest, those who were best fitted to cope with their circumstances, but not the best in any other way, survived. Life was a continuous free fight, and em leader a war of each against all was the normal state of existence." (Huxley 1888)