Bottlenose dolphins (Tursiops truncatus) prefer to cooperate when petted: Integrating proximate and ultimate explanations II.

Cooperation poses theoretical problems because the behaviors of individuals can benefit others. Evolutionary and game-theory explanations that focus on maximizing one's own material outcomes are usually supported by experimental models with isolated and anonymous subjects. Cooperation in the natural world, however, is often a social act whereby familiar individuals coordinate behaviors for shared outcomes. Social cooperation is also associated with a cooperation bias expressed as a preference for cooperation even when noncooperation is immediately more beneficial. The authors report on evidence for such a bias in a captive group of bottlenose dolphins that voluntarily preferred to receive petting from human guides by using a pairwise coordinated approach, even though this was more difficult, and total petting amount was thereby reduced. To explain why this bias occurs, the authors propose an integrated behavioral-evolutionary approach whereby performance is determined by two kinds of immediate outcomes: material gains and intrinsic affective states associated with cooperating. The latter can provide reinforcement when immediate material gains are reduced, delayed, or absent. Over a lifetime, this proximate mechanism can lead to cooperative relationships whose long-term ultimate consequences can be adaptive.