Abstract: Coalitionary support in agonistic interactions represents cooperation because intervening in a fight is potentially costly to the donor of support but benefits the recipient. Here, we first review the characteristics of, and evolutionary forces favoring, intragroup coalitions in 49 species and find that patterns of intragroup coalition formation are remarkably similar between primates and nonprimates. We then test hypotheses suggesting kin selection, reciprocal altruism, and direct benefits as adaptive explanations for coalitionary interventions among adult female spotted hyenas (Crocuta crocuta) belonging to a large social group in Kenya. As predicted by kin selection theory, females supported close kin most often, and the density (connectedness) of cooperation networks increased with genetic relatedness. Nevertheless, kinship failed to protect females from coalitionary attacks. We found no evidence of enduring alliances based on reciprocal support among unrelated adult females. Instead, donors generally minimized costs to themselves, intervening most often during low-intensity fights and when feeding opportunities were unavailable. Females also gained direct benefits from directing coalitionary attacks toward subordinates. Finally, females monitored the number of dominant bystanders in the “audience” at fights and modified their level of cooperation based on this knowledge. Overall, hyenas made flexible decisions regarding whether or not to intervene in fights, modifying their tendency to cooperate based on multiple types of information about their immediate social and ecological environments. Taken together, these findings indicate that the combined evolutionary forces of kin selection and direct benefits derived from reinforcing the status quo drive coalitionary interventions among adult female spotted hyenas.

Abstract: Triver's model of reciprocal altruism, and its descendants based on the Prisoner's Dilemma model, have dominated thinking about cooperation and altruism between non-relatives. However, there are three alternative models of altruism directed to non-relatives. These models, which are not based on the Prisoner's Dilemma, may explain a variety of phenomena, from allogrooming among impala to helping by non-relatives in cooperatively breeding birds and mammals.

Abstract: Prior modeling work has found that pure winner and loser effects (i.e., changing the estimation of your own fighting ability as a function of direct prior experience) can have important consequences for hierarchy formation. Here these models are extended to incorporate “bystander effects.” When bystander effects are in operation, observers (i.e., bystanders) of aggressive interactions change their assessment of the protagonists' fighting abilities (depending on who wins and who loses). Computer simulations demonstrate that when bystander winner effects alone are at play, groups have a clear omega (bottom-ranking individual), while the relative position of other group members remains difficult to determine. When only bystander loser effects are in operation, wins and losses are randomly distributed throughout a group (i.e., no discernible hierarchy). When pure and bystander winner effects are jointly in place, a linear hierarchy, in which all positions (i.e., {alpha} to {delta} when N = 4) are clearly defined, emerges. Joint pure and bystander loser effects produce the same result. In principle one could test the predictions from the models developed here in a straightforward comparative study. Hopefully, the results of this model will spur on such studies in the future.

Abstract: We present the results of a series of computer simulations that examined the impact of winner, loser, and bystander effects on hierarchy formation in fused groups. These effects and their implications for hierarchy structure and aggressive interactions were first examined in small four-member groups. Subsequent to this, the two small groups were fused into a single larger group. Further interactions took place in this fused group, generating a new hierarchy. Our models demonstrate clearly that winner, loser, and bystander effects strongly influence both the structure and types of interactions that emerge from the fusion of smaller groups. Four conditions produced results in which the same general patterns were uncovered in pre- and postfusion groups: (1) winner effects alone, (2) bystander loser effects alone, (3) winner and bystander winner effects operating simultaneously, and (4) all four effects in play simultaneously. Outside this parameter space, hierarchy structure and the nature of aggressive interactions differed in pre- and postfusion groups. When only loser effects were in play, one of the two clear alphas from the prefused groups dropped in rank in the eight-member fused group. When bystander winner effects were in play, it was difficult to rank any of the eight individuals in the fused group, and players interacted almost exclusively with those that were not in their original four-member group. When loser and bystander loser effects operated simultaneously, two top-ranking individuals emerged in the fused groups, but the relative rank of the other players was difficult to assign.

Abstract: Summary The introduction of game-theoretical thinking into evolutionary biology has laid the groundwork for a heuristic view of animal behaviour in which individuals employ “strategies” – rules that instruct them how to behave in a given circumstance to maximize relative fitness. Axelrod and Hamilton (1981) found that a strategy called Tit-For-Tat (TFT) is one robust cooperative solution to the iterated Prisoner's Dilemma game. There exists, however, little empirical evidence that animals employ TFT. Predator inspection in fish provides one ecological context in which to examine the use of the TFT strategy.