Abstract: To examine when and why animals break up fights between others in their group, I modelled whether ‘winner’ and ‘loser’ effects might be one element driving the evolution of intervention behaviour. I considered one particular type of intervention: when the intervener simply breaks up fights between two others, but does not favour either party in so doing. When victories at time T + 1 are more likely given a victory at time T (i.e. winner effects), intervention is often favoured. Intervention is favoured in these circumstances because the intervening party in essence stops others from ‘getting on a roll’ and climbing up any hierarchy that exists. However, when loser effects alone are at work (defeats at time T + 1 are more likely given a defeat at time T), breaking up fights between others is never selected. If both winner and loser effects are operating simultaneously, then the likelihood of intervention behaviour evolving is a function of the relative strength of these two effects. The greater the winner effect relative to the loser effect, the more likely intervention behaviour is to evolve.

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: 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.

Abstract: Bekoff [J. Consci. Stud. 8 (2001) 81] argued that mammalian social play is a useful behavioral phenotype on which to concentrate in order to learn more about the evolution of fairness. Here, we build a game theoretical model designed to formalize some of the ideas laid out by Bekoff, and to examine whether `fair' strategies can in fact be evolutionarily stable. The models we present examine fairness at two different developmental stages during an individual's ontogeny, and hence we create four strategies--fair at time 1/fair at time 2, not fair at time 1/not fair at time 2, fair at time 1/not fair at time 2, not fair at time 1/fair at time 2. Our results suggest that when considering species where fairness can be expressed during two different developmental stages, acting fairly should be more common than never acting fairly. In addition, when no one strategy was evolutionarily stable, we found that all four strategies we model can coexist at evolutionary equilibrium. Even in the absence of an overwhelming database from which to test our model, the general predictions we make have significant implications for the evolution of fairness.

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.