Abstract: A key aspect of human behaviour is cooperation1, 2, 3, 4, 5, 6, 7. We tend to help others even if costs are involved. We are more likely to help when the costs are small and the benefits for the other person significant. Cooperation leads to a tension between what is best for the individual and what is best for the group. A group does better if everyone cooperates, but each individual is tempted to defect. Recently there has been much interest in exploring the effect of costly punishment on human cooperation8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23. Costly punishment means paying a cost for another individual to incur a cost. It has been suggested that costly punishment promotes cooperation even in non-repeated games and without any possibility of reputation effects10. But most of our interactions are repeated and reputation is always at stake. Thus, if costly punishment is important in promoting cooperation, it must do so in a repeated setting. We have performed experiments in which, in each round of a repeated game, people choose between cooperation, defection and costly punishment. In control experiments, people could only cooperate or defect. Here we show that the option of costly punishment increases the amount of cooperation but not the average payoff of the group. Furthermore, there is a strong negative correlation between total payoff and use of costly punishment. Those people who gain the highest total payoff tend not to use costly punishment: winners don't punish. This suggests that costly punishment behaviour is maladaptive in cooperation games and might have evolved for other reasons.

Abstract: In recent years evidence has been accumulating that personalities are not only found in humans but also in a wide range of other animal species. Individuals differ consistently in their behavioural tendencies and the behaviour in one context is correlated with the behaviour in multiple other contexts. From an adaptive perspective, the evolution of animal personalities is still a mystery, because a more flexible structure of behaviour should provide a selective advantage. Accordingly, many researchers view personalities as resulting from constraints imposed by the architecture of behaviour (but see ref. 12). In contrast, we show here that animal personalities can be given an adaptive explanation. Our argument is based on the insight that the trade-off between current and future reproduction often results in polymorphic populations in which some individuals put more emphasis on future fitness returns than others. Life-history theory predicts that such differences in fitness expectations should result in systematic differences in risk-taking behaviour. Individuals with high future expectations (who have much to lose) should be more risk-averse than individuals with low expectations. This applies to all kinds of risky situations, so individuals should consistently differ in their behaviour. By means of an evolutionary model we demonstrate that this basic principle results in the evolution of animal personalities. It simultaneously explains the coexistence of behavioural types, the consistency of behaviour through time and the structure of behavioural correlations across contexts. Moreover, it explains the common finding that explorative behaviour and risk-related traits like boldness and aggressiveness are common characteristics of animal personalities.

Abstract: Living in large, stable social groups is often considered to favour the evolution of enhanced cognitive abilities, such as recognizing group members, tracking their social status and inferring relationships among them. An individual's place in the social order can be learned through direct interactions with others, but conflicts can be time-consuming and even injurious. Because the number of possible pairwise interactions increases rapidly with group size, members of large social groups will benefit if they can make judgments about relationships on the basis of indirect evidence. Transitive reasoning should therefore be particularly important for social individuals, allowing assessment of relationships from observations of interactions among others. Although a variety of studies have suggested that transitive inference may be used in social settings, the phenomenon has not been demonstrated under controlled conditions in animals. Here we show that highly social pinyon jays (Gymnorhinus cyanocephalus) draw sophisticated inferences about their own dominance status relative to that of strangers that they have observed interacting with known individuals. These results directly demonstrate that animals use transitive inference in social settings and imply that such cognitive capabilities are widespread among social species.

Abstract: Animals that forage socially often stand to gain from coordination of their behaviour. Yet it is not known how group members reach a consensus on the timing of foraging bouts. Here we demonstrate a simple process by which this may occur. We develop a state-dependent, dynamic game model of foraging by a pair of animals, in which each individual chooses between resting or foraging during a series of consecutive periods, so as to maximize its own individual chances of survival. We find that, if there is an advantage to foraging together, the equilibrium behaviour of both individuals becomes highly synchronized. As a result of this synchronization, differences in the energetic reserves of the two players spontaneously develop, leading them to adopt different behavioural roles. The individual with lower reserves emerges as the 'pace-maker' who determines when the pair should forage, providing a straightforward resolution to the problem of group coordination. Moreover, the strategy that gives rise to this behaviour can be implemented by a simple 'rule of thumb' that requires no detailed knowledge of the state of other individuals.