Abstract: Cooperation among unrelated individuals can evolve not only via reciprocal altruism but also via trait-group selection or by-product mutualism (or some combination of all three categories). Therefore the (iterated) prisoner's dilemma is an insufficient paradigm for studying the evolution of cooperation. We replace this game by the cooperator's dilemma, which is more versatile because it enables all three categories of cooperative behavior to be examined within the framework of a single theory. Controlled studies of cooperation among fish provide examples of each category of cooperation. Specifically, we describe reciprocal altruism among simultaneous hermaphrodites that swap egg parcels, group-selected cooperation among fish that inspect dangerous predators and by-product mutualism in the cooperative foraging of coral-reef fish.

Abstract: An altruistic individual has to gamble on cooperation to a stranger because it does not know whether the stranger is trustworthy before direct interaction. Nowak and Sigmund (Nature 393 (1998a) 573; J. Theor. Biol. 194 (1998b) 561) presented a new theoretical framework of indirect reciprocal altruism by image scoring game where all individuals are informed about a partner's behavior from its image score without direct interaction. Interestingly, in a simplified version of the image scoring game, the evolutionarily stability condition for altruism became a similar form of Hamilton's rule, i.e. inequality that the probability of getting correct information is more than the ratio of cost to benefit. Since the Hamilton's rule was derived by evolutionarily stable analysis, the evolutionary meaning of the probability of getting correct information has not been clearly examined in terms of kin and group selection. In this study, we applied covariance analysis to the two-score model for deriving the Hamilton's rule. We confirmed that the probability of getting correct information was proportional to the bias of altruistic interactions caused by using information about a partner's image score. The Hamilton's rule was dependent on the number of game bouts even though the information reduced the risk of cooperation to selfish one at the first encounter. In addition, we incorporated group structure to the two-score model to examine whether the probability of getting correct information affect selection for altruism by group selection. We calculated a Hamilton's rule of group selection by contextual analysis. Group selection is very effective when either the probability of getting correct information or that of future interaction, or both are low. The two Hamilton's rules derived by covariance and contextual analyses demonstrated the effects of information and group structure on the evolution of altruism. We inferred that information about a partner's behavior and group structure can produce flexible pathways for the evolution of altruism.

Abstract: Mule deer, Odocoileus hemionus, females actively defend fawns against predators, including nonoffspring conspecific fawns and heterospecific white-tailed deer, O. virginianus, fawns. We hypothesized that the defence of nonoffspring fawns was due to a recognition error. During a predator attack, females may have to decide whether to defend a fawn with imperfect information on its identity obtained from hearing only a few distress calls. We examined fawn distress calls to determine whether calls made by the two species and by different individuals within each species were acoustically distinctive. The mean and maximum fundamental frequencies of mule deer fawns were nearly double those of white-tailed deer fawns, with no overlap, enabling us to classify 100% of calls to the correct species using a single trait. A large proportion of calls was also assigned to the correct individual using a multivariate analysis (66% and 70% of mule deer and white-tailed deer fawns, respectively, chance = 6% and 10%); however, there was considerable statistical uncertainty in the probability of correct classification. We observed fawns approach conspecific females in an attempt to nurse; females probed most offspring fawns with their noses before accepting them, and always probed nonoffspring fawns before rejecting them, suggesting that close contact and olfactory information were needed to unequivocally distinguish nonoffspring from offspring fawns. Taken together, these results suggest that acoustic variation alone would probably be sufficient to permit rapid and reliable species discrimination, but it may not be sufficient for mothers to unequivocally distinguish their own fawn from conspecific fawns.

Abstract: Recent experimental research has revealed forms of human behavior involving interaction among unrelated individuals that have proven difficult to explain in terms of kin or reciprocal altruism. One such trait, strong reciprocity is a predisposition to cooperate with others and to punish those who violate the norms of cooperation, at personal cost, even when it is implausible to expect that these costs will be repaid. We present evidence supporting strong reciprocity as a schema for predicting and understanding altruism in humans. We show that under conditions plausibly characteristic of the early stages of human evolution, a small number of strong reciprocators could invade a population of self-regarding types, and strong reciprocity is an evolutionary stable strategy. Although most of the evidence we report is based on behavioral experiments, the same behaviors are regularly described in everyday life, for example, in wage setting by firms, tax compliance, and cooperation in the protection of local environmental public goods.