Indirect reciprocity: song sparrows distrust aggressive neighbours based on eavesdropping

The evolution of cooperation between unrelated individuals has been a central issue in evolutionary biology. The main problem in most theories of cooperation is how a cooperative player selects individuals to ‘trust’ so that he does not get exploited by noncooperators. While early models emphasized the role of direct experience with individuals in deciding who to trust, more recent work has shown that individuals can eavesdrop on interactions between other individuals to identify cooperators and noncooperators. This second route to cooperation is called indirect reciprocity. In spatially structured populations with repeated interactions between players, both sources of information (direct experience and observed reputation) are readily available. Most models and empirical studies to date, however, have considered indirect reciprocity only in one-shot interactions when direct experience is not available. We examined the role of indirect reciprocity in the maintenance of mutual restraint in aggression (Dear Enemy cooperation) between territorial male song sparrows, Melospiza melodia. We found that territory owners eavesdropped on simulated defections by a neighbour (intrusions onto a third bird’s territory) and subsequently retaliated against these defecting neighbours. Taken together with our previous results, these results suggest that both direct and indirect reciprocity can be at work in repeated-interaction scenarios, and together lead to emergence of cooperation.