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Many biological and social systems show significant levels of collectiveaction. Several cooperation mechanisms have been proposed, yet they have beenmostly studied independently. Among these, direct reciprocity supportscooperation on the basis of repeated interactions among individuals. Signalsand quorum dynamics may also drive cooperation. Here, we resort to anevolutionary game theoretical model to jointly analyse these two mechanisms andstudy the conditions in which evolution selects for direct reciprocity,signalling, or their combination. We show that signalling alone leads to higherlevels of cooperation than when combined with reciprocity, while offeringadditional robustness against errors. Specifically, successful strategies inthe realm of direct reciprocity are often not selected in the presence ofsignalling, and memory of past interactions is only exploited opportunisticallyin case of earlier coordination failure. Differently, signalling alwaysevolves, even when costly. In the light of these results, it may be easier tounderstand why direct reciprocity has been observed only in a limited number ofcases among non-humans, whereas signalling is widespread at all levels ofcomplexity.

Signalling boosts the evolution of cooperation in repeated group interactions