Abstract
Cooperation is the backbone of modern human societies, making it a priority to understand how successful cooperation-sustaining mechanisms operate. Cyclic dominance, a non-transitive set-up comprising at least three strategies wherein the first strategy overrules the second, which overrules the third, which, in turn, overrules the first strategy, is known to maintain biodiversity, drive competition between bacterial strains, and preserve cooperation in social dilemmas. Here, we present a novel route to cyclic dominance in voluntary social dilemmas by adding to the traditional mix of cooperators, defectors and loners, a fourth player type, risk-averse hedgers, who enact tit-for-tat upon paying a hedging cost to avoid being exploited. When this cost is sufficiently small, cooperators, defectors and hedgers enter a loop of cyclic dominance that preserves cooperation even under the most adverse conditions. By contrast, when the hedging cost is large, hedgers disappear, consequently reverting to the traditional interplay of cooperators, defectors, and loners. In the interim region of hedging costs, complex evolutionary dynamics ensues, prompting transitions between states with two, three or four competing strategies. Our results thus reveal that voluntary participation is but one pathway to sustained cooperation via cyclic dominance.