Second-order cooperation: Cooperative offspring as a living public good arising from second-order selection on...

Switching rate between cooperating and non-cooperating genotypes is a crucial social evolution factor, often neglected by game theory-inspired theoretical and experimental frameworks. We show that the evolution of alleles increasing the mutation or phenotypic switching rates toward cooperation is in itself a social dilemma. Although cooperative offspring are often unlikely to reproduce, due to high cost of cooperation, they can be seen both as a living public good and a part of the extended parental phenotype. The competition between individuals that generate cooperators and ones that do not is often more relevant than the competition between cooperators and non-cooperators. The dilemma of second-order cooperation we describe relates directly to eusociality, but can be also interpreted as a division of labor or a soma-germline distinction. The results of our simulations shine a new light on what Darwin had already termed a "special difficulty" of evolutionary theory and describe a novel type of cooperation dynamics.