Sex ratio and sexual conflict in a collective action problem

The maintenance of cooperation is difficult whenever collective action problems are vulnerable to freeriding (reaping the benefits without contributing to the maintenance of the good). We identify a novel factor that can make a system tolerate an extent of freeriding. If a population consists of discrete types with demographically distinct roles, such that the success of one type does not imply it can spread to replace other types in the population, then collective goods may persist in the presence of free-riders because they are necessarily kept in a minority role. Biased sex ratios (e.g. in haplodiploids) create conditions where individuals of one sex are a minority. We show that this can make the less common sex contribute less to a public good in a setting where the relevant life-history stage -- larval group defence against predators -- does not feature any current breeding opportunities that might lead to confounding reasons behind sex-specific behaviour. We test our model with haplodiploid pine sawfly larvae, showing that female larvae are the main contributors to building the antipredator defence against predators. Significance statementIndividuals in groups can cooperate to achieve something together, but with an evolutionary difficulty: if benefits of cooperation are shared equally among all, freeriders get the same benefit as others while paying less for it. We propose a novel reason why freeriding does not automatically spread until the collectively beneficial outcome is destroyed: sometimes groups consist of individuals of distinct categories, limiting freerider spread. If, for example, there are always fewer males than females, then even if every male becomes a freerider, the whole group still survives simply because not everyone can be male. Pine sawfly larvae defend against predators by regurgitating sticky fluids, but females contribute more to this common defence, and we show this example fits our model.