Group size and direct fitness in social queues.

We explore the effects of group size on the direct reproductive success of subordinate helpers in eusocial animals where only a single, dominant individual reproduces at one time. Helpers can reproduce directly if they inherit dominance, but when dominance is age based, an individual born into a larger group has a longer wait to inherit. We show that this disincentive to help can potentially be offset by increased productivity, increased life span, and insurance-based benefits for helpers if they survive to inherit dominance in larger groups. We analyze a field experiment in which group size was manipulated in the hover wasp Liostenogaster flavolineata. Productivity increased linearly with group size, larger groups were less likely to fail, and dominants in larger groups may have lived longer. Combined with the probability of inheriting dominance, these effects led overall to a negative correlation between group size and expected direct fitness, mainly because group size decreased during our study period, so that helpers could not expect to inherit as large a group as they started queuing in. Our analysis suggests that the relationship between group size and productivity plays a central role in determining the fitness consequences of helping.