When to bee social: interactions among environmental constraints, incentives, guarding, and relatedness in a facultatively social carpenter bee

In the facultatively social carpenter bee, Xylocopa pubescens,foundresses usually establish nests solitarily. However, nestsmay become social if a second foundress (referred to as alpha)successfully usurps the nest, with the original foundress (referredto as beta) remaining as a guard. Reproductive skew theory predictsthat beta foundresses should remain as helpers only if alphausurpers allow them a share of reproduction. Because alpha femalesdestroy much of beta's brood and beta females do not lay eggsafter takeovers, studies have concluded that usurpers offerno staying incentives or concessions in return for helping behavior.This conclusion is paradoxical, and we suggest that by refrainingfrom destroying all of beta's brood, alpha females do indeedoffer concessions to beta females. We constructed a model toexamine the conditions under which social nesting is favoredby both alpha and beta females. Female preference for socialversus solitary nesting is proportional to expected fitnessin either setting and is affected by current environmental conditions,the value of guarding behavior in protecting brood from pollenrobbery, the size of the concession offered by alpha, and thedegree of genetic relatedness between the foundresses. Our modelshows that at a minimum, establishing sociality after unrelatedusurpations always requires a concession, whereas in relatedusurpations, a concession is not always required. Generally,agreement between alpha and beta is difficult because alpharequires a much higher premium in pollen robbery protectionthan beta in order for sociality to be advantageous. Alpha femalesprefer social nesting only under the most severe environmentalconditions because usually they gain less by the presence ofa guard than by replacing beta brood with their own. In contrast,beta females always strongly prefer social nesting because thechances of successful renesting elsewhere are low and rarelyoutweigh the advantages of guarding their own brood that surviveusurpation. Effects of relatedness between foundresses on preferencefor social nesting are nonintuitive: first, alpha's preferenceincreases with relatedness, whereas beta's preference declines,and second, unrelated beta females prefer sociality more stronglythan related ones. This is because replacement of beta's offspringwith related alpha offspring partially compensates her for theloss of her own brood, even should she leave the nest.