Sex Ratio Is Synergistically Modulated by Local Resource Competition and Enhancement: the Case of a Primitively Eusocial Wasp Under Female Philopatry
Abstract BackgroundIn animals, the offspring sex ratio is modulated by kin conflict and cooperation, and determining the ratio is a main concern in evolutionary biology. Male competition for access to local mates is predictive of a female-biased sex ratio in the offspring (local mate competition; LMC). Conversely, female competition for access to local resources is predictive of a male-biased sex ratio in the offspring (local resource competition; LRC). However, several factors other than competition should synergistically operate in real-world populations. In the Australian paper wasp Ropalidia plebeiana, LRC and local resource enhancement (LRE) may operate simultaneously. To determine whether this is the case, we evaluated colony sex ratios and examined whether competition and/or enhancement operates at the population level in this species. ResultsIn spring, many foundress queens started their colonies by comb-cutting, in which nest combs from the previous season were divided into several combs to be reused. Genetic relatedness among foundresses did not differ before and after comb-cutting. Relatedness among foundresses was 0.339, whereas relatedness among new foundresses was 0.589, revealing nearly functional monogyny. The global FST value calculated with mtDNA markers was higher than that calculated with microsatellite markers, even after we corrected for differences in effective population sizes between sexes. This finding indicates female philopatry, which was also confirmed by mark–release–recapture before and after the hibernation of new foundresses. The colony sex ratio of reproductives became slightly biased toward males in larger colonies. In addition, both the number of foundresses and number of workers were positively associated with the number of reproductives, which indicates that LRE was also operating.ConclusionsOur results suggest that although the population structure seems to meet the requirements of LRC, the sex ratio is not modulated solely by LRC. Instead, the availability of female helpers at the founding stage likely mitigates the sex ratio predicted by LRC through LRE. Thus, LRC at the founding stage and LRE at the reproductive stage synergistically modulate the colony sex ratio in R. plebeiana.