Nutrition induced direct fitness for workers in a primitively eusocial wasp

Eusocial insects are characterised by a reproductive caste differentiation such that egg-laying is restricted to a small number of queens. The majority of the colony members function as non-reproducing workers and gain indirect fitness by rearing the offspring of the reproductives. In primitively eusocial species, some workers can also get direct fitness by sneaking in some eggs in the presence of the queens, replacing the queens and becoming new queens, or initiating new nests. Here we aimed to understand the factors that permit some workers to gain direct fitness and alter the number of workers doing so, using the primitively eusocial wasp Ropalidia marginata. We transplanted 12 naturally occurring colonies into large laboratory cages where there was adequate space for the workers to leave their natal nests and initiate new colonies. We compared six control colonies that we provided with ad libitum food placed near the nest to six test colonies in which we hand-fed the wasps in addition to the ad libitum food. Wasps in test colonies consumed more food, showed more aggression, replaced their queens, and initiated new nests significantly more often than those in control colonies. When considering all 12 colonies, the rates of queen replacement and nest foundation were significantly positively correlated with food consumption rates. The additional nutrition gained by hand-fed wasps appears to help workers in test colonies to develop their ovaries and lay eggs, implying that they are nutritionally castrated in control colonies.

Sex Ratio Is Synergistically Modulated by Local Resource Competition and Enhancement: the Case of a Primitively Eusocial Wasp Under Female Philopatry

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.

The effect of age on non-reproductive division of labour in the tropical primitively eusocial wasp, Ropalidia cyathiformis

Division of labour among workers (non-reproductive division of labour), a characteristic feature of eusocial insects enables the efficient functioning of their colonies. In many advanced insect societies division of labour is based on age (age polyethism). Primitively eusocial insects however are believed to have a weak age polyethism. Here we investigated the role of age in non-reproductive division of labour in the tropical primitively eusocial wasp, Ropalidia cyathiformis and compared it with that in Ropalidia marginata, a congeneric species that exhibits relatively strong age polyethism. Age had a significant effect on the first performance of the four tasks studied; tasks were initiated in the sequence feed larva, build, bring food and bring building material. We measured task performance as the absolute frequency of tasks performed (FTP) and the probability of performing a task relative to other tasks (PTP) and age as absolute age in days since eclosion as well as relative age compared to nestmates. FTP varied significantly with both absolute and relative age, although absolute age explained more variance. PTP varied significantly with absolute age but not always with relative age. This is contrary to R. marginata, where more variation is explained by relative age than by absolute age. There was no trade-off between intranidal and extranidal tasks in R. cyathiformis unlike in R. marginata where the frequency of intranidal tasks decreased and that of extranidal tasks increased with age. We conclude that age polyethism is weak and less flexible in R. cyathiformis compared to that in R. marginata.

A place for everything and everything in its place: spatial organization of individuals on nests of the primitively eusocial wasp Ropalidia marginata

Non-random space use is common among animals across taxa and habitats. Social insects often use space non-randomly, outside as well as inside their nests. While such non-random space use outside the nest may improve foraging efficiency, inside the nest, it is often associated with the efficient division of labour. Non-random space use by adults on their nests has been hypothesized to result from dyadic dominance interactions, non-random distribution of tasks, differential activity levels, workers avoiding their queens or prophylactic avoidance of disease spread. These hypotheses are generally derived from species in which the tasks of the workers are themselves non-randomly distributed on the nest. Here, we study the primitively eusocial wasp Ropalidia marginata , in which tasks are not distributed non-randomly, and show that 62.4% ± 16.2% of the adults nevertheless use space on their nest non-randomly. In this species, we find that non-random space use may help optimizing nutritional exchange between individuals while prophylactically minimizing disease spread among nest-mates. We did not find evidence for the roles of dominance interactions, activity levels or location of larvae in non-random space use. Spatial organization appears to be a mechanism of minimizing the costs and maximizing the benefits of social life.