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One of the hallmarks of eusociality is that workers forego their own reproduction to assist their mother in raising siblings. This seemingly altruistic behaviour may benefit workers if gains in indirect fitness from rearing siblings outweigh the loss of direct fitness. If worker presence is advantageous to mothers, however, eusociality may evolve without net benefits to workers. Indirect fitness benefits are often cited as evidence for the importance of inclusive fitness in eusociality, but have rarely been measured in natural populations. We compared inclusive fitness of alternative social strategies in the tropical sweat bee, Megalopta genalis, for which eusociality is optional. Our results show that workers have significantly lower inclusive fitness than females that found their own nests. In mathematical simulations based on M. genalis field data, eusociality cannot evolve with reduced intra-nest relatedness. The simulated distribution of alternative social strategies matched observed distributions of M. genalis social strategies when helping behaviour was simulated as the result of maternal manipulation, but not as worker altruism. Thus, eusociality in M. genalis is best explained through kin selection, but the underlying mechanism is likely maternal manipulation., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

Social castes of eusocial insects may have arisen through an evolutionary modification of an ancestral reproductive ground plan, such that some adults emerge from development physiologically primed to specialize on reproduction (queens) and others on maternal care expressed as allo-maternal behaviour (workers). This hypothesis predicts that variation in reproductive physiology should emerge from ontogeny and underlie division of labour. To test these predictions, we identified physiological links to division of labour in a facultatively eusocial sweat bee, Megalopta genalis. Queens are larger, have larger ovaries and have higher vitellogenin titres than workers. We then compared queens and workers with their solitary counterparts-solitary reproductive females and dispersing nest foundresses-to investigate physiological variation as a factor in caste evolution. Within dyads, body size and ovary development were the best predictors of behavioural class. Queens and dispersers are larger, with larger ovaries than their solitary counterparts. Finally, we raised bees in social isolation to investigate the influence of ontogeny on physiological variation. Body size and ovary development among isolated females were highly variable, and linked to differences in vitellogenin titres. As these are key physiological predictors of social caste, our results provide evidence for developmental caste-biasing in a facultatively eusocial bee.

Optimal-skew models (OSMs) predict that cooperative breeding occurs as a result of dominants conceding reproductive benefits to subordinates, and that division of reproduction within groups reflects each cooperator's willingness and ability to contest aggressively for dominance. Polistine paper wasps are a leading model system for testing OSMs, and data on reproduction and aggression appear to support OSMs. These studies, however, measure aggression as a single rate rather than by the activity patterns of individuals. This leads to a potential error: if individuals are more likely to receive aggression when active than when inactive, differences in aggression across samples can reflect changes in activity rather than hostility. This study replicates a field manipulation cited as strongly supporting OSMs. We show that fundamentally different conclusions arise when controlling for individual activity states. Our analyses strongly suggest that behaviours classified as 'aggression' in paper wasps are unlikely to function in establishing, maintaining or responding to changes in reproductive skew. This illustrates that OSM tests using aggression or other non-reproductive behaviour as a metric for reproductive partitioning must demonstrate those links rather than assume them.

Recent evolutionary models of reproductive partitioning within animal societies (known as 'optimal skew', 'concessions' or 'transactional' models) predict that a dominant individual will often yield some fraction of the group's reproduction to a subordinate as an incentive to stay in the group and help rear the dominant's offspring. These models quantitatively predict how the magnitude of the subordinate's 'staying incentive' will vary with the genetic relatedness between dominant and subordinate, the overall expected group output and the subordinate's expected output if it breeds solitarily. We report that these predictions accord remarkably well with the observed reproductive partitioning between conesting dominant and subordinate queens in the social paper wasp Polistes fuscatus. In particular, the theory correctly predicts that (i) the dominant's share of reproduction, i.e. the skew, increases as the colony cycle progresses and (ii) the skew is positively associated both with the colony's productivity and with the relatedness between dominant and subordinate. Moreover, aggression between foundresses positively correlated with the skew, as predicted by transactional but not alternative tug-of-war models of societal evolution. Thus, our results provide the strongest (quantitative support yet for a unifying model of social evolution.