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Results: The results from phylogenetic analyses reveal that the inquilinism (or social parasite) evolved once in the clade of yellowjackets and hornets. The divergence of inquilines from their hosts is estimated to have occurred 2.18 (sd = 0.52) million years ago.

Methods: A total of 12 gene fragments were sequenced from 12 species of yellowjackets, Vespula*, and hornets, Vespa*. A Bayesian phylogenetic analysis, using the molecular data, was performed with MrBayes. Dating analyses of the phylogeny were conducted with BEAST. The results were mapped on the species tree by the DIVALIKE program. Bayesian molecular dating was used to estimate the divergence dates of inquilinism among the studied species. The rate of synonymous substitution was used as the molecular clock.

Abstract: Social parasites exploit the brood-care behaviour and social structure of one or more host species. Within the social Hymenoptera there are different types of social parasitism. In its extreme form, species of obligate social parasites, or inquilines, do not have the worker caste and depend entirely on the workers of a host species to raise their reproductive offspring. The strict form of Emery's rule states that social parasites share immediate common ancestry with their hosts. Moreover, this rule has been linked with a sympatric origin of inquilines from their hosts. Here, we conduct phylogenetic analyses of yellowjackets and hornets based on 12 gene fragments and evaluate competing evolutionary scenarios to test Emery's rule.

In addition to the identification of the closest relatives of social parasites, we discuss and reject other hypotheses that may explain why this group of organisms has evolved to live in social colonies. We conclude that the evolution of inquilinism in yellowjackets and hornets is due to species-specific social and behavioral traits and not to phylogenetic constraints. This finding also demonstrates that the evolution of sociality in social insects is not a consequence of ecological conditions, but rather a response to selection for other important evolutionary innovations and conditions. The evolution of inquilines in social Hymenoptera is thus a fascinating example of the interplay between ecological, behavioral and social traits.

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