The advanced colonial state of eusociality has evolved in insects as a defense of nest sites within foraging distance of persistent food sources. In the Hymenoptera, the final step in the approach to eusociality is through a suite of preadaptations comprising simultaneous provisioning, fidelity to the nest, and a preexisting propensity toward dominance behavior and the selection of tasks according to opportunity. The only genetic change needed to cross the threshold to the eusocial grade is the foundress’s possession of an allele that holds the foundress and her offspring to the nest. The preadaptations provide the phenotypic flexibility required for eusociality, as well as the key emergent traits arising from interactions of the group members. Group (colony-level) selection then immediately acts on both of these traits. The rarity of the origin of eusociality is evidently due to the rarity of the combination of progressive provisioning with environments of the kind that gives an edge to group selection over the individual direct selection, causing offspring to stay at the natal nest rather than disperse. Several lines of evidence, examined here, suggest that collateral kin selection does not play a significant role.
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