Origin of an evolutionary novelty: the worker phenotype of eusocial wasps
AbstractComplex problems in evolutionary biology can be approached in two ways, top down using theoretical constructs and bottom up using empirical studies . Theoretical concepts predominate evolutionary interpretations of eusociality in a literature that is small relative to an enormous literature of natural history and basic research that is not synthesized into a conceptual whole. Here, I draw insights from this literature to show how paper wasps’ allomaternal non-reproductive worker phenotype originates in every colony cycle via confluence of multiple factors of paper wasp biology. These include behavior, development, nutrient dynamics, indirect genetic effects, sex ratio, and demography. A novel perspective on the colony cycle, based on individuals’ reproductive physiology, serves as context to examine of each of these. It will be shown that the allomaternal non-reproductive worker phenotype does not require relatedness among colony members to originate. Allomaternal care of non-relatives is frequent and can occur in at least twelve contexts. Life histories of living species as they will be presented here show that relatedness among colony members is not the target of selection in simple eusociality. However, the novel allomaternal non-reproductive worker phenotype had to be present at the ancestral origins of complex eusociality in which relatedness among colony members is essential.