What can mechanisms underlying derived traits tell us about the evolution of social behavior?

Social behavior, although rare, is a highly successful form of living that has reached its most extreme forms in eusocial insects. A tractable framework to understand social evolution is the study of major transitions in social behavior. This includes the transitions between solitary to social living, from species exhibiting intermediate degrees of sociality to species exhibiting true sociality, and from primitive to advanced eusocial species. The latter transition is characterized by the emergence of traits not previously found in primitive eusocial species, such as fixed morphological differences between castes and task specialization within the sterile caste. Such derived traits appear to exist in a binary fashion, present in advanced eusocial species, and absent or rare in primitive ones, and thus do not exist in a gradient that is easily tracked and compared between species. Thus, they may not be viewed as valuable to explore ultimate questions related to social evolution. Here, we argue that derived traits can provide useful insights on social evolution even if they are absent or rare in species with a lower social organization. This applies only if the mechanism underlying the trait, rather than the function it regulates for, can be traced back to the solitary ancestors. We discuss two examples of derived traits, morphological differences in female castes and primer pheromones regulating female reproduction, demonstrating how their underlying mechanisms can be used to understand major transitions in the evolution of social behavior and emphasize the importance of studying mechanistic, rather than functional continuity of traits.