Eusociality is not a major evolutionary transition, and why that matters

Major evolutionary transitions (METs) across Earth's biological history describe fusions of lower-level entities into higher-level individuals (evolutionary transitions in individuality: ETIs) as well as novel forms of information storage and transmission (Information Leaps). Obligate eusociality is frequently listed as a MET-most often in the context of being an ETI and with an extrapolation that the ETI requires inclusive fitness maximization for all parties. However, obligate eusociality neither fundamentally alters how information is stored and transmitted nor meets the various criteria proposed for an ETI. We argue that rather than representing a higher-level individual, the evolution of non-reproductive worker castes is more analogous to a novel 'organ' that maintains homeostasis and nurtures the gonadal tissue of mated queens. Worker castes benefit queens by performing dangerous but necessary functions such as foraging, while enabling the gamete-producing functions to be kept relatively safe. This is an ecologically successful and significant evolutionary innovation, which can be thought of as a major competitive transition (MCT). In this context, we hypothesize that worker castes are most likely to evolve through parental manipulation. Employing such a MCT perspective generates a broad series of predictions about eusocial life histories.