Convergent evolution of wingless reproductives across all subfamilies of ants, and sporadic loss of winged queens (Hymenoptera: Formicidae)

Flight is a one-off event in ants, hence after mating, the wing muscles of winged queens can function as protein reserves during independent colony foundation (ICF). Another strategy occurring in many unrelated lineages is dependent colony foundation (DCF). DCF does not require queens with expensive wing muscles because dispersal is on foot, and a found- ress relies on nestmate workers to feed her first brood of workers. The shift to DCF seems the reason why wingless reproductives (ergatoid queens, short-winged queens, and gamergates) evolved independently in more than 50 genera belonging to 16 subfamilies. In various species they occur together with winged queens (in the same or different popu- lations), in other species winged queens were replaced completely. Because wingless reproductives are the product of convergence, there is tremendous heterogeneity in morphological characteristics as well as selective contexts. These novel reproductive phenotypes cannot function without nestmate workers (foundresses forage in only few species), hence addi- tional investment in workers is needed.