In less than 100 species of ponerine ants, queens no longer exist and have been replaced by mated egg-laying workers. Workers in other subfamilies can lay haploid eggs when queens are removed, but they never reproduce sexually. Ponerine workers are able to mate because they have a spermatheca in most species, foreign males are sexually active near their nests, and their pygidial gland secretions can assume a sexual meaning. Furthermore, ponerine queens are seldom very fecund, and one or several gamergates are able to approximate their egg production. Finally, opportunities for colony fragmentation occur consequent to their life history, and this is a necessary precondition because gamergates cannot start new colonies independently. Many of these characteristics are associated with the limited caste divergence exhibited in this phylogenetically primitive group. Although a few non-ponerine species exhibit some of these preconditions, gamergates have not been found outside the Ponerinae, which alone exhibit the combination of traits leading to queen elimination and worker mating.