Sex ratio theory in social insects with swarming

Abstract

In social insects which reproduce by colony fission there is often only one queen in each swarm (e.g. honeybees, army ants) and the number of males greatly exceeds the number of new queens produced at swarming time. This seems to contradict Fisher's principle that there should be equal investment in male and female reproductives. Hamilton 1975 has suggested that the principle can be saved by counting the investment in the swarm as part of the investment in female reproductives. Craig 1980, on the other hand, argues that few queens are produced because any further investment in queens would be wasted since a queen without a swarm is valueless; on this view the sex ratio is male-biased because of local competition between queens for swarms. The present paper investigates from first principles how many males should be produced by a species which reproduces by colony fission and how the workers should divide themselves between the new colonies. The results of the analysis do not support Hamilton's conjecture but show that a number of factors are involved in the evolution of male production, including kin selection and the relationship between the “fitness” of a colony and the number of workers in it.