The ecology and evolution of male reproductive behaviour in the bees and wasps

Males of aculeate Hymenoptera differ in the behavioural adaptations employed to locate and secure mates. The ecological and evolutionary bases of these differences are explored in this paper. Male bees and wasps search for females by patrolling widely within emergence-nesting areas or within patches of flowers attractive to conspecific females, or by waiting at landmarks, at specific emergence sites, or at nests. Nest dispersion, flower distribution, the type of female mating system and the nature of male-male competition appear to be key factors in determining the mate-locating behaviour of males. Of special interest in multiple-mating by females, which may be an evolutionary response to the costs of attempting to resist copulation in certain situations. When polyandry occurs, males are under selection pressure to be the last male to copulate with a female prior to oviposition if sperm precedence occurs. In species in which females mate just once, a selective premium is placed on being the first male to reach a virgin female. In either case, because receptive females are a limited resource, there is intense competition among males for access to the resource. The density of competitor males may play an important role in determining whether holding a relatively restricted territory is preferable to the strategy of patrolling widely at various sites which may have females. Territoriality is practiced by males of several species of aculeate Hymenoptera when the number of male competitors is relatively few in number and the distribution of emergence sites or foraging areas of females is clumped in space.