Abstract

Ever since the seminal papers of Hamilton (1964), a common focus of sociobiology has been the genetic relationships among individuals of insect societies. At issue are agents that reduce the average relatedness among colony members since they are difficult to interpret in light of the evolution of sociality by kin selection. One such factor is polyandry or female multiple mating. Low levels of polyandry are common among the social Hymenoptera, occurring across a broad range of taxa and social systems (Page and Metcalf 1982; Page 1986; Crozier and Pamilo 1996). In spite of that, high mating frequencies (‚2 mates/ female) are relatively rare among the highly eusocial Hymenoptera, limited to only one or a few genera in each of the ants, bees, and wasps (Boomsma and Ratnieks 1996). These extraordinary levels of polyandry have been of particular interest among students of social insect research since their adaptive significance is still in question and highly debated (Kraus and Page 1998; Sherman et al. 1998). The genus Apis has been studied in depth for both queen reproductive behavior and mating frequency, facts that allow insight into the evolution of extreme polyandry in these species. The reproductive behavior of honey bee queens (Apis mellifera) occurs early in their lifetimes. Virgin queens initiate mating behavior when they are about 1 wk old by briefly exiting the colony and orienting to local landmarks (Ruttner 1956). After such orientation * To whom correspondence should be addressed; e-mail: drtarpy@

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