A central issue in evolutionary biology is the extent to which complex social organization is under genetic control. We have found that a single genomic element marked by the protein-encoding gene Gp-9 is responsible for the existence of two distinct forms of social organization in the fire ant Solenopsis invicta. This genetic factor inf luences the reproductive phenotypes and behavioral strategies of queens and determines whether workers tolerate a single fertile queen or multiple queens per colony. Furthermore, this factor affects worker tolerance of queens with alternate genotypes, thus explaining the dramatic differences in Gp-9 allele frequencies observed between the two social forms in the wild. These findings reveal how a single genetic factor can have major effects on complex social behavior and inf luence the nature of social organization.Organic evolution is marked by a small number of major transitions, one of which is the evolution of complex social behavior (1, 2). Social life can take a variety of forms, each distinguished by features such as group size and the reproductive roles of group members (3,(4)(5)(6)(7)(8). A critical need in evolutionary biology is to identify the causes of social behavior and its conspicuous variation and especially to determine the extent to which social organization is under genetic control (9-15). Such information is crucial for reconstructing pathways of social evolution (16,17) and can lead to improved genetic models for the origin and spread of new social variants (18-20) that may constitute adaptive solutions to changed environments or transitional stages during speciation (3,(21)(22)(23)(24)(25). Current views on insect social evolution stress the importance of ecological and behavioral environments in molding what are largely plastic social behaviors (3,17,21,22,(25)(26)(27)(28)(29)(30). As a result of this emphasis on behavioral plasticity and an immediate role of the environment, studies of the genetic underpinnings of social behavior have languished. Here we present evidence that major variation in the social organization of fire ant colonies is under simple genetic control, providing a unique demonstration of a strong genetic component to complex social behavior.The fire ant Solenopsis invicta is an introduced pest species in the southern United States that exists in two distinct social forms. The monogyne form features colonies with a single fertile (egg-laying) queen, whereas the polygyne form features colonies with multiple fertile queens. The two social forms differ in other major aspects of their reproductive biology, including the phenotypes and reproductive strategies of young queens and the modes of colony founding (3). Monogyne nests produce heavy queens that accumulate large fat reserves and exhibit rapid oogenesis, whereas polygyne nests produce mostly lighter queens with lesser reserves and more gradual oogenesis (31). These phenotypic differences are adaptive because young monogyne queens found colonies independently (without workers), a...