Male fighting and "territoriality" within colonies of the ant Cardiocondyla venustula

*[The ant genus Cardiocondyla is characterized by a bizarre male polymorphism with wingless fighter males & winged disperser males. Winged males have been lost convergently in several clades, & in at least one of them, wingless males have evolved mutual tolerance. To better understand the evolutionary pathways of reproductive tactics, we investigated Cardiocondyla venustula, a species, which in a phylogenetic analysis clusters with species with fighting & species with mutually tolerant, wingless males. Wingless males of C. venustula use their strong mandibles to kill freshly eclosed rival males & also engage in short fights with other adult males, but in addition show a novel behavior hitherto not reported from social insect males: they spread out in the natal nest & defend territories against other males. Ant males therefore show a much larger variety of reproductive tactics than previously assumed.]