Territorial-sneaker games with non-uniform interactions and female mate choice

Male territorial-sneaker polymorphisms are common in nature. To understand how these polymorphisms evolve, we developed a game theoretical model analogous to the classical Hawk-Dove model, but with two important differences. First, we allowed non-uniform interaction rates of strategies to account for the possibility that some interactions between male strategies are disproportionately more frequent than others. Second, we allowed females to exhibit a preference for one type of male and thereby choose mates adaptively. Selection dynamics were modeled using coupled replicator equations. The model confirms that there is a broad range of conditions under which a male polymorphism will arise. We applied the model to understand the genetic polymorphism in adult male Mnais damselflies (Zygoptera). Here, orange-winged adult males defend oviposition sites and mate with females when they arrive, while clear-winged ‘sneaker’ males are typically non-territorial and opportunistically mate with females. Intriguingly, in allopatry, the males of Mnais costalis and M. pruinosa both exhibit the same orange-clear winged polymorphism but where the species co-occur, males of M. costalis evolve orange wings while males of M. pruinosa tend to evolve clear wings. To understand this phenomenon and evaluate the importance of female choice in mediating it, we extended our game-theoretical model to two interacting species. While both competitive and reproductive interference can explain the male monomorphisms in sympatry, reproductive interference explains the phenomenon under a wider set of conditions. When females of the rarer species change their male preferences to facilitate species discrimination, it can generate runaway selection on male phenotypes.