Habitat generalist species constrain the diversity of mimicry rings in heterogeneous habitats.

How evolution creates and maintains trait patterns in species-rich communities is still an unsolved topic in evolutionary ecology. One classical example of community-level pattern is the unexpected coexistence of different mimicry rings, each of which is a group of mimetic species with the same warning signal. The coexistence of different mimicry rings in a community seems paradoxical because selection among unpalatable species should favor convergence to a single warning pattern. We combined mathematical modeling based on network theory and numerical simulations to explore how different types of selection, such as mimetic and environmental selections, and habitat use by mimetic species influence the formation of coexisting rings. We show that when habitat and mimicry are strong sources of selection, the formation of multiple rings takes longer due to conflicting selective pressures. Moreover, habitat generalist species decrease the distinctiveness of different mimicry rings' patterns and a few habitat generalist species can generate a "small-world effect", preventing the formation of multiple mimicry rings. These results may explain why the coexistence of mimicry rings is more common in groups of animals that tend towards habitat specialism, such as butterflies.