A Müllerian mimicry ring in Appalachian millipedes

Few biological phenomena provide such an elegant and straightforward example of evolution by natural selection as color mimicry among unrelated organisms. By mimicking the appearance of a heavily defended aposematic species, members of a second species gain protection from predators and, potentially, enhanced fitness. Mimicking a preexisting warning advertisement is economical because a potentially costly novel one can be avoided; simultaneously, the addition of more aposematic individuals enhances the overall warning effect. The better-known mimetic systems comprise tropical taxa, but here, we show a remarkable example of color mimicry in 7 species of blind, cyanide-generating millipedes endemic to the Appalachian Mountains of temperate North America. Because these millipedes lack eyes, there is no sexual selection or intraspecific signaling for coloration, providing an ideal system for mimicry studies. We document a Müllerian symbiosis where unrelated species vary in color and pattern over geographical space but appear identical where they co-occur. By using spectral color data, estimations of evolutionary history, and detailed field observations of species abundance, we test 4 predictions of Müllerian mimicry theory and begin to unravel the story of an elaborate mimetic diversification in the forests of Appalachia.