The evolution and function of pattern diversity in snakes

Species in the suborder Serpentes present a powerful model for understanding processes involved in visual signal design. Although vision is generally poor in snakes, they are often both predators and prey of visually oriented species. We examined how ecological and behavioral factors have driven the evolution of snake patterning using a phylogenetic comparative approach. The appearances of 171 species of Australian and North American snakes were classified using a reaction-diffusion model of pattern development, the parameters of which allow parametric quantification of various aspects of coloration. The main findings include associations between plain color and an active hunting strategy, longitudinal stripes and rapid escape speed, blotched patterns with ambush hunting, slow movement and pungent cloacal defense, and spotted patterns with close proximity to cover. Expected associations between bright colors, aggressive behavior, and venom potency were not observed. The mechanisms through which plain and longitudinally striped patterns might support camouflage during movement are discussed. The flicker-fusion hypothesis for transverse striped patterns being perceived as uniform color during movement is evaluated as theoretically possible but unlikely. Snake pattern evolution is generally phylogenetically conservative, but by sampling densely in a wide variety of snake lineages, we have demonstrated that similar pattern phenotypes have evolved repeatedly in response to similar ecological demands.