Warning Signals Promote Morphological Diversification In Fossorial Uropeltid Snakes (Squamata: Uropeltidae)

Many species possess warning colourations that signal unprofitability to predators. Warning colourations are also thought to provide prey with a 'predator-free space' and promote niche expansion. However, how such strategies release a species from environmental constraints and facilitate niche expansion is not clearly understood. Fossoriality in reptiles imposes several morphological limits on head and body size to facilitate burrowing underground, but many fossorial snakes live close to the surface and occasionally move above ground, exposing them to predators. In such cases, evolving antipredator defences that reduce predation on the surface could potentially relax the morphological constraints associated with fossoriality and promote morphological diversification. Fossorial uropeltid snakes possess varying degrees of conspicuous warning colourations that reduce avian predation when active above ground. We predicted that species with more conspicuous colourations will exhibit more robust body forms and show faster rates of morphological evolution because constraints imposed by fossoriality are relaxed. Using a comparative phylogenetic approach on the genus Uropeltis, we show that more conspicuous species tend to have more robust morphologies and have faster rates of head-shape evolution. Overall, we find that the evolution of warning colourations in Uropeltis can facilitate niche expansion by influencing rates of morphological diversification.