Rates of morphological evolution in Captorhinidae: an adaptive radiation of Permian herbivores.

The evolution of herbivory in early tetrapods was crucial in the establishment of terrestrial ecosystems, although it is so far unclear what effect this innovation had on the macro-evolutionary patterns observed within this clade. The clades that entered this under-filled region of ecospace might be expected to have experienced an "adaptive radiation": an increase in rates of morphological evolution and speciation driven by the evolution of a key innovation. However such inferences are often circumstantial, being based on the coincidence of a rate shift with the origin of an evolutionary novelty. The conclusion of an adaptive radiation may be made more robust by examining the pattern of the evolutionary shift; if the evolutionary innovation coincides not only with a shift in rates of morphological evolution, but specifically in the morphological characteristics relevant to the ecological shift of interest, then one may more plausibly infer a causal relationship between the two. Here I examine the impact of diet evolution on rates of morphological change in one of the earliest tetrapod clades to evolve high-fibre herbivory: Captorhinidae. Using a method of calculating heterogeneity in rates of discrete character change across a phylogeny, it is shown that a significant increase in rates of evolution coincides with the transition to herbivory in captorhinids. The herbivorous captorhinids also exhibit greater morphological disparity than their faunivorous relatives, indicating more rapid exploration of new regions of morphospace. As well as an increase in rates of evolution, there is a shift in the regions of the skeleton undergoing the most change; the character changes in the herbivorous lineages are concentrated in the mandible and dentition. The fact that the increase in rates of evolution coincides with increased change in characters relating to food acquisition provides stronger evidence for a causal relationship between the herbivorous diet and the radiation event.