Species and morphological diversification in snakes of the family Viperidae: patterns and processes

Species and phenotypic diversity may vary considerably between taxonomic groups and through time for a given lineage. The study of such variation became one of the main goals of evolutionary biology and provides important information related to the possible mechanisms regulating biodiversity. The general goal of the present thesis was to investigate the patterns of species and morphological diversification in a cosmopolitan group of snakes, the family Viperidae, and the potential underlying processes. First, (1) we estimated the phylogenetic relationships and divergence times between lineages of the family Viperidae using a Bayesian approach; then we (2) applyed a recently developed method (BAMM) to explore how speciation and extinction rates varied during the radiation of the group suggesting possible underlying processes. Finally, (3) we analyzed if body size evolution and speciation rates showed distinct patterns among vipers occurring in different habitats (terrestrial vs arboreal). Herein we generated the most complete molecular phylogeny for vipers until this moment using sequences from 11 mitochondrial and nuclear genes comprising 79% of extant species (264 terminals) and all except one genus. In general, we were able to recover well supported phylogenetic relationships with most genera being monophyletic. Divergence time estimates suggested that vipers started to diversify around the late Paleocene/middle Eocene finding older ages than previous studies. During the group radiation, an increase in speciation rates seems to have occurred during the diversification of crotalines (pit vipers) not only due to the evolution of loreal pits but also as a result of climatic and geological changes in Asia and the invasion of the New World. After this rapid initial increase, speciation rates decelerated toward the present. Lastly, the results presented here suggest that although arboreal habitats constrain morphological evolution in vipers the evolution of arboreality does not seem to affect speciation rates, which remain similar among arboreal and terrestrial lineages. Our results suggest two distinct scenarios: (1) speciation could be independent of morphological evolution in vipers; or (2) geographic isolation would be an important mechanism underlying species diversification in arboreal lineages offsetting decreases in speciation opportunities potentially related to the selective pressures imposed by the arboreal environment. The present thesis contribute to increase our understanding about how vipers evolved during their ∼50 million years. In addition to providing scenarios and hypotheses to be further explored with vipers, we elaborated a broad and conceptual discussion about the possible mechanisms underlying species and morphological diversification that might apply to other groups of organisms. Therefore, this thesis comprises a contribution that goes beyond the understanding of mechanisms generating and maintaining the diversity of snakes, but will hopefuly enrich the discussion of mechanisms that generate and maintain biodiversity as a whole (AU)