Phylogenetic relationships within Bothrops neuwiedi group (Serpentes, Squamata): geographically highly-structured lineages, evidence of introgressive hybridization and Neogene/Quaternary diversification

Eight current species of snakes of the Bothrops neuwiedi group are widespread in South American open biomes from northeastern Brazil to southeastern Argentina. In this paper, 140 samples from 93 different localities were used to investigate species boundaries and to provide a hypothesis of phylogenetic relationships among the members of this group based on 1122bp of cyt b and ND4 from mitochondrial DNA and also investigate the patterns and processes occurring in the evolutionary history of the group. Combined data recovered the B. neuwiedi group as a highly supported monophyletic group in maximum parsimony, maximum likelihood and Bayesian analyses, as well as four major clades (Northeast I, Northeast II, East-West, West-South) highly-structured geographically. Monophyly was recovered only for B. pubescens. By contrast, B. diporus, B. lutzi, B. erythromelas, B. mattogrossensis, B. neuwiedi, B. marmoratus, and B. pauloensis, as currently defined on the basis of morphology, were polyphyletic. Sympatry, phenotypic intergrades and shared mtDNA haplotypes, mainly between B. marmoratus and B. pauloensis suggest recent introgressive hybridization and the possible occurrence of a narrow hybrid zone in Central Brazil. Our data suggest at least three candidate species: B. neuwiedi from Espinhaço Range, B. mattogrossensis (TM173) from Serra da Borda (MT) and B. diporus (PT3404) from Castro Barros, Argentina. Divergence estimates highlight the importance of Neogene events in the origin of B. neuwiedi group, and the origin of species and diversification of populations of the Neotropical fauna from open biomes during the Quaternary climate fluctuations. Data reported here represent a remarkable increase of the B. neuwiedi group sampling size, since representatives of all the current recognized species from a wide geographic range are included in this study, providing basic information for understanding the evolution and conservation of Neotropical biodiversity.