Molecular phylogeny, species limits, and biogeography of the Brazilian endemic lizard genus Enyalius (Squamata: Leiosauridae): An example of the historical relationship between Atlantic Forests and Amazonia

The endemic Brazilian Enyalius encompasses a diverse group of forest lizards with most species restricted to the Atlantic Forest (AF). Their taxonomy is problematic due to extensive variation in color pattern and external morphology. We present the first phylogenetic hypothesis for the genus based on 2102 bp of the mtDNA (cyt-b, ND4, and 16S) and nuclear (c-mos) regions, uncovering all previously admitted taxa (9 spp). Different methods of tree reconstruction were explored with Urostrophus vautieri, Anisolepis grilli and A. longicauda as outgroups. The monophyly of Enyalius and its split into two deeply divergent clades (late Oligocene and early Miocene) is strongly supported. Clade A assembles most lineages restricted to south and southeastern Brazil, and within it Enyalius brasiliensis is polyphyletic; herein full species status of E. brasiliensis and E. boulengeri is resurrected. Clade B unites the Amazonian E. leechii as sister-group to a major clade containing E. bilineatus as sister-group to all remaining species from northeastern Brazil. We detected unrecognized diversity in several populations suggesting putative species. Biogeographical analyses indicate that Enyalius keeps fidelity to shadowed forests, with few cases of dispersal into open regions. Ancient dispersal into the Amazon from an AF ancestor may have occurred through northeastern Brazil.This study was supported by the Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP (2003/10335-8 and 2011/50146-6), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), and partially by the Dimensions of Biodiversity Program [FAPESP (BIOTA, 2013/50297-0), NSF (DOB 1343578), and NASA]. The authors are grateful to IBAMA for collecting permits; to S. Baroni, M. Conscistré, B. Freire, and M. Antunes for assisting with lab work; and to D. Borges, D. Pavan, D. Vrcibradic, F. Curcio, F. Juncá, J. Cassimiro, J. Gasparini, M.B. Souza, H. Zaher, M.A. Freitas, M. Dixo, M.B. Martins, M. Teixeira Jr., N. Liou, R. Feio, S. Favorito, T. Tomé, V. Verdade, and V. Xavier for tissues samples and fieldwork. Thanks also to M. Teixeira Jr. for helping with figures, to Cyberinfrastructure for Phylogenetic Research (CIPRES) for providing accessibility of computational resources used in part of our analyses, and Mariana Morando and an anonymous reviewer for helpful comments. M. Carvalho and M.A. Carvalho kindly revised the idiom of the manuscript