title: An integrative systematic revision and biogeography ofRhynchocalamussnakes (Reptilia, Colubridae) with a description of a new species from Israel.
abstract: BACKGROUND
  The colubrid snakes of the genusRhynchocalamusare seldom studied and knowledge of their ecology and life history is scarce. Three species ofRhynchocalamusare currently recognized,R. satunini(from Turkey eastwards to Iran),R. arabicus(Yemen and Oman), andR. melanocephalus(from the Sinai Peninsula northwards to Turkey). All are slender, secretive, mainly nocturnal and rare fossorial snakes. This comprehensive study is the first to sample all knownRhynchocalamusspecies in order to review the intra-generic phylogenetic relationships and historical biogeography of the genus.
  
  
  METHODS
  We revised the systematics ofRhynchocalamususing an integrative approach and evaluated its phylogeography. The phylogenetic position within the Colubridae and the phylogenetic relationships within the genus were inferred using 29 individuals belonging to the three known species, with additional sampling of two other closely-related genera,MuhtarophisandLytorhynchus. We analysed three mitochondrial (12S,16S, cytb) and one nuclear (c-mos) gene fragments. Phylogenetic trees were reconstructed using maximum likelihood and Bayesian inference methods; the latter method also used to provide the first time-calibrated molecular phylogeny of the genus. We generated a nuclear network and carried out a topology test and species delimitation analysis. Morphological comparisons were used to differentiate among species and to describe a new species from Israel. The studied material was comprised of 108 alcohol-preserved specimens, 15 photographs, and data from the literature for the examination of 17 mensural, 14 meristic, and two categorical characters.
  
  
  RESULTS
  The molecular results supportRhynchocalamusas monophyletic, and as having split from its sister genusLytorhynchusduring the Late Oligocene. The three recognized species ofRhynchocalamuscomprise four independently evolving groups. The molecular results reveal that the genus began to diverge during the Middle Miocene. We revealed that the best-studied species,R. melanocephalus,is paraphyletic. A population, formally ascribed to this species, from the Negev Mountain area in southern Israel is phylogenetically closer toR. arabicusfrom Oman than to the northern populations of the species from Israel, Syria and Turkey. Herein we describe this population as a new species:Rhynchocalamus dayanaesp. nov.
  
  
  CONCLUSIONS
  We identify four species withinRhynchocalamus:R. satunini, R. arabicus, R. melanocephalus, andR. dayanaesp. nov., the latter, to the best of our knowledge, is endemic to southern Israel. The onset ofRhynchocalamusdiversification is very old and estimated to have occurred during the Middle Miocene, possibly originating in the Levant region. Radiation probably resulted from vicariance and dispersal events caused by continuous geological instability, sea-level fluctuations and climatic changes within the Levant region.
authors.affiliation: The Steinhardt Museum of Natural History, Israel National Center for Biodiversity Studies, Tel Aviv University, Tel Aviv, Israel; Department of Zoology, Tel Aviv University, Tel Aviv, Israel.
authors.forename: Karin
authors.initials: K
authors.lastname: Tamar
authors.affiliation: South African National Biodiversity Institute , Cape Town , South Africa.
authors.forename: Jiří
authors.initials: J
authors.lastname: Šmíd
authors.affiliation: Department of Biology, Zoology Section, Ege University , Bornova-Izmir , Turkey.
authors.forename: Bayram
authors.initials: B
authors.lastname: Göçmen
authors.affiliation: The Steinhardt Museum of Natural History, Israel National Center for Biodiversity Studies, Tel Aviv University, Tel Aviv, Israel; Department of Zoology, Tel Aviv University, Tel Aviv, Israel.
authors.forename: Shai
authors.initials: S
authors.lastname: Meiri
authors.affiliation: Animal Biodiversity and Evolution, Institute of Evolutionary Biology (CSIC-Pompeu Fabra University) , Barcelona , Spain.
authors.forename: Salvador
authors.initials: S
authors.lastname: Carranza
keywords: Arabia; Diversification; Evolution; Middle east; Phylogeny; Phylogeography; Reptiles; Taxonomy