An integrative systematic revision and biogeography ofRhynchocalamussnakes (Reptilia, Colubridae) with a description of a new species from Israel

Abstract

BackgroundThe colubrid snakes of the genus Rhynchocalamus are seldom studied and knowledge of their ecology and life history is scarce. Three species of Rhynchocalamus are currently recognized, R. satunini (from Turkey eastwards to Iran), R. arabicus (Yemen and Oman), and R. 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 known Rhynchocalamus species in order to review the intra-generic phylogenetic relationships and historical biogeography of the genus.MethodsWe revised the systematics of Rhynchocalamus using 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, Muhtarophis and Lytorhynchus. 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.ResultsThe molecular results support Rhynchocalamus as monophyletic, and as having split from its sister genus Lytorhynchus during the Late Oligocene. The three recognized species of Rhynchocalamus comprise 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 to R. arabicus from Oman than to the northern populations of the species from Israel, Syria and Turkey. Herein we describe this population as a new species: Rhynchocalamus dayanae sp. nov.DiscussionWe identify four species within Rhynchocalamus: R. satunini, R. arabicus, R. melanocephalus, and R. dayanae sp. nov., the latter, to the best of our knowledge, is endemic to southern Israel. The onset of Rhynchocalamus diversification 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.

Additions to the phylogeny of colubrine snakes in Southwestern Asia, with description of a new genus and species (Serpentes: Colubridae: Colubrinae)

“…The number of supralabials and infralabials is not totally unique in dwarf snakes, hence 7-8 supralabials and 7-9 infralabials are present in the genus Eirenis and in D. andreanus as well. (Olgun et al, 2007) Although previous phylogenetic studies did not unambiguously resolve the phylogenetic position of Muhtarophis barani or Baran's Black-headed Dwarf Snake (Avcı et al, 2015;Šmíd et al, 2015;Tamar et al, 2016), our tree surprisingly places it strongly as the sister group to the genus Scaphiophis. African Shovel-nosed Snakes (S. albopunctatus (Peters, 1870) and S. raffreyi (Bocourt, 1875) are large-sized snakes, maximum total length around 150 centimeters (Broadley, 1994), distributed around the periphery of the Central African rain forest from Ghana to western Ethiopia and adjacent Sudan (Largen & Rasmussen, 1993).…”

“…Muhtarophis barani -ZMHRU2014/60-5 Tamar et al (2016) Wallaceophis gujaratensis -NCBS HA-105 Mirza et al (2016) Wallophis brachyura -- Mirza & Patel (2018) Lytorhynchus maynardi Iran MVZ234499 Tamar et al (2016) Lytorhynchus maynardi Pakistan MVZ248463 Tamar et al (2016) Lytorhynchus gaddi Iran MVZ234500 Tamar et al (2016) Lytorhynchus diadema Oman CN4093 Tamar et al (2016) Lytorhynchus diadema Morocco IBES1329 Tamar et al (2016) Lytorhynchus diadema Egypt SPM002589 Tamar et al (2016) Rhynchocalamus satunini Iran CAS228723 Tamar et al (2016) Rhynchocalamus satunini Turkey ZMHRU2015/0 Tamar et al (2016) Rhynchocalamus arabicus Oman CN4780 Tamar et al (2016) Rhynchocalamus dayanae Israel TAU. R17093 Tamar et al (2016) Rhynchocalamus melanocephalus Israel HUJ.…”

The biogeography ofElaphe sauromates(Pallas, 1814), with a description of a new rat snake species

“…nov. Given the commonly observed biogeographic pattern in this region (Jandzik, Avcı & Gvoždík, 2013; Jandzik et al, 2018; Stümpel et al, 2016; Tamar et al, 2016; Kornilios, 2017), it is also possible that these, likely, isolated populations have diverged from one of these species (or their common ancestor) and represent a yet undiscovered taxon (Zinner, 1972).…”

“…2) in mtDNA trees. The uncorrected p -distances between both lineages in mtDNA sequences reach values around 7% (Table 3), which is very similar to distances observed in many other closely related snake species, for example, E. dione and E. bimaculata (~9%) , E. quadrivirgata and E. carinata (~8.6%), E. quadrivirgata and E. schrenckii (~8.2%), Z. persicus Werner, 1913 and Z. longissimus (Laurenti, 1768) (~8%), Vipera ursinii (Bonaparte, 1835), V. graeca Nilson & Andrén, 1988, and V. renardi (Christoph, 1861) (~4.5%), Rhynchocalamus melanocephalus (Jan, 1862) and R. dayanae Tamar, Šmíd, Göçmen, Meiri, Carranza, 2016 (4–10.2%) (Utiger et al, 2002; Gvoždík et al, 2012; Huang et al, 2012; Tamar et al, 2016; Mizsei et al, 2017). Two out of four nuclear genes ( MC1R and RAG1 ) are also clearly differentiated, though closely related between E. urartica sp.…”

Snakes of Şanlıurfa Province

“…Rhynchocalamus cinsine ait 5 tür bulunmaktadır. Bunlar; R. melanocephalus (Jan, 1862), R. satunini (Nikolsky, 1899), R. arabicus Schmidt, 1933, R. dayanae (Tamar, Šmíd, Göçmen, Meiri, & Carranza, 2016) ve R. levitoni Torki, 2017. R. satunini (Nikolsky, 1899, ilk kez Contia satunini olarak Nikolsky tarafından 1899 yılında Megri (Ermenistan yakınlarından)'den tanımlanmıştır.…”