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Phylogenetic Relationships of Old World Ratsnakes Based on Visceral Organ Topography, Osteology, and Allozyme Variation

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... Estudios sobre el amplio género Elaphe Fitzinger, 1833, tanto mor-Elaphe fológicos (Helfenberger, 2001), inmunológicos (Minton, 1976), de electroforesis (Dowling et al., 1983, Helfenberger, 2001, o de análisis de ADN (Lenk et al., 2001;Utiger et al., 2002;Nagy et al., 2004), plantean su polifiletismo, y varios de ellos sugieren la revalidación del género Rhinechis Michahelles, 1833 para denominar a Elaphe scalaris (Schinz, 1822). La combinación propuesta sería Rhinechis scalaris (Schinz, 1822). ...
... Estudios sobre el amplio género Elaphe Fitzinger, 1833, tanto mor-Elaphe fológicos (Helfenberger, 2001), inmunológicos (Minton, 1976), de electroforesis (Dowling et al., 1983, Helfenberger, 2001, o de análisis de ADN (Lenk et al., 2001;Utiger et al., 2002;Nagy et al., 2004), plantean su polifiletismo, y varios de ellos sugieren la revalidación del género Rhinechis Michahelles, 1833 para denominar a Elaphe scalaris (Schinz, 1822). La combinación propuesta sería Rhinechis scalaris (Schinz, 1822). ...
... La razón que se aduce para su conservación es su amplia utilización durante más de 50 años. La combinación Rhinechis scalaris ya ha sido utilizada al menos por Helfenberger (2001), Utiger et al. (2002) y Nagy et al. (2004. ...
... Among the most emblematic members are the rat snakes of the genus Elaphe Fitzinger in Wagler, 1833, once a large genus with distribution in temperate, subtropical, and tropical zones of both eastern and western hemisphere (Schulz, 1996). With the rise of molecular taxonomy in the last decades of the 20th century, the complicated relationships among dozens of species have been studied with a new rigor, resulting in the New-World as well as most Old-World species being identified as new or resurrected genera (Helfenberger, 2001;Lenk, Joger & Wink, 2001;Utiger et al., 2002;Chen et al., 2017). As a consequence, the genus Elaphe is comprised of 15 species (Wallach, Williams & Boundy, 2014;Chen et al., 2017), with most occurring exclusively in Asia. ...
... It was described as a separate species under the name Elaphis sauromates (see : Strauch, 1873), followed by being synonymized with Coluber quatuorlineatus Bonnaterre, 1790, and subsequently treated as its subspecies C. quatuorlineatus sauromates (Boulenger, 1894). At the beginning of the 21st century, morphological and molecular-phylogenetic studies justified the species status of E. sauromates (Helfenberger, 2001;Lenk, Joger & Wink, 2001;Utiger et al., 2002) and this view has been largely accepted to date (Sindaco, Venchi & Grieco, 2013;Sillero et al., 2014;Wallach, Williams & Boundy, 2014;Jablonski, Soltys & Simonov, 2019). Aside from confirming the separate species status of E. sauromates, early genetic studies found a surprisingly high genetic distance between populations from Kazakhstan and eastern Turkey (5.0-6.5% in Lenk, Joger & Wink, 2001 or even 7.6% in Huang et al., 2012), which was quite similar to the distances between E. sauromates and E. quatuorlineata, E. dione and E. bimaculata Schmidt, 1925, E. quadrivirgata (Boie, 1826 and E. carinata (Günther, 1864), or between E. quadrivirgata and E. schrenckii (Strauch, 1873) (Utiger et al., 2002;Huang et al., 2012). ...
... Since the publication of Boulenger's Catalogue of the snakes in the British Museum (1894) E. sauromates had been considered an eastern subspecies of E. quatuorlineata. Based solely on genetic distances between the two taxa, Helfenberger (2001) and Lenk, Joger & Wink (2001) proposed their split into two sister species, which has been largely accepted. The distribution of both species is parapatric (Sindaco, Venchi & Grieco, 2013;Kornilios et al., 2014). ...
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Background: The rat snake genus Elaphe once comprised several dozens of species distributed in temperate through tropical zones of the New and Old World. Based on molecular-genetic analyses in early 2000s, the genus was split into several separate genera, leaving only 15 Palearctic and Oriental species as its members. One of the three species also occurring in Europe is Elaphe sauromates, a robust snake from the Balkans, Anatolia, Caucasus, Ponto-Caspian steppes, and Levant that has been suspected to be composed of two or more genetically diverse populations. Here, we studied the genetic structure and morphological variation of E. sauromates, aiming to better understand its inter-population relationships and biogeography, and subsequently revise its taxonomy. Methods: We reconstructed the phylogeography and analyzed the genetic structure of E. sauromates populations originating from most of its geographic range using both mitochondrial (COI, ND4) and nuclear (C-MOS, MC1R, PRLR, RAG1) DNA gene fragments. We employed Maximum likelihood and Bayesian inference methods for the phylogenetic tree reconstructions, supplemented with species delimitation methods, analysis of haplotype networks, and calculation of uncorrected p-distances. Morphological variation in 15 metric and 18 meristic characters was studied using parametric univariate tests as well as multivariate general linearized models. In total, we analyzed sequences originating from 63 specimens and morphological data from 95 specimens of E. sauromates sensu lato. Results: The molecular phylogeny identified two clearly divergent sister lineages within E. sauromates, with both forming a lineage sister to E. quatuorlineata. The genetic distance between them (5.80–8.24% in mtDNA) is similar to the distances among several other species of the genus Elaphe. Both lineages are also moderately morphologically differentiated and, while none of the characters are exclusively diagnostic, their combination can be used for confident lineage identification. Here, following the criteria of genetic and evolutionary species concepts, we describe the lineage from eastern Anatolia and parts of the Lesser and Great Caucasus as a new species E. urartica sp. nov. Discussion: Elaphe urartica sp. nov. represents a cryptic species whose ancestors presumably diverged from their common ancestor with E. sauromates around the Miocene-Pliocene boundary. The intraspecific genetic structure indicates that the recent diversity of both species has been predominantly shaped by Pleistocene climatic oscillations, with glacial refugia mainly located in the Balkans, Crimea, and/or Anatolia in E. sauromates and Anatolia and/or the Caucasus in E. urartica sp. nov.
... The head is distinguishable from the neck, the trunk vertebra lack a hypapophysis, and the posterior maxillary teeth are not significantly differentiated from the others. With the rise of molecular taxonomy in the last decades of the 20 th century, a series of major taxonomic changes occurred at generic and species levels, resulting in the establishment or resurrection of several genera and species (Helfenberger 2001;Lenk Joger and Wink 2001;Utiger et al. 2002;Huang et al. 2012;. Recent molecular phylogenetic studies suggest that the genus Orthriophis should be subsumed within the genus Elaphe, because the generic status of Orthriophis renders Elaphe paraphyletic, where E. zoigeensis Huang, Ding, Burbrink, Yang, Huang, Ling, Chen & Zhang, 2012 is sister to all the other Elaphe plus Orthriophis (Chen et al. 2017;Li et al. 2020; but see Figueroa et al. 2016). ...
... Recent molecular phylogenetic studies suggest that the genus Orthriophis should be subsumed within the genus Elaphe, because the generic status of Orthriophis renders Elaphe paraphyletic, where E. zoigeensis Huang, Ding, Burbrink, Yang, Huang, Ling, Chen & Zhang, 2012 is sister to all the other Elaphe plus Orthriophis (Chen et al. 2017;Li et al. 2020; but see Figueroa et al. 2016). Currently, the genus Elaphe sensu stricto is comprised of 16 species of which most, are widely distributed in eastern Asia and the south slopes of the Himalaya, although the range of the genus extends east to eastern Russia, south to the Indonesia-Malayan region, and west to as far as Italy (Helfenberger 2001;Zhao 2006;Huang et al. 2012;Uetz et al. 2021). Previous biogeographic, phylogenetic, and phylogenomic studies support the hypothesis that Elaphe originated in the Eastern Palearctic (Lenk, Joger and Wink 2001;Utiger et al. 2002;Burbrink and Lawson 2007;Burbrink and Pyron 2010;Chen et al. 2017). ...
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A sheep in wolf's clothing: Elaphe xiphodonta sp. nov. (Squamata, Colubridae) and its possible mimicry to Protobothrops jerdonii. Abstract Based on combined morphological and osteological characters and molecular phylogenetics, we describe a new species of the genus Elaphe that was discovered from the south slope of the Qinling Mountains, Shaanxi, China, namely Elaphe xiphodonta sp. nov. It is distinguished from the other congeners by a combination of the following characters: dorsal scales in 21-21-17 rows, the medial 11 rows keeled; 202-204 ventral scales, 67-68 subcaudals; two preoculars (including one subpreocular); two postoculars; two anterior temporals, three posterior temporals; reduced numbers of maxillary teeth (9+2) and dentary teeth (12); sharp cutting edges on the posterior or posterolateral surface of the rear maxillary teeth and dentary teeth; dorsal head yellow, three distinct markings on the head and neck; a distinct black labial spot present in supralabials; dorsum yellow, 46-49 complete (or incomplete) large black-edged reddish brown blotches on the body and 12-19 on the tail, two rows of smaller blotches on each ventrolateral side; ventral scales yellow with mottled irregular black blotches, a few irregular small red spots dispersed on the middle of the ventral. Based on molecular phylogenetic analyses, the new species forms the sister taxon to E. zoigeensis. The discovery of this new species increases the number of the recognized species in the genus Elaphe to 17. A peer-reviewed open-access journal Shuo Qi et al. / ZooKeys 1048: 23-47 (2021) 24
... Modern materials have not yet been used to study the osteology of Elaphe dione and Elaphe schrenckii. The exception is their vertebrae (Helfenberger, 2001;Ratnikov, 2004). The aim of the first study was to estimate the phylogenetic relationships between ratsnake species in the Old World by using, among other things, vertebra features. ...
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The taxonomic identification of Quaternary remains requires a comparison of fossil bones with the skeletons of existing taxa based on the morphological criteria. However, the osteology of most modern snake species has not yet been studied. This work is aimed at finding differences in the bone morphology of Amur and Dione’s ratsnakes and the criteria for their identification. The variety of bone morphology depends on several parameters. First, these are species differences which determine the systematic position of the animal. Second, it is individual variability. Third, these are progressive changes in the vertebrae morphology along the vertebral column from the first vertebra to the last. Fourth, it is age-related variability. 20 skeletonised specimens: 12 specimens of Elaphe dione and 8 specimens of Elaphe schrenckii were used to define diagnostic osteological characters. The bones of the two studied species are very similar and their variability overlaps. However, differences and identification criteria were found for 13 skull bones: nasal, frontal, parietal, supraoccipital, maxilla, quadrate, palatine, pterygoid, ectopterygoid, basiparasphenoid, basioccipital, dentary and the compound bone. In addition, the variability and differences of the cervical, trunk, cloacal, and caudal vertebrae were examined.
... Smith (1943) considered Coluber helena as a member of the genus Elaphe. Later, Helfenberger (2001) revalidated the genus Coelognathus and transferred E. helena to Coelognathus. Now, C. helena has three subspecies, namely C. h. ...
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We provide an account on the distribution, morphology and biology of the Indian trinket snake, Coelognathus helena (Daudin) from Gujarat, India, and report the first record of the subspecies, Coelognathus helena nigriangularis Mohapatra, Schulz, Helfenberger, Hofmann, and Dutta from the Gujarat state based on reptile surveys throughout the state. We show that our understanding regarding the morphology of this species is not fully known as our series of specimens shows range of 204-245 ventral scales in C. h. helena and 219-279 ventral scales, 78-98 subcaudal scales in C. h. monticollaris.
... 7 Elaphe (quatuorlineata) sauromates is native to Romania. The separate species status for E. quatuorlineata and E. sauromates, as opposed to both being subspecies of E. quatuorlineata, has been long argued (Helfenberger 2001;Lenk et al. 2001;Utiger et al. 2002) and widely accepted (e.g. Uetz et al. 2020, Speybroeck et al. 2020) but, as also discussed by Iftime, 2010, reproductive isolation/ incompatibility between the two forms has not been actually investigated, quoted authors advocating for the split on the grounds of genetic distance and parapatric distribution with limited contact. ...
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The paper presents a review of literature data, supplemented with original observations, on the presence, establishment, distribution and invasive status of alien fish, amphibian and reptile species in Romania. Consistent criteria were followed in defining alien species records, establishment and invasive status. From the 48 alien fish species, 1 fish hybrid, 1 amphibian and 18 reptile species recorded, only 16 fishes and 3 reptiles can be regarded as established. Of these we consider the criteria for invasive status as being probably fulfilled by one fish species ( Perccottus glenii ), and less likely by six more fish species. The presence and the alien status of the one amphibian are debatable. No reptile species can be considered invasive at present.
... Most Eurasian "colubrine" snakes have a neural spine which is inclined anteriorly, e.g., members of the genera Elaphe Fitzinger in Wagler, 1833 (sensu lato). Exceptions include the extinct Elaphe paralongissima Szyndlar, 1984, and the extant Elaphe rudoforsata (Cantor, 1842) (currently placed in its own genus Oocatochus Helfenberger, 2001) where the anterior margin of the neural spine is vertical (see Szyndlar, 1991a;Ratnikov, 2004), Coronella Laurenti, 1768, Hierophis Fitzinger in Bonaparte, 1834, most members of Dolichophis Gistel, 1868, and others as well (Szyndlar, 1991a;Ikeda, 2007 [Gilmore, 1938] and Pantherophis pliocenicus [Holman, 1968]), Phyllorhynchus Stejneger, 1890, and Rhinocheilus Baird and Girard, 1853. In contrast, the posteriorly inclined anterior margin has been documented only in the extinct Pseudocemophora antiqua Auffenberg, 1963, from the early Miocene of USA and Lampropeltis vetustum Auffenberg, 1963, from the late Miocene of USA (Auffenberg, 1963;Holman, 2000). ...
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We herein describe the fossil amphibians and reptiles from the Neogene (latest Miocene or earliest Pliocene; MN 13/14) locality of Maramena, in northern Greece. The herpetofauna is shown to be extremely diverse, comprising at least 30 different taxa. Amphibians include at least six urodelan (Cryptobranchidae indet., Salamandrina sp., Lissotriton sp. [Lissotriton vulgaris group], Lissotriton sp., Ommatotriton sp., and Salamandra sp.), and three anuran taxa (Latonia sp., Hyla sp., and Pelophylax sp.). Reptiles are much more speciose, being represented by two turtle (the geoemydid Mauremys aristotelica and a probable indeterminate testudinid), at least nine lizard (Agaminae indet., Lacertidae indet., ?Lacertidae indet., aff. Palaeocordylus sp., ?Scincidae indet., Anguis sp., five morphotypes of Ophisaurus, Pseudopus sp., and at least one species of Varanus), and 10 snake taxa (Scolecophidia indet., Periergophis micros gen. et sp. nov., Paraxenophis spanios gen. et sp. nov., Hierophis cf. hungaricus, another distinct “colubrine” morphotype, Natrix aff. rudabanyaensis, and another distinct species of Natrix, Naja sp., cf. Micrurus sp., and a member of the “Oriental Vipers” complex). The autapomorphic features and bizarre vertebral morphology of Periergophis micros gen. et sp. nov. and Paraxenophis spanios gen. et sp. nov. render them readily distinguishable among fossil and extant snakes. Cryptobranchids, several of the amphibian genera, scincids, Anguis, Pseudopus, and Micrurus represent totally new fossil occurrences, not only for the Greek area, but for the whole southeastern Europe. The four different types of serration within the Varanus teeth from Maramena raise questions on the taxonomic importance or the variability of this feature. The large number of distinct amphibian and reptile taxa in Maramena makes this Greek locality by far the most diverse and speciose among all European localities across the latest Miocene and earliest Pliocene. An estimation of the palaeoprecipitation value of the locality is provided. The biogeographic origins of the Maramena herpetofauna are not fully resolved, though certain of its elements were previously only known from the early and middle Miocene of Central Europe.
... To construct the dichotomous key presented herein, we relied heavily upon the works of Taylor (1917Taylor ( , 1918Taylor ( , 1919Taylor ( , 1922aTaylor ( -c, 1923Taylor ( , 1925Taylor ( , 1963, Leviton (1957Leviton ( , 1964aLeviton ( -d, 1965aLeviton ( -c, 1967Leviton ( , 1970aLeviton ( -c, 1979Leviton ( , 1983, Inger and Marx (1965), Leviton et al. (2014), andWynn et al. (2016). We also used character state data from additional publications for Philippine members of the following genera: Acutotyphlops (Wallach et al. 2007); Ahaetulla (Gaulke 1994); Boiga (Peters 1861(Peters , 1867Gaulke 2004a); Calliophis and Hemibungarus ; Cerberus (Murphy et al. 2012;Barrera Jr. et al. 2017); Coelognathus (Helfenberger 2001); Dendrelaphis (Gaulke 2004b;Rooijen and Vogel 2012;Vogel and van Rooijen 2008); Gerrhopilus (Savage 1950); Gonyosoma (Dowling 1958); Hologerrhum (Brown et al. 2001); Hydrophis (Kharin 1984;Rasmussen 1989Rasmussen 2011Rasmussen , 2014Kharin and Hallerman 2009;Sherratt et al. 2018); Laticauda (Kharin 2005); Lycodon (Ota and Ross 1994;Lanza 1999;Gaulke 2002;Ota 2000); Malayotyphlops (Wynn and Leviton 1993;Hedges et al. 2014); Naja (Wüster and Thorpe 1996); Oligodon (Gaulke 1981;Green 2010); Opisthotropis (Brown and Leviton 1961;Yang et al. 2011); Pseudorabdion (Leviton and Brown 1959;Inger and Leviton 1966;Brown et al. 1999;Doria and Petri 2010); Ptyas (Ross et al. 1987;Malkmus et al. 2002); Ramphotyphlops (Gaulke 1995;Wallach 1993); Sibynophis (Gaulke 1993); Stegonotus (Boulenger 1893; Sanguila et al. 2016)); Trimeresurus (Malhotra and Thorpe 2004;David et al 2011); Tropidonophis (Malnate and Underwood 1988); and Tropidolaemus (Vogel et al. 2007). In addition to synthesizing data from earlier publications, we examined formalin/alcohol-preserved specimens at the University of Kansas Biodiversity Institute (KU). ...
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Aims: Yunnan Province has the richest biodiversity among all administrative regions in China. Therefore, having detailed, updated checklists of different fauna and flora groups of Yunnan are particularly important for the conservation and scientific utilization of biodiversity in China. Methods: Based on published literatures and examination of relevant specimens in natural history museums in China, we update the checklist of the reptilian fauna of Yunnan. Following the update, we revised the zoogeographic division of reptilian fauna of Yunnan and compiled diversity-related statistics for each zoogeographic region. Results: As of 31th December, 2021, there are 235 recognized species of reptiles in 82 genera, 25 families, and 2 orders recorded from Yunnan Province of China, including 16 species of Testudines in 12 genera, 4 families, 72 species of Lacertilia in 20 genera, 6 families, and 147 species of Serpentes in 50 genera, 15 families. Comparing to the latest monograph, Amphibia and Reptilia of Yunnan, which was published in 2008, our updated checklist added 82 new records, retained 21 questionable records, and removed records of 23 recognized species from Yunnan. With the revised taxonomy and distribution data of Yunnan’s reptilian fauna, we continued to recognize six zoogeographic regions in Yunnan, namely Northwestern Hengduan Mountains of Yunnan, Western Hills of Yunnan, Southern Hills of Yunnan, Southeastern Hills of Yunnan, Northern and Central Yunnan Plateau, and Northeastern Hills of Yunnan; but we adjusted the ranges for four of these regions, namely Northwestern Hengduan Mountains of Yunnan, Western Hills of Yunnan, Southeastern Hills of Yunnan, and Northern and Central Yunnan Plateau. While the three southern zoogeographic regions have the highest overall diversity, the Northwestern Hengduan Mountains of Yunnan and Central Yunnan Plateau have the highest percentage of endemic species. In total, 13% of the recorded taxa are endemic to Yunnan, 33% of the taxa are only found in Yunnan within China, and 26% of the taxa have been initially described from Yunnan. Taxonomically, Lacertilia constitutes the highest percentage of endemic taxa, which is followed by Serpentes and Testudines. For conservation, about 34% of the assessed reptile species of Yunnan are considered threatened based on China’s Red List of Biodiversity•Vertebrates (Vol. III): Reptiles, and about 16% of the total species of Yunnan still lack conservation assessments. In contrast, only 12% of the recorded species are nationally protected. Of the six zoogeographic regions of Yunnan, the Southern Hills of Yunnan have the highest percentage of threatened species and the highest number of nationally protected species. Conclusion: The reptilian diversity of Yunnan is still underestimated, and the taxonomy of the recorded species is changing regularly. Taxonomy should continue to be the focus of herpetological studies in the future, and detailed distribution data at higher resolution are needed, preferably to the county level. The percentage of endemic species of Yunnan and the conservation threat of Yunnan’s reptilian fauna are both high. Habitat conservations of endemic species in northwest and central Yunnan warrants particular attention. Lastly, as taxonomy and conservation status of species are changing regularly, and given many threatened species are not currently protected by the List of Wild Animals under Special State Protection, we call for the update of the List of Wild Animals under Special Provincial Protection of Yunnan, so that the overlooked, threatened species and their habitats can have legal protection converge.
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The herpetology collection of the Ewha Womans University Natural History Museum (EWNHM) represents one of the oldest and largest institutional collections in the Republic of Korea. The specimens deposited in the EWNHM represent a major historical collection of the native herpetofauna, both in species diversity and time span. However, the full inventory of the herpetology collection has never been conducted and thus the collection has received little attention from researchers. Here, we provide the first full account of the herpetology specimens held at the EWNHM. We provide voucher information for all documented specimens to make the collection accessible for future studies.
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