DOI: 10.3897/vz.76.e169219

Authors: Diego Omar Di Pietro, Julieta Sánchez, Sebastián Poljak, Leandro Alcalde

Abstract: We analysed the genetic and morphological variation in Philodryas patagoniensis, a widely distributed South American racer snake. Two well-differentiated haplogroups were identified using mitochondrial gene sequences (12S and 16S) and the nuclear gene c-mos. Genetic divergence between these haplogroups correlates strongly with morphological differences, allowing the recognition of two morphotypes within P. patagoniensis. We integrated genetic and morphological data into a total evidence analysis using parsimony. Our results support the distinction between the two haplogroups/morphotypes, consistent with recognising two species within P. patagoniensis. Accordingly, we re-describe P. patagoniensis, refining its morphological variation and geographical distribution to reflect the observed genetic differentiation, and describe a new species. Morphological characteristics can distinguish the two species, including body measurements, scale patterns, and cranial osteology. The new species differs from P. patagoniensis sensu stricto in traits associated with arboreal habits, which are strongly correlated with the distribution of the two taxa across forested and open habitats in South America. Furthermore, P. patagoniensis sensu stricto has a significant Lycosa spider component in its diet, which is absent in the newly described species.

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DOI: 10.3897/vz.75.e158031

Authors: Shuo Qi, Hai Ngoc Ngo, L. Lee Grismer, Hao-Tian Wang, Han-Ming Song, Xiao-Yu Zhu, Zhu-Qing He, Zi-Chen Zhou, Pi-Peng Li, Ji-Chao Wang, Ying-Yong Wang

Abstract: Understanding the taxonomy of the Goniurosaurus lichtenfelderi group has long been obscured by limited sampling, inconsistent morphological characters, and a lack of publicly available molecular and distribution data. In this study, we conducted the most comprehensive morphological and molecular assessment of this group to date, integrating extensive sampling from Hainan Island and adjacent mainland regions. Phylogenetic analyses using mitochondrial and nuclear genes, along with SNPs delivered from ddRAD-seq, revealed that G. hainanensis is polyphyletic, consisting of two deeply divergent clades (i.e., western and eastern) on Hainan Island. The western clade, from the type locality at Mt. Wuzhi, shows small genetic divergence from G. sinensis (= G. kwanghua), supporting their synonymy. In contrast, the eastern clade is genetically more closely related to the continental G. lichtenfelderi than to its western counterpart. Divergence time estimates further indicate that the eastern and western Hainan populations have distinct evolutionary histories. Based on integrated evidence, we synonymize G. kwanghua and G. sinensis with G. hainanensis, and delimit the eastern Hainan clade as G. cf. lichtenfelderi, pending further morphological confirmation. Consequently, the number of valid species in the G. lichtenfelderi group is revised from five to four (including G. lichtenfelderi, G. hainanensis, G. bawanglingensis, and G. zhoui). In addition, a revised taxonomic account and updated distribution information for all recognized species are provided in this study.

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DOI: 10.3897/vz.75.e142775

Authors: Anita Malhotra, Mrinalini Walter, Guillem Limia Russel, Roger S. Thorpe

Abstract: Abstract In this study, we present a comprehensive analysis of species in the Trimeresurus albolabris complex (comprising the white-lipped pitviper, its former subspecies, and closely related species) to identify genetic and phenotypic lines of evidence to assess whether these taxa represent independently evolving lineages or populations with wide-ranging variants. Our morphometric data set includes over 400 live and preserved specimens spanning almost the entire range of T. albolabris sensu stricto and its relatives. We compare patterns of differentiation in three mitochondrial gene fragments for over 300 specimens and conduct a population genetic analysis of nuclear NT3 sequences from over 200 specimens. This level of detail allowed us to identify instances of incongruence between morphological affinities, mitochondrial clades, and nuclear haplotype distribution in putative taxa throughout Indochina. Although recently described species from this region, including T. salazar, T. caudornatus, and T. uetzi, are related to T. septentrionalis in the mitochondrial phylogeny, they extensively share nuclear haplotypes with the purpureomaculatus group, particularly with T. erythrurus. The most common haplotype of T. albolabris sensu stricto is also present throughout Indochina. However, populations in southern Indochina and western Java are morphologically differentiated, belong to distinct mitochondrial clades, and have a high proportion of private NT3 alleles. As they appear to intergrade extensively with populations further north, we herein resurrect the Javan taxon Bothrops viridis var. fario Jan, 1859 as a nomen for this population at the subspecies level, as Trimeresurus albolabris fario comb. nov. Furthermore, we lower the rank of taxa from the extensive zone of intergradation that stretches from Myanmar to central Vietnam to that of subspecies under T. albolabris (T. a. guoi) or T. septentrionalis (T. s. salazar, T. s. caudornatus, and T. s. uetzi), pending detailed further investigations of the extent of gene flow between them and other recognised species in the complex.

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DOI: 10.3897/vz.74.e123485

Authors: Evgeniy Simonov, Artem Lisachov, Spartak Litvinchuk, Anastasia Klenina, Polina Chernigova, Alexander Ruchin, Andrey Bakiev, Kazhmurat Akhmedenov

Abstract: Abstract Most of the geographic range of the grass snake Natrix natrix is occupied by the subspecies N. n. scutata, which occurs from Eastern Europe to East Siberia. This study addressed the phylogeography of this subspecies via analysis of variation of partial cytochrome b (cyt b) sequences from 135 individuals sampled at 91 localities across its range, in addition to species distribution modelling. A haplotype network was constructed and identified a major star-like haplogroup that harbours most of the analysed specimens and is considered the main source of recolonisation of vast territories of northern Eurasia after the Last Glacial Maximum (LGM). Most of this subspecies’ current range is occupied by haplotypes from a single haplogroup, with probable refugia in the south of the European part of Russia or the North Caucasus. The most frequent (ancestral) haplotype in this group is the only one discovered in West Siberia among 25 specimens from 13 sampling sites, indicating relatively recent colonisation of Siberia. According to species distribution modelling, no relatively suitable areas were present in Central Asia or Siberia during the LGM. Nonetheless, the only two specimens examined from the easternmost area of the species’ geographic range, near Lake Baikal, had unique haplotypes that differed from the most common haplotype by one and two substitutions. The most probable explanation for this pattern is that N. n. scutata colonised the area during a previous interglacial period and survived here during the last glacial. The Mugodzhar Range in western Kazakhstan also showed cyt b differentiation as compared to surrounding areas but warrants further sampling to test competing hypotheses.

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DOI: 10.3897/vz.74.e123824

Authors: Simona Papežíková, Martin Ivanov, Petr Papežík, Adam Javorčík, Konrad Mebert, Daniel Jablonski

Abstract: Abstract The Western Palearctic harbours a diverse snake fauna, including numerous endemic species and yet unnamed clades, identified through molecular analyses. However, morphological characteristics of these clades, even of common species, often remain relatively unexplored. In this study, we provide an examination of the morphology and cranial anatomy of the semi-aquatic snake species Natrix tessellata (Laurenti, 1768), with a focus on populations of the so-called ‘Europe’ and ‘Greece’ clades. Utilising both museum collections and field data, we first morphologically examined 541 individuals of N. tessellata, categorising them according to previously established clades and lineages that resulted in relatively low morphometric and meristic variation across the species’ range. When assessing the 448 specimens from the ‘Europe’ and the ‘Greece’ clades separately, we similarly observed little variation in meristic characteristics. On the other hand, individuals of the ‘Greece’ clade displayed smaller and more slender body and head proportions compared to those of the ‘Europe’ clade and the pigmentation of the labial scales is distinctively paler in the ‘Greece’ clade, whereas the overall body colouration remains largely similar between the two. Our osteological analysis of 47 N. tessellata skulls also indicated slight differences in the frontoparietal portion of the braincase between the ‘Europe’ and the ‘Greece’ clades, warranting further examination with a larger dataset and extending to other skull components. These findings hold significance for ongoing enquiries into the species’ biogeography, morphology and ecological adaptations. In summary, the integration of morphological and osteological data with genetic information offers a promising avenue for potential taxonomic revisions of N. tessellata in the future.

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DOI: 10.3897/vz.74.e112132

Authors: Parinya Pawangkhanant, Nikolay A. Poyarkov, Harry Ward-Smith, Rupert Grassby-Lewis, Montri Sumontha, Nikita S. Kliukin, Sabira S. Idiiatullina, Alexei V. Trofimets, Chatmongkon Suwannapoom, Justin L. Lee

Abstract: Abstract We describe a new species of kukri snake (Oligodon Fitzinger, 1826) from the limestone karst formations of Satun and Trang Provinces in southern Thailand. Phylogenetic analyses based on three mitochondrial DNA fragments (12S–16S ribosomal rRNA and cytochrome b) recover the new species within the Oligodon cinereus species complex, where it forms a deeply divergent yet poorly supported clade sister to Oligodon saiyok Sumontha et al., 2017 and another unnamed lineage currently referred to Oligodon cinereus (Günther, 1864) from southwest Myanmar. Morphologically, the new species is distinguished from all other members of the genus by the following combination of characters: ventral scales 189–193 with distinct lateral keeling; subcaudal scales 47–54, paired; anterior dorsal scale rows 17–19, with the reduction from 19 to 17 rows occurring above the 28th–30th ventral scale when present; maxillary teeth 8, blade-like and laterally compressed; dorsum olive–gray, plain; ventral surface white anteriorly, dark gray posteriorly; underside of tail dark gray, smeared with white. We briefly discuss the natural history and conservation status of this new species and provide observations of other kukri snakes inhabiting limestone karst habitats. Our study also incorporates genetic samples of four recently described Oligodon endemic to Thailand, all of which are recovered in the O. cinereus species complex. In agreement with previous studies, we demonstrate that species-level diversity within the O. cinereus species complex is underestimated, and additional sampling is necessary to revise this taxonomically challenging clade.

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DOI: 10.3897/vz.74.e113347

Authors: Sabira S. Idiiatullina, Tan Van Nguyen, Parinya Pawangkhanant, Chatmongkon Suwannapoom, Lawan Chanhome, Zeeshan A. Mirza, Patrick David, Gernot Vogel, Nikolay A. Poyarkov

Abstract: Abstract Despite recent progress in our understanding of diversity within the genus Trimeresurus Lacépède, 1804, the subgenus Popeia Malhotra & Thorpe, 2004, distributed across most parts of East and Southeast Asia, remains taxonomically challenging. We applied an integrative taxonomic approach including analyses of morphological data and four mitochondrial genes (12S and 16S rRNA, cytochrome b, and NADH dehydrogenase subunit 4), along with examination of available type material, to address longstanding taxonomic questions in one clade within Popeia, the T. popeiorum group, and reveal a high level of hidden diversity of these snakes in the Indo-Burma Biodiversity Hotspot. We confirm that T. popeiorum Smith, 1937 sensu stricto is restricted to Northeast India, eastern Nepal, southern Bhutan, southeastern Bangladesh, western Yunnan Province (China), and northern and southwestern Myanmar. We further confirm that the recently described species T. yingjiangensis Chen et al., 2019 is a junior synonym of T. popeiorum. In addition, we discovered that the combination Trimesurus [sic] elegans Gray, 1853 is a valid senior synonym of T. popeiorum and threatens the stability of the latter taxon. Therefore, in order to protect the nomen popeiorum and in accordance with Article 23.9 of the International Code of Zoological Nomenclature, we regard the taxon Trimesurus elegans as a nomen oblitum and render Trimeresurus popeiorum a nomen protectum. Examination of a larger series of specimens allows us to describe two new cryptic species of Trimeresurus from the Indo-Burma Region. This study brings the total number of species in the subgenus Popeia to six and also suggests that the subspecific taxonomy of the T. sabahi complex requires further investigation. We urge adequate actions regarding the conservation of the newly discovered species and recommend further studies on their toxicology.

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DOI: 10.3897/vz.74.e116992

Authors: Sang Ngoc Nguyen, Manh Van Le, Amy Lathrop, Thi-Dieu-Hien Vo, Robert W. Murphy, Jing Che

Abstract: Abstract Homalopsid snakes of the genus Myrrophis include only two species distributed in southern China and northern Vietnam. Here, we describe a third species from southern Vietnam based on morphological data and nucleotide sequences from the mitochondrial gene cyt b. Myrrophis dakkrongensis sp. nov. is diagnosed by the following morphological characters: Medium-sized mud snake (largest total length 452 mm); internasal single and distinctly separated from loreals; dorsal scales smooth, in 23 rows at midbody, reduced to 19 or 20 rows before vent; ventrals 133–138; subcaudals 34–42, paired; cloacal plate divided; supralabials 8, fourth entering orbit; second pair of chin-shields small and oblique; maxillary teeth 17 or 18; gland-like tubercles present in the cloacal region; hemipenis short, forked and spinose, reaching 7th subcaudal; dorsum dark brown to black; and a white or yellow to orange lateroventral stripe present. The new species differs from its congeners by an uncorrected p distance in cyt b sequences of at least 10.5%.

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DOI: 10.3897/vz.74.e106238

Authors: Vinícius Sudré, Albedi Andrade-Junior, Manuella Folly, Josué A. R. Azevedo, Robson Waldemar Ávila, Felipe Franco Curcio, Pedro M. Sales Nunes, Paulo Passos

Abstract: Abstract Currently, the proposed diagnoses for the Chironius bicarinatus complex reflect a wide variation in color pattern and pholidosis. Herein, we review the Chironius bicarinatus complex based on morphological and molecular data from a sample of 485 specimens covering the species distribution. Our results corroborate the recognition of C. bicarinatus and C. gouveai, and diagnose a distinct lineage without an available name. Thus, here we describe this new species restricted to the Baturité Massif, a relictual rainforest isolated in the Caatinga xerophytic domain, in the state of Ceará, northeastern Brazil. The new species can be distinguished from its congeners by its unique combination of qualitative and quantitative morphological characters (scale counts, morphometric, color pattern), and is also supported by molecular and ecological evidence. Additionally, we rectify data on the distribution and morphological variability of C. gouveai to accurately infer the boundaries between this taxon and C. bicarinatus, which was not properly addressed. Finally, we discuss our results in the light of previous studies that suggest diversification hypotheses in the Atlantic Forest already detected for other taxa, highlighting the importance of conserving the areas of “Brejos de Altitude”, in northeastern Brazil, and the southern limit of Serra do Mar up to Serra do Tabuleiro, in southern Brazil.

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DOI: 10.3897/vz.73.e101372

Authors: Zbigniew Szyndlar, Georgios L. Georgalis

Abstract: Abstract We here present a thorough documentation of the vertebral morphology and intracolumnar variation across non-caenophidian snakes. Our studied sample of multiple individuals covers a large number of genera (67) and species (120), pertaining to almost all extant non-caenophidian families. Detailed figuring of multiple vertebrae across the trunk, cloacal, and caudal series for many different individuals / taxa documents the intracolumnar, intraspecific, and interspecific variation. An emphasis is given in the trunk-to-caudal transition and the pattern of the subcentral structures in that region of the column. Extant non-caenophidian snakes show an astonishing diversity of vertebral morphologies. Diagnostic vertebral features for extant families and many genera are given, though admittedly vertebral distinction among genera in certain groups remains a difficult task. A massive compilation of vertebral counts for 270 species, pertaining to 78 different genera (i.e., almost all known valid genera) and encompassing all extant non-caenophidian families, is provided based on our observations as well as an extensive literature overview. More particularly, for many taxa, detailed vertebral counts are explicitly given for the trunk, cloacal, and caudal portions of the column. Extant non-caenophidian snakes witness an extremely wide range of counts of vertebrae, ranging from 115 up to 546. A discussion on the diagnostic taxonomic utility and potential phylogenetic value of certain vertebral structures is provided. Comparisons of the subcentral structures of the cloacal and caudal vertebral series are also made with caenophidian lineages. We anticipate that this illustrative guide will set the stage for more vertebral descriptions in herpetological works but will also be of significant aid for taxonomic identifications in ophidian palaeontology and archaeozoology.

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DOI: 10.3897/vz.73.e109854

Authors: Sabira S. Idiiatullina, Parinya Pawangkhanant, Tanapong Tawan, Thanawut Worranuch, Bunyarit Dechochai, Chatmongkon Suwannapoom, Tan Van Nguyen, Lawan Chanhome, Nikolay A. Poyarkov

Abstract: Abstract We describe a new species of pitvipers from Trang Province of Thailand, near the Thailand–Malaysian border, based on morphological and molecular (2427 bp from cyt b, ND4, and 16S rRNA mitochondrial DNA genes) lines of evidence. Morphologically, Trimeresurus ciliaris sp. nov. is distinguished from its congeners by the following combination of morphological characters: a long papillose hemipenis; first supralabial and nasal scale fused; three to four small supraocular scales; internasals not in contact; small scale between nasal and the scale formed by the fused second supralabial and loreal present; dorsal scales in 17–17–15 rows across the body; ventral scales 172–175 in males, 171 in female; subcaudal scales 59–63 in males, 61 in female, all paired; in life an emerald-green dorsum with reddish-brown bands; creamy-white venter lacking dark dots or stripes on the lateral sides of the ventrals; white vertebral spots present in both sexes on every two or three dorsal scales; dark brown spots forming discontinuous pattern present on 1–3 lateral dorsal scale rows; males with reddish-brown postocular stripe. The new species forms a distinct clade on the phylogenetic tree of the genus Trimeresurus and differs from the morphologically similar species T. venustus by a significant divergence in cytochrome b mitochondrial DNA gene sequences (p = 12.5%). The new species is currently known from a small karstic area in the Nakawan Range spanning the border of Thailand and Malaysia, in particular in limestone forests in Trang and Satun provinces (Thailand); it likely also occurs in the adjacent parts of Perlis State (Malaysia). Our study also suggests that the taxonomy of T. kanburiensis species complex requires further studies; in particular our study suggests that the status of populations from Chumphon Province of Thailand and Pulau Langkawi Island of Malaysia should be re-assessed.

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DOI: 10.3897/vz.73.e108402

Authors: Mariana Chuliver, Agustín Scanferla

Abstract: Abstract Snakes lay their eggs in clutches of different size, which are usually attached to each other forming a cluster. Egg-clustering is a widespread phenomenon across alethinophidian snakes, mostly recorded in Pythonoidea and caenophidian clades. Here we report a new type of egg-clustering for threadsnakes (Leptotyphlopidae) that departs from the alethinophidian type. We found that females of Epictia australis and Leptotyphlops sylvicolus lay their eggs connected to each other through a filament, and we dubbed it ‘string-egg clustering’. The histomorphology of the filament linking the eggs in E. australis showed an outer calcareous layer underlain by a thick layer of collagen fibers, demonstrating that it is an integral part of the eggshell formed during its deposition process in the oviduct. String egg-clustering seems to be present only among species belonging to both subfamilies of threadsnakes, Epictinae and Leptotyphlopinae. Egg-clustering in alethinophidians has been demonstrated to have several advantages for embryo development and post-hatching survival, including fixing the position of the embryo within the egg, protection against predators, and embryo-to-embryo communication. The presence of a filament connecting the eggs in leptotyphlopid species might be relevant for maintaining the position of the embryo in the egg, to avoid the dispersion of the egg in the nesting site, and potentially for the transmission of physical cues. Thus, we hypothesize that the string-egg clustering constitutes an advantageous reproductive trait among threadsnakes.

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DOI: 10.3897/vz.73.e103319

Authors: Uwe Fritz, L. Lee Grismer, Marika Asztalos

Abstract: Using georeferenced photographic records of 2944 grass snakes from Germany, Austria, and northern Italy as well as previously published mtDNA sequences (n = 1062) and microsatellite data (n = 952) for grass snakes from the same regions, we examined whether or not coloration and pattern reliably allow to differentiate between Natrix natrix and N. helvetica and if so, whether the distribution patterns revealed by phenotypes and genetics are congruent. Furthermore, we used cline analyses across hybrid zones to test whether the phenotypic transition from one species to the other parallels the steep clines unveiled by genetics. Our results suggest that the two species can be reliably differentiated using coloration and pattern. The most powerful diagnostic traits are the presence/absence of side bars on the body flanks, the number of occipital spots, and the shape of the posterior dark occipital spot. The distributions of morphologically identified N. natrix and N. helvetica match their genetically confirmed ranges. Single conflicting individuals morphologically identified as N. natrix or hybrids within the distribution range of N. helvetica either represent misidentifications or translocated snakes. For the genetic markers and phenotypes, our cline analyses revealed concordant steep clines across hybrid zones. However, the southern part of the hybrid zone in Italy, for which no sufficient genetic data are available, should be studied in more detail because the phenotypic data suggest a smooth cline in this region. The unexpected high percentage of putative hybrids with dorsal stripes in this region also calls for further research. For northwestern Germany, another region for which no genetically verified records are available, iNaturalist data suggest that the contact zone of N. natrix and N. helvetica is near the Ems River and extends from there southeastwards to the region of Höxter, North Rhine-Westphalia.

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DOI: 10.3897/vz.73.e98170

Authors: Angele Martins, Manuella Folly, Guilherme Nunes Ferreira, Antônio Samuel Garcia da Silva, Claudia Koch, Antoine Fouquet, Alessandra Machado, Ricardo Tadeu Lopes, Roberta Pinto, Miguel Trefaut Rodrigues, Paulo Passos

Abstract: Threadsnakes are known for their conserved external morphology and historically controversial systematics, challenging taxonomic, biogeographic and evolutionary researches in these fields. Recent morphological studies—mostly based on µCT data of the skull and lower jaw—have resolved systematic issues within the group, for instance leading to the description of new taxa or re-positioning little-known scolecophidian taxa in the tree of life. Herein we describe a new polymorphic species of the genus Siagonodon from Amazonia based on morphological (external, osteology and hemipenis) and molecular data, and provide the first hemipenial description for the genus. We also reassign Siagonodon acutirostris to the genus Trilepida based on osteological data in combination with molecular evidence. The new species described represents an evolutionary paradox for scolecophidians because the species displays a remarkable variation in the shape of the snout region that is otherwise always highly conserved in this clade. Finally, this study reinforces the importance of protected areas as essential in maintaining vertebrate populations, including those that are not yet formally described.

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DOI: 10.3897/vz.73.e96958

Authors: Platon V. Yushchenko, Justin L. Lee, Hieu Minh Pham, Peter Geissler, Elena V. Syromyatnikova, Nikolay A. Poyarkov Jr.

Abstract: We investigated the taxonomic status of the recently described kukri snake Oligodon arenarius Vassilieva, 2015 and the morphologically similar Oligodon macrurus (Angel, 1927), two species endemic to the southern coast of Vietnam. Based on phylogenetic analyses using three mitochondrial genes (12S–16S rRNA, cytochrome b), we recovered O. arenarius and O. macrurus in a clade within the O. cyclurus-taeniatus species group, agreeing with previous intrageneric classifications. Genetic distances between O. arenarius and O. macrurus are extremely low (less than 0.5% based on 12S–16S) and render O. arenarius paraphyletic. All preserved specimens of O. arenarius and O. macrurus convey little to no differences in color pattern, hemipenial morphology and osteological features; the latter of which is based on three dimensional micro computer tomography (µCT) scans of one specimen per species. Contrasting these results, univariate and multivariate analyses revealed significant differences in relative tail length, and the number of ventral and subcaudal scales between both species. Although the molecular and morphological datasets present conflicting results, integrating the evidence leads us to synonymize O. arenarius with O. macrurus. We provide a formal redescription of O. macrurus, designate a neotype specimen to avoid future taxonomic confusion, and provide the first detailed osteological description of this species. Oligodon macrurus sensu stricto is endemic to coastal dunefields and adjacent forest habitats in southern Vietnam, where ongoing human development, tourism and road mortality pose significant threats to its conservation. Consequently, we suggest that O. macrurus should be listed as “Vulnerable” based on the assessment criteria of the International Union for Conservation of Nature (IUCN).

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DOI: 10.3897/vz.73.e91230

Authors: Platon V. Yushchenko, Justin L. Lee, Thy Neang, Hun Seiha, Nguyen Van Tan, Gernot Vogel, Nikolay A. Poyarkov Jr.

Abstract: The ashy kukri snake Oligodon cinereus (Günther, 1864) is a widely distributed and morphologically variable species found throughout mainland Southeast Asia. In this paper, we re-assessed the taxonomic status of O. cinereus populations found in southern Indochina (southern Vietnam, Cambodia, and southern Laos), including the recently described Cat Tien kukri snake Oligodon cattienensis Vassilieva et al., 2013, which was previously confused with this species. Phylogenetic analyses using mitochondrial DNA from the 12S–16S ribosomal subunit and cytochrome b gene revealed that O. cattienensis is embedded in a mixed clade containing samples of the subspecies O. cinereus pallidocinctus, which bears a dorsal color pattern with white crossbars and black edges. This clade forms a strongly supported sister group with a topotypic sample of O. cinereus cinereus, representing populations bearing a uniform dorsal color pattern and slight reticulate markings, however the genetic divergence between the two clades is very low. The morphological characters used to distinguish O. cattienensis from O. cinereus sensu lato broadly overlap and supposed differences in hemipenial morphology between the two taxa are due to outdated terminologies used to describe the organ. We relegate both O. cattienensis and O. cinereus pallidocinctus to the junior synonymy of O. cinereus and consider all color patterns of this species found near the type locality in Cambodia, southern Laos, and southern Vietnam to represent O. cinereus sensu stricto. Future integrative investigations across the range of O. cinereus sensu lato are needed to resolve the status of the remaining subspecies and synonyms associated with this taxon. Problems associated with hemipenial morphology and Oligodon systematics are also discussed.

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DOI: 10.3897/vz.72.e89647

Authors: Fred Kraus, Hinrich Kaiser, Mark O’Shea

Abstract: With its conservative set of scalation characters, Toxicocalamus loriae is a morphologically confusing species to which a wide array of phenotypes has been assigned. Careful analysis of 224 museum specimens reveals that multiple distinct species remain hidden under the name T. loriae and that diagnostic, species-level differences are more nuanced in this group of snakes than among other members of the genus. Our taxonomic reassessment leads us to resurrect the species T. lamingtoni comb. nov., T. loennbergii comb. nov., and T. nymani comb. nov. from synonymy with T. loriae, retain only T. pratti as a synonym, and describe three new species. As a consequence, T. loriae is no longer recognized as ranging throughout the entire island of New Guinea but is instead restricted to the southern versant of the Papuan Peninsula, and T. lamingtoni and T. spilorhynchus sp. nov. are species restricted to that same peninsula’s northern versant. Toxicocalamus loennbergii is known only from the type series taken on the Onin Peninsula in West Papua, Indonesia, Toxicocalamus atratus sp. nov. is a high-elevation (800–2200 m) Central Highlands endemic, and T. vertebralis sp. nov. ranges from the Central Highlands of Papua New Guinea eastward into the Wau area of Morobe Province. Toxicocalamus nymani inhabits a geologically more heterogenous region, occurring from the Central Highlands eastward to the Huon Peninsula, including Karkar Island, and adjacent areas of Madang Province as well as the northernmost reaches of the Papuan Peninsula. We expect that denser geographic sampling across New Guinea and focussed specimen collection of a few known populations will result in the recognition of additional species in this complex.

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DOI: 10.3897/vz.72.e87313

Authors: Teddy Angarita-Sierra, Sergio Daniel Cubides-Cubillos, Juan Pablo Hurtado-Gómez

Abstract: Bothrocophias microphthalmus (Cope, 1875) currently comprises most mid- to highland populations of the genus Bothrocophias in the eastern versant of the Andes. We describe two new species of Bothrocophias from the highlands of the Colombian Andes previously referred to as B. microphthalmus based on morphological and genetic evidence. Our phylogenetic analysis revealed that B. micropthalmus as currently recognized is paraphyletic with respect to B. hyoprora, and the two new taxa are sister lineages. These new toadheaded pitvipers can be morphologically distinguished from their congeners based on the presence of contact between the internasal scales, the number of prefoveal scales, the presence of a lacunolabial scale, the arrangement of supralabial scales, ventral scale counts, the color pattern of the dorsal and ventral surfaces of the body and tail, and hemipenial morphology. We discuss some possible taxonomic scenarios for the lineages found within the B. microphthalmus species complex but refrain from making additional taxonomic changes given our reduced sampling of the southern lineages.

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DOI: 10.3897/vz.72.e87426

Authors: Uwe Fritz, Flora Ihlow

Abstract: Abstract We used a dataset of georeferenced photos of 5,751 grass snakes from iNaturalist to evaluate subspecific variation of Natrix natrix in coloration and pattern. Our results provide evidence that all four genetically delineated subspecies differ morphologically, although unstriped individuals of N. n. vulgaris are difficult to tell apart from the nominotypical subspecies. The iNaturalist dataset shows that the frequency of dark body coloration increases from south to north and from west to east. This trend is both concordant with taxonomic variation (the easternmost subspecies, N. n. scutata, being the darkest taxon) and variation within the same subspecies (in N. n. natrix and N. n. scutata more northern populations harbor more dark or melanistic individuals than more southern populations). Although available characters were limited to coloration and pattern traits, our study suggests that photo material from iNaturalist and similar platforms can be a valuable data source for studies on morphological variation. However, investigations using such databases can only supplement, but not replace, studies using museum material because only then measureable, meristic and genetic characters will be accessible.

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DOI: 10.3897/vz.72.e84516

Authors: Ho Yuen Yeung, Michael W. N. Lau, Jian-Huan Yang

Abstract: A new species of the genus Calamaria Boie, 1827, Calamaria arcana sp. nov., is described based on a single male specimen collected from Mt. Dadongshan, Guangdong, southern China. The new species can be distinguished from all known congeners by the significant genetic divergence in the mitochondrial cytochrome-b gene fragment (p-distance ≥ 13.9%), and morphologically by the combination of the following characters: (1) ten modified maxillary teeth; (2) four supralabials, second and third supralabials entering orbit; (3) preocular present; (4) mental not touching anterior chin shields; (5) six scales and shields surrounding the paraparietal; (6) 170 ventral scales; (7) 22 paired subcaudals; (8) tail not gradually tapering, abruptly tapering at the tip; (9) dorsal scales reduced to five rows above last subcaudal at tail; (10) dorsum of body and tail brownish; (11) dark collar on nuchal region absent; (12) two outermost dorsal scale rows light yellow with upper margins partly dark pigmented; (13) ventral scales immaculate, without dark outermost corners and pigmentation anteriorly; and (14) absence of distinct dark longitudinal line or scattered spots on the underside of tail. Calamaria arcana sp. nov., represents the fifth species of the genus recorded in China. Following the IUCN Red List Categories and Criteria, we propose the new species to be listed as Data Deficient.

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DOI: 10.3897/vz.71.e76453

Authors: Marika Asztalos, Dinçer Ayaz, Yusuf Bayrakcı, Murat Afsar, Cemal Varol Tok, Carolin Kindler, Daniel Jablonski, Uwe Fritz

Abstract: Using two mitochondrial DNA fragments and 13 microsatellite loci, we examined the phylogeographic structure and taxonomy of two codistributed snake species (Natrix natrix, N. tessellata) in their eastern distribution area, with a focus on Turkey. We found evidence for frequent interspecific hybridization, previously thought to be extremely rare, and for backcrosses. This underscores that closely related sympatric species should be studied together because otherwise the signal of hybridization will be missed. Furthermore, the phylogeographic patterns of the two species show many parallels, suggestive of a shared biogeographic history. In general, the phylogeographies follow the paradigm of southern richness to northern purity, but the dice snake has some additional lineages in the south and east in regions where grass snakes do not occur. For both species, the Balkan Peninsula and the Caucasus region served as glacial refugia, with several mitochondrial lineages occurring in close proximity. Our results show that the mitochondrial divergences in both species match nuclear genomic differentiation. Yet, in the former glacial refugia of grass snakes there are fewer nuclear clusters than mitochondrial lineages, suggesting that Holocene range expansions transformed the glacial hotspots in melting pots where only the mitochondrial lineages persisted, bearing witness of former diversity. On the other hand, the deep mitochondrial divergences in N. tessellata across its entire range indicate that more than one species could be involved, even though lacking microsatellite data outside of Turkey prevent firm conclusions. On the contrary, our microsatellite and mitochondrial data corroborate that N. megalocephala is invalid and not differentiated from sympatric populations of N. natrix. For Cypriot grass snakes, our analyses yielded conflicting results. A critical assessment of the available evidence suggests that N. natrix is a genetically impoverished recent invader on Cyprus and taxonomically not distinct from a subspecies also occurring in western Anatolia and the southern Balkans. Based on combined mitochondrial and nuclear genomic evidence we propose that for grass snakes the following subspecies should be recognized in our study region: (1) Natrix natrix vulgaris Laurenti, 1768, southeastern Central Europe and northern Balkans; (2) Natrix natrix moreotica (Bedriaga, 1882), southern Balkans, western Anatolia, and Cyprus; and (3) Natrix natrix scutata (Pallas, 1771), eastern Anatolia, Caucasus region, Iran, northeastern distribution range (from eastern Poland and Finland to Kazakhstan and the Lake Baikal region). Thus, Natrix natrix cypriaca (Hecht, 1930) becomes a junior synonym of N. n. moreotica and Natrix natrix persa (Pallas, 1814) becomes a junior synonym of N. n. scutata. Due to insufficient material, we could not resolve the status of Natrix natrix syriaca (Hecht, 1930) from the Gulf of İskenderun, southeastern Turkey.

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DOI: 10.3897/vz.71.e66239

Authors: Ashok Kumar Mallik, Achyuthan Needamangalam Srikanthan, Sumaithangi Rajagopalan Ganesh, Seenapuram Palaniswamy Vijayakumar, Patrick D. Campbell, Anita Malhotra, Kartik Shanker

Abstract: Abstract Asian pit vipers belonging to the genus Craspedocephalus are a complex group of vipers, distributed in South and Southeast Asia. Their taxonomy is unresolved in many lineages across their distributional range. Here, we reassess the taxonomy and systematics of pit vipers of the genus Craspedocephalus in Peninsular India based on extensive field sampling, in particular in the Western Ghats. We build and expand on the previous findings of genetic relatedness between the peninsular Indian lineages with the Sundaic clade (C. puniceus complex) with greater evidence, based on additional taxa sequenced herein. We reconstruct the phylogeny of the group using three mitochondrial genes and delineated lineages using coalescent species delimitation methods. We then used multiple criteria including genetic divergence and separation in morphological and geographic space to designate taxonomic units. Our work revealed the presence of a South Asian radiation of the clade Craspedocephalus, with a few Sundaic members. Our study reveals the systematic relationships of four Peninsular Indian species of Craspedocephalus, including Peltopelor macrolepis and C. strigatus, sequenced here for the first time, that are classified or confirmed as members of Craspedocephalus. Hence, we place the genus Peltopelor in the synonymy of Craspedocephalus. Using our multi-criteria approach, we delimit four new cryptic evolutionary lineages within the Western Ghats escarpment of Peninsular India. These cryptic lineages belong to the C. malabaricus, C. gramineus and C. macrolepis complexes and are geographically and/or ecologically (in terms of habitat association) distinct from their sister lineages across their distributional range, while others are separated in morphological space. Our new phylogenetic tree and delimitation analysis thus reveals the presence of multiple clades with several cryptic lineages separated by geographical barriers or habitat association.

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DOI: 10.3897/vz.71.e70438

Authors: Duong T. T. Le, Thinh G. Tran, Huy D. Hoang, Bryan L. Stuart

Abstract: Abstract We describe a new species of pareid snake from the Di Linh Plateau in Lam Dong Province of southern Vietnam based on morphological and molecular evidence. Pareas temporalis sp. nov. is distinguished from its congeners by having the combination of yellow-brown body colouration; hexagonal-shaped frontal, with lateral sides parallel to the body axis; 16–17 temporals, with 4–5 anterior temporals; loreal and prefrontal not contacting eye; 2–3 preoculars; two suboculars; 2–3 postoculars; 8–9 supralabials; 8–9 infralabials; 15–15–15 dorsal scale rows, all keeled, three vertebral scale rows enlarged; 191 (+1 preventral) ventrals, smooth; 92 subcaudals, all divided; undivided anal scale; two postocular stripes; and a solid dark brown vertebral stripe extending from rear of nuchal collar along the entire length of body and tail. Phylogenetic analyses of mitochondrial DNA data recovered the new species to be nested within the P. carinatus complex and to be the sister taxon to P. nuchalis from Borneo.

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DOI: 10.3897/vz.71.e65097

Authors: Omar M. Entiauspe-Neto, Claudia Koch, Michael B. Harvey, Guarino R. Colli, Thaís B. Guedes

Abstract: Apostolepis is a diverse genus of dipsadid snakes, currently comprising 34 species occurring in most cis-Andean South America. The taxonomy of the group is highly unstable. Upon discovering its type series, we redescribe the rare species A. ambiniger (Peters, 1869) and provide an account of its geographic distribution and morphological variation in pholidosis, osteology, and hemipenial characters. We also discuss some aspects of the taxonomy of Apostolepis.

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DOI: 10.3897/vz.71.e63986

Authors: Surya Narayanan, Pratyush P. Mohapatra, Amirtha Balan, Sandeep Das, David J. Gower

Abstract: We reassess the taxonomy of the Indian endemic snake Xylophis captaini and describe a new species of Xylophis based on a type series of three specimens from the southernmost part of mainland India. Xylophis deepaki sp. nov. is most similar phenotypically to X. captaini, with which it was previously confused. The new species differs from X. captaini by having a broader, more regular and ventrally extensive off-white collar, more ventral scales (117–125 versus 102–113), and by lack of flounces on the body and proximal lobes of the hemipenis. Phylogenetic analysis of mitochondrial 16S DNA sequences strongly indicates that the new species is most closely related to X. captaini, differing from it by an uncorrected pairwise genetic distance of 4.2%. A revised key to the species of Xylophis is provided.

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