DOI: 10.3897/vz.76.e176241

Authors: Aurélien Miralles, Mark D. Scherz, Sam Hyde Roberts, Andolalao Rakotoarison, Frank Glaw, Miguel Vences

Abstract: Abstract Madascincus is a genus of quadrupedal skinks endemic to Madagascar, with 12 recognized species described between the 19th and 21st centuries, occupying diverse habitats from humid forests to arid southern regions and even high-altitude areas. Recent field expeditions uncovered two morphologically distinct forms that did not match any known species, prompting integrative taxonomic analyses that combine multilocus phylogenetics and morphology. Results revealed that each of these forms represents a highly divergent and likely ancient lineage, as evidenced by the substantial branch lengths in both mitochondrial and nuclear phylogenetic trees. These investigations led to the formal description of two new species with restricted distribution ranges: Madascincus irery sp. nov., likely endemic to the island of Nosy Be, and Madascincus minotaurus sp. nov., apparently restricted to the Tsingy de Namoroka karst system.

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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.76.e154822

Authors: Zijia Hong, M. S. Shahrul Anuar, L. Lee Grismer, Evan S. H. Quah

Abstract: Phylogenetic analyses of newly discovered populations of Hemiphyllodactylus from sky-islands across Peninsular Malaysia using the mitochondrial gene ND2, recovered two new upland species embedded within the harterti group. Hemiphyllodactylus puncak sp. nov. from Langkawi Island and H. jeraiensis sp. nov. from Gunung Jerai are sister species with an uncorrected pairwise sequence divergence of 3.6% between them. Together, they formed the sister lineage to H. cicak from Penang Hill, with an uncorrected pairwise sequence divergence of 3.9–4.8% and 4.5–5.7%, respectively. Given that these three populations occur on mountain tops isolated by lowland habitat and the Straits of Malacca, gene flow between them is highly unlikely, and there are unique combinations of characters that differentiate them from one another, as well as from all other species of the harterti group. The time-calibrated BEAST phylogeny shows that the harterti group diversified across the uplands of Peninsular Malaysia in the Oligocene and Pleistocene, which is consistent with climatic fluctuations during this period. During the Middle Oligocene, the ancestral population of the harterti group diverged into two distinct populations, one in the Banjaran Titiwangsa and another in the Banjaran Timur. These two populations then radiated independently across Peninsular Malaysia, giving rise to at least nine additional species within the harterti group.

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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.e167366

Authors: Werner Conradie, Chad Keates, Eli Greenbaum, Javier Lobón-Rovira, Krystal A. Tolley, Max Benito, Pedro Vaz Pinto, Reuben V. van Breda, Luke Verburgt

Abstract: Abstract Ichnotropis is a genus of medium-sized lacertids endemic to sub-Saharan Africa, characterised by rough head shields. The genus currently comprises six nominal species distributed across much of southern, central, and eastern Africa. Some species are apparently active at only certain times of the year, resulting in limited specimen collections and severely hampering research. This scarcity of material has historically made comprehensive systematic reviews of the genus difficult and has led to the description of numerous regional morphological variants as distinct species or subspecies. Material collected in recent years has enabled us to provide a new phylogenetic hypothesis of Ichnotropis using two mitochondrial genes (16S and ND4) and two nuclear genes (c-mos and RAG-1). Our phylogenetic dataset includes 56 individuals representing five of the six currently recognised species (excluding I. chapini). Additionally, the broad geographical sampling of the widespread I. capensis group has allowed us to explore the taxonomic status of several species and subspecies within the group. As a result, we demonstrate the monophyly of Ichnotropis in relation to other African lacertids and present the most comprehensive phylogeny of the genus to date. We also provide the first phylogenetic placements for I. tanganicana and I. grandiceps, which allows us to validate their taxonomic statuses. Furthermore, we recovered a new cryptic species closely related to I. grandiceps, and identified several well-supported clades within the I. capensis group, all corroborated by multi-locus species delimitation analyses. One of these clades is described herein as a new species, while the remaining taxa of interest are discussed and highlighted for future investigation. Based on our findings, we recommend the following taxonomic revisions: Ichnotropis longipes and I. macrolepidota should remain synonyms of I. capensis; I. bivittata pallida and I. capensis nigrescens are treated as a junior synonyms of I. bivittata; and I. overlaeti is considered a junior synonym of I. tanganicana. Although we could not determine the phylogenetic placement of I. chapini due to the lack of genetic material, its head morphology and scalation support its reassignment to the I. bivittata group. Thus, it is retained as a valid species pending the availability of new material for further taxonomic actions. In conclusion, this study resolves several long-standing taxonomic issues within one of Africa’s most understudied lacertid genera and lays a solid foundation for future research on the genus Ichnotropis.

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DOI: 10.3897/vz.75.e162650

Authors: Amirtha Balan, Abhijit Das, Bitupan Boruah, Frank Tillack, Samuel Lalronunga, Veerappan Deepak

Abstract: Abstract The genus Ptyctolaemus Peters, 1864 is currently represented by three species; P. gularis, P. collicristatus and P. chindwinensis. Previous molecular phylogenetic studies have highlighted additional lineages within P. gularis. Phylogenetic analysis using the ND2 mitochondrial gene recovered three distinct lineages in India. Multivariate analyses using morphological data placed the type specimen of P. gularis (ZMB 5004) with the samples collected from Meghalaya. We assign this lineage as P. gularis and describe the other two lineages as new species. Ptyctolaemus siangensis sp. nov. found west of Siang River in Arunachal Pradesh and Ptyctolaemus namdaphaensis sp. nov. found in Namdapha, Changlang district of Arunachal Pradesh. Although phenotypically similar the two new species differ from its congeners in gular colouration and subtle morphological differences.

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DOI: 10.3897/vz.75.e162071

Authors: Carlos M. Marín, Daniel Bocanumenth, Juan M. Daza

Abstract: Abstract The lizard genus Anolis is the second most diverse genus of terrestrial vertebrates. Within Anolis, the highly speciose clade Dactyloa comprises six species series, including the latifrons series. Despite previous efforts to reconstruct its phylogeny, earlier studies have excluded a substantial proportion of the clade’s species diversity. Here, we integrated both historical and newly generated genetic data to reconstruct the most comprehensive molecular phylogeny of the latifrons series to date including 88% of the current species diversity. We also conducted a thorough morphological examination of museum specimens representing ten species, primarily distributed in Colombia, including vouchers of A. danieli from the Central Cordillera used in previous molecular phylogenies. Our phylogeny also included genetic samples of A. danieli from several localities in the Western Cordillera, samples of Anolis limon and A. mirus (two species previously lacking genetic data), and sequences from the Central American species A. kathydayae and A. brooksi. Our results recovered topological differences for A. limon and A. mirus compared to previous hypotheses and revealed that specimens assigned to A. danieli in earlier studies were misidentified and are not phylogenetically related to this species. Instead, our results showed that the true A. danieli is sister to a green anole clade distributed across the Central Cordillera, Pacific region, and Panama. Based on our phylogenetic and genetic distance analyses, we conclude that A. kathydayae should be considered a junior synonym of A. brooksi. Lastly, we describe the taxon previously confused with A. danieli and comment on the taxonomic implications of our findings.

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DOI: 10.3897/vz.75.e153514

Authors: François S. Becker, Graham J. Alexander, Krystal A. Tolley

Abstract: Abstract Barking geckos, Ptenopus Gray, 1866 are burrowing geckos that occur across the xeric regions of southern Africa. They possess unique vocal abilities, with males producing loud advertisement calls to attract females. The taxonomy of the genus has remained stable for six decades, with three recognised species: Ptenopus garrulus (Smith, 1849), P. kochi Haacke, 1964, and P. carpi Brain, 1962. Within P. garrulus, two subspecies have been recognised since 1935: the nominotypical form (P. g. garrulus) and P. g. maculatus Gray, 1866. A recent phylogenetic analysis of the genus found that it contains eight to ten putative species. We used an integrated taxonomic approach to delimit a total of nine species, including evidence from phylogenetics, ecology, calls, and morphology. Ptenopus g. maculatus is elevated to full species, thereby restricting the geographic range of P. garrulus sensu stricto to the greater Kalahari. Additionally, four new species are named which were previously included in ‘P. g. maculatus’: Ptenopus adamanteus sp. nov. from the southern Namib Desert, P. circumsyrticus sp. nov. from the central Namib Desert, P. kenkenses sp. nov. from the northern Nama Karoo, and P. australis sp. nov. from southern Nama Karoo. As a result, the range of P. maculatus sensu stricto is restricted to the central northern Namib Desert. Furthermore, one new species previously included in P. carpi is named P. sceletus sp. nov. from the Skeleton Coast (northern coastal Namib Desert), thereby restricting the range of P. carpi sensu stricto to a small strip of coastal Namib Desert between the Swakop and Kuiseb rivers. The Namib Desert is the centre of diversity for the genus Ptenopus, containing seven of the nine species including the oldest divergent lineages. Two species-level keys are provided: a morphological key and a unique bioacoustic key to the advertisement calls.

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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.75.e152054

Authors: Javier Torres, Dexter Reilly, Claudia Nuñez-Penichet, R. Graham Reynolds, Richard E. Glor

Abstract: Abstract Cuba is the only landmass with more than one species in the Anolis carolinensis subgroup. We test the hypothesis that three rather than two distinct species occur on Cuba, based on substantial prior evidence of paraphyly. To test this hypothesis, we collected phenotypic data from all described species in the subgroup, including eastern and west-central Cuban populations of A. porcatus, and assessed phenotypic diagnosability using uni- and multivariate analyses. We also examined geographic isolation using all available occurrence records for Cuban lineages. Additionally, we conducted ecological niche modeling and niche overlap analyses, considering only Cuban lineages, to test for ecological differentiation. Finally, we reconstructed phylogenetic trees, incorporating all species from the subgroup for the first time. Our results support the recognition of three species in Cuba: A. allisoni and eastern and west-central A. porcatus as two distinct cryptic species, showing minimal phenotypic differentiation but clear geographic isolation, distinct ecological niches, and deep genetic divergence. We restrict the name A. porcatus to west-central Cuba, with Havana as the type locality, and formally describe the eastern Cuban populations as Anolis torresfundorai sp. nov., designating Baracoa, Guantánamo, as the type locality.

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DOI: 10.3897/vz.74.e128775

Authors: Peter J. McDonald, Aaron L. Fenner, Janne Torkkola, Paul M. Oliver

Abstract: Abstract Stable upland habitats in arid zone biomes are often characterised by locally endemic lineages. Explanations for this pattern include habitat or substrate specialisation (ecological specialisation) or intensifying aridity driving retreat into climatically buffered habitats (climatic refugia). Here we present an analysis of these alternative models using genetic, morphological and climate data for Diplodactylus galeatus, a gecko from central Australia that occurs in a series of isolated populations associated with dissected tablelands and mountain ranges. Analyses of mtDNA and SNP data support four distinct lineages, and dating analyses suggest divergence through the Pliocene. Morphological data show slight differences across lineages. Investigation of climate niche shows that two lineages are restricted to areas more arid than the intervening uninhabited region. These data suggest that specialisation to rocky substrates, potentially with subsequent paleo-erosion of dissected tablelands after a Pliocene wet pulse, was the key driver of divergence in this clade. Based on their deep genetic divergence, and differences in morphology and pattern, we recognise two isolated populations as new species.

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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.e130147

Authors: Omar Torres-Carvajal, Camila Sandoval, Diego A. Paucar

Abstract: The taxonomic status of the skinks from Ecuador has never been carefully addressed. In this paper we examine populations of Mabuya lizards across Amazonian Ecuador in an attempt to establish their taxonomic identity and phylogenetic affinities. We confirm the presence of both M. altamazonica and M. nigropunctata and describe a new species from Yasuní National Park, one of the most biodiverse places on Earth. The new species differs from its congeners in lepidosis and color patterns. For the first time, we include samples from Ecuador in a molecular phylogenetic analysis of Mabuya, which confirms the monophyly of the new species and the taxonomic identity of both M. altamazonica and M. nigropunctata from Ecuador. The new species is closely related to M. bistriata. Finally, we present an identification key for species of Ecuadorian Mabuya.

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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.e124752

Authors: Bitupan Boruah, Surya Narayanan, Neelavar Ananthram Aravind, Samuel Lalronunga, V. Deepak, Abhijit Das

Abstract: Abstract We describe six new species of Cyrtodactylus from the khasiensis group using morphological characteristics, supported by the molecular analyses based on the mitochondrial ND2 gene. We used four different molecular species delimitation analyses that recovered six distinct undescribed lineages distributed across four states in northeastern India. Our phylogenetic analyses using ML and Bayesian approaches recovered a clade where the recently described C. arunachalensis and C. cayuensis align together with our other samples from Arunachal Pradesh, north of Brahmaputra River. Based on these results and overlapping morphological characteristics we synonymize C. arunachalensis with C. cayuensis. We provide updated comparative morphological characters for species in the khasiensis group and where available these characters are tabulated for males and females separately. Including the six new species the khasiensis group now contains 35 species, of which 26 are endemic to India.

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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.e109088

Authors: Kai Wang, V. Deepak, Abhijit Das, L. Lee Grismer, Shuo Liu, Jing Che

Abstract: Abstract Owing to the harsh terrain, few biodiversity surveys have been carried out in the Pan-Himalaya Region. Among the understudied taxa from this region, Jerdon’s forest lizard, Calotes jerdoni, is believed to have a wide distribution, from northeast India to southwestern China. However, given the heterogeneous environment across its range and the lack of studies on this species, its taxonomy remains questionable. Using integrative taxonomic methods, we combined both morphological and genetic data from the type and topotypic specimens and examined the current taxonomic hypothesis of C. jerdoni across its range. Molecular data reveal that C. jerdoni as currently recognized, contains three deeply diverged lineages: one from the type locality in Northeast India, one from Western Myanmar, and another one from Southwestern China. The uncorrected genetic distances of mitochondrial coding gene ND2 among these three clades ranged over 10%. The Chinese population is sister to C. medogensis and paraphyletic to the remaining two clades of C. jerdoni. Morphological analyses confirm the results of the molecular analyses, where the Myanmar and Chinese populations can be diagnosed statistically in both univariate and multivariate space from the true C. jerdoni, as well as by a suite of reliable categorical morphological characters, including the size and shape of gular scales and ventral scales. To resolve the current taxonomic confusion, we resurrect the junior synonym, C. yunnanensis, for the Chinese population and expand its distribution to Myanmar, redescribe the elusive C. maria and C. medogensis based on its type material, and describe the remaining western Myanmar population as a new species. We further discuss the possibility of additional cryptic species within the complex in the Pan-Himalaya Region and provide a diagnostic key to all recognized members of the C. jerdoni complex.

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DOI: 10.3897/vz.74.e113084

Authors: Sandeep Das, Saunak Pal, Surya Narayanan, K. Subin, Muhamed Jafer Palot, K. P. Rajkumar, V. Deepak

Abstract: Abstract We describe a new species of Agasthyagama from Idukki district, Kerala, India. The new species is distinguished from its sister species Agasthyagama beddomii by a combination of scale characters and are also genetically different from each other with 11% uncorrected pairwise difference in ND2 gene and 3% in 16S gene. The two species are also geographically separated, the closest distributional records are approximately 80 km apart.

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DOI: 10.3897/vz.74.e112782

Authors: Alex Slavenko, Stephen J. Richards, Stephen C. Donnellan, Allen Allison, Paul M. Oliver

Abstract: Abstract Skinks are the most diverse component of the reptile fauna in the mountains of New Guinea and many seemingly specialised high-elevation species remain undescribed. Here we describe two spectacular new gold-patterned skinks in the montane-specialist genus Papuascincus. Both species can be diagnosed from all congeners by their distinctive colouration, in addition to aspects of scalation and body size. One new species is mainly recorded from lower montane forest in karst habitats spanning more than five hundred kilometres along the southern edge of New Guinea’s Central Cordillera and is likely to warrant an IUCN conservation status of Least Concern. The second new species has thus far only been recorded from cloud forest on the summit of Mt. Menawa in the North Coastal Ranges and we suggest it should be considered Data Deficient. However, if further survey work confirms a restricted distribution with little scope for upslope elevational retreat under future warming climates it will likely qualify for Endangered or Critically Endangered status.

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DOI: 10.3897/vz.74.e113899

Authors: Hao-Tian Wang, Shuo Qi, Dan-Yang Zhou, Ying-Yong Wang

Abstract: Abstract A new species of the genus Gekko Laurenti, 1768, Gekko paucituberculatus sp. nov., is described here, based on two specimens from Tianyang District, Baise City, Guangxi Zhuang Autonomous Region, China. It was placed in subgenus Japonigekko on the basis of morphological and molecular phylogenetic analysis and can be distinguished from all congeners in this subgenus by significant divergences in the mitochondrial 16S and ND2 genes and by a combination of the following morphological characters: Moderate body size, SVL 77.2 mm in the adult male and SVL 85.9 mm in the adult female; tubercles only present along dorsolateral trunk and absent on other regions; fingers and toes with weak webbing; continuous precloacal pores 12 in the male, absent in the female; a single postcloacal tubercle on each side; a light-coloured vertebral line from nape to tip of tail; dorsum greyish-brown, with 7–8 dirty-white bands between nape and sacrum. Meanwhile, the distribution of G. palmatus in China has been confirmed as occurring in Guangxi and Guangdong Provinces. This study brings the total species of the subgenus Japonigekko in China to 19.

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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.74.e110674

Authors: Ishan Agarwal, Tejas Thackeray, Akshay Khandekar

Abstract: Abstract Subdoluseps is a recently described genus of Lygosomine skinks distributed in peninsular India and Southeast Asia. We conduct the first revision of Indian Subdoluseps based on range-wide sampling including 89 specimens from 33 localities. We use two mitochondrial and three nuclear markers, 58 morphological characters, and ecological data to reconstruct the evolutionary history of Indian Subdoluseps and assess their diversity and distribution, providing insights into lygosominin biogeography. We formally describe the Indian clade as a new genus, Dravidoseps gen. nov. and name five new species from Tamil Nadu, India in an integrative taxonomic framework – D. gingeeensis sp. nov., D. jawadhuensis sp. nov., D. kalakadensis sp. nov., D. srivilliputhurensis sp. nov., and D. tamilnaduensis sp. nov.. We transfer Riopa goaensis, Subdoluseps pruthi and S. nilgiriensis to the new genus and designate neotypes for the former two. Members of Dravidoseps gen. nov. are the first known viviparous skinks from peninsular India and the only known viviparous lygosominins apart from a few species of east African Mochlus. The Lygosomini have a Southeast Asian origin and began diversifying in the Eocene with three dispersals between India and Southeast Asia. Species level diversification in Dravidoseps gen. nov. was likely driven by a combination of niche conservatism, paleoclimate and past forest distribution. The discovery of a new genus and five new species reiterates the high levels of diversity and endemism present in peninsular India and how much more remains to be discovered.

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DOI: 10.3897/vz.73.e110626

Authors: Karin Tamar, Marek Uvizl, Mohammed Shobrak, Mohammed Almutairi, Salem Busais, Al Faqih Ali Salim, Raed Hamoud M. AlGethami, Abdulaziz Raqi AlGethami, Abdulkarim Saleh K. Alanazi, Saad Dasman Alsubaie, Laurent Chirio, Salvador Carranza, Jiří Šmíd

Abstract: Abstract A recent molecular phylogeny of the agamid genus Pseudotrapelus, distributed in the rocky areas of North Africa and the Arabian Peninsula, revealed the presence of a genetically distinct lineage around the city of Riyadh in central Saudi Arabia. With the inclusion of additional specimens, we were able to describe this lineage as a new species, P. tuwaiqensis sp. nov., confined to the Tuwaiq Escarpment, thus endemic to central Saudi Arabia. Our results of morphological examinations and molecular analyses, using three mitochondrial (COI, 16S, ND4-tRNAs) and two nuclear (c-mos, MC1R) gene fragments, show the new species is genetically differentiated and phylogenetically close to P. sinaitus and P. chlodnickii.

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DOI: 10.3897/vz.73.e110512

Authors: Akshay Khandekar, Tejas Thackeray, Ayuthavel Kalaimani, Ishan Agarwal

Abstract: Abstract We describe two new small-bodied, sympatric species of south Asian Cnemaspis belonging to the mysoriensis + adii clade from the Gingee Hills in Tamil Nadu, peninsular India. The two new species can be easily distinguished from the other eight described members of the mysoriensis + adii clade by their dorsal pholidosis, the configuration of femoral and precloacal pores in males, a number of meristic characters and subtle differences in colouration, beside 6.7–20.8 % uncorrected pairwise ND2 sequence divergence. The two species represent different ecomorphs, one a stouter, microhabitat generalist and the other a more slender, elongate rock specialist. The discovery of two new species from granite boulder habitats and Tropical Dry Evergreen Forests is indicative of the importance of these areas for biodiversity. It is likely that similar rocky habitats across southern peninsular India will harbour many more undescribed species.

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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.e95639

Authors: Michael F. Bates, Javier Lobón-Rovira, Edward L. Stanley, William R. Branch, Pedro Vaz Pinto

Abstract: Cordylus angolensis (Bocage, 1895) was described 128 years ago on the basis of a single specimen collected at Caconda in the west-central highlands of Angola. Additional specimens referred to this species were collected at ‘Mombolo’ (also in the central highlands) during the Vernay Angola Expedition in 1925. As the holotype was apparently destroyed in the fire of 1978 at the Museu Bocage in Lisbon and no additional specimens have been collected, its taxonomic status and phylogenetic relationships has remained uncertain. The species has eluded all efforts aimed at its re-discovery in the vicinity of the type locality, with a single specimen from near Condé, north of Mombolo—collected in 1970 by Wulf Haacke—the only other specimen of Cordylus known from west-central Angola. Recent field work in the Angolan highlands resulted in the collection of a series of specimens from Taqueta Mountain (west of Caconda), Monte Verde (Sandula, ‘Mombolo’) and Uassamba (Vondo). A phylogenetic analysis, using three mitochondrial and six nuclear genes, indicated the existence of two distinct species-level lineages in the Angolan highlands. These two species are allopatric and morphologically distinct, differing especially in terms of their colour patterns, eye colour and certain scalation characteristics. We therefore confirm that C. angolensis is a valid species and designate a neotype, and describe a new species, Cordylus momboloensis sp. nov.

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DOI: 10.3897/vz.73.e104494

Authors: Surya Narayanan, Peter Christopher, Kothandapani Raman, Nilanjan Mukherjee, Ponmudi Prabhu, Maniezhilan Lenin, Sivangnanaboopathidoss Vimalraj, V. Deepak

Abstract: A new large-bodied (SVL 101–109 mm) gecko of the genus Hemidactylus is described from the Gingee Hills in the Eastern Ghats of India. The new species is closely related to H. graniticolus and the recently described H. easai, from which it can be distinguished by its lower femoral pores count. The new species described here was previously identified as H. cf. graniticolus based only on the molecular data, pending its formal description. Our findings were consistent with the results from the molecular DNA analyses, showing that this population is morphologically distinct from other closely related species. As a result, we formally describe this lineage as a new species, providing a comprehensive description of its morphological characteristics based on a type series of five specimens and compare it with its congenerics.

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DOI: 10.3897/vz.73.e102602

Authors: Ishan Agarwal, Tejas Thackeray, Akshay Khandekar

Abstract: We describe two new spotted species of ground-dwelling gecko of the genus Cyrtodactylus (Geckoella) from southeastern India in an integrative taxonomic framework. The new species are recovered as sister taxa within the C. collegalensis species complex, with 13.0–16.7% uncorrected mitochondrial sequence divergence from the other eight members of the C. collegalensis complex and 10.4% from one another. The new species are morphologically diagnosed by a spotted dorsal pattern of four pairs of spots (occasionally fused into figure 8-shaped markings) from the banded species C. aravindi, C. speciosus, C. rishivalleyensis and C. yakhuna; and from the spotted species with three or fewer pairs of spots in C. collegalensis and C. srilekhae; and from C. chengodumalaensis by the absence of any enlarged dorsal scales and from C. varadgirii by the absence of a patch of enlarged roughly hexagonal scales on the canthus rostralis and beneath the angle of the lower jaw. The two new species can only be differentiated from each other based on slight differences in body size, relative body width and other statistically significant, size-corrected morphometric characters. These are among the first endemic lizards from Tropical Dry Evergreen habitats along the southeast coast of India.

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DOI: 10.3897/vz.73.e101871

Authors: Akshay Khandekar, Tejas Thackeray, Rameshwaran Mariappan, Satpal Gangalmale, Vivek Waghe, Swapnil Pawar, Ishan Agarwal

Abstract: We describe a new species of Hemidactylus based on an integrative taxonomic framework from scrub habitats at the southern tip of India, in Thoothukudi District, Tamil Nadu. The new species has the most densely packed tubercles among Indian Hemidactylus, almost resembling the most tuberculate Indian Cyrtopodion. Hemidactylus quartziticolus sp. nov. is phylogenetically placed within the brookii group of Indian Hemidactylus, where it is sister to the H. gleadowi complex from western-central India. The new species is 14.5–23.7% divergent in ND2 mitochondrial sequence data from other brookii group members, and can be easily diagnosed from regional congeners by its unique dorsal scalation, the number and arrangement of precloacal-femoral pores, the number of dorsal tubercle rows at midbody, number of lamellae under digit I and IV of manus and pes. The new species is currently known only from two isolated, low quartzite hillocks 45 km apart with scrubby, thorn forests and loose, stony soil.

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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.e90979

Authors: Suranjan Karunarathna, Kanishka D. B. Ukuwela, Anslem De Silva, Aaron M. Bauer, Majintha Madawala, Nikolay A. Poyarkov, Madhava Botejue, Dinesh Gabadage, L. Lee Grismer, Vladislav A. Gorin

Abstract: Sri Lanka is a local hotspot for Cnemaspis day geckos with 40 currently known species with 100% endemism. In this paper, we evaluate the phylogenetic relationships of Cnemaspis species belonging to the alwisi group of the podihuna clade and describe two additional new species of Cnemaspis from Sri Lanka; one from Galgiriya mountain, Kurunegala District, and another from Ethagala mountain, Ampara District. These new species were recorded from granite caves within forested areas in isolated mountains in the dry bioclimatic zone (point-endemics). Both new species are microhabitat specialists with narrow niches limited to humid, cool, canopy-shaded granite caves and old buildings associated with granite caves, where they are camouflaged by their cryptic morphology and body colouration. Furthermore, both species prefer narrow (~ 6–12 mm), long (~ 120–450 mm) and deep (~ 80–260 mm) crevices as refugia. The regions in which these habitats are located receive relatively low annual rainfall (1,000–1,500 mm). These new species are medium in size (28.5–36.8 mm SVL) and can be differentiated from all other Sri Lankan Cnemaspis by the presence of clearly enlarged, subhexagonal subcaudal scales and the absence of precloacal pores in males. Both species described here are categorised herein as Critically Endangered (CR) under the IUCN Red List criteria. The major threats for these new species are habitat loss due to expansion of commercial-scale agriculture, illicit forest encroachments, and forest fires. Therefore, we recommend that relevant authorities take immediate conservation action to ensure the protection of these forest areas with their buffer zones in the near future.

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DOI: 10.3897/vz.73.e101311

Authors: Surya Narayanan, Saunak Pal, L. Lee Grismer, N. A. Aravind

Abstract: We describe a new species of rupicolous Cnemaspis from the Male Mahadeshwara Wildlife Sanctuary in the Eastern Ghats. The current description is based on a type series of eight specimens that bear a unique combination of morphological and colour pattern characteristics that do not occur in other closely related species. Maximum likelihood and Bayesian analyses based on the mitochondrial gene ND2 recovered the new species to be nested within the gracilis clade and the sister species to the recently described C. agarwali from the Eastern Ghats.

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DOI: 10.3897/vz.73.e94265

Authors: Rafael A. Moreno-Arias, Miguel A. Méndez-Galeano, Iván Beltrán, Mario Vargas-Ramírez

Abstract: The Anolis heterodermus group comprises eight big-headed and short-legged lizard species from the highlands of the northernmost South American Andes. Recent studies revealed unknown lineages within this group that had previously been categorized as a species complex. By widely sampling and applying an integrative taxonomic framework, we (1) assessed the species diversity of the group using a molecular dataset (two mitochondrial and one nuclear markers) along with an inclusive morphological study (scalation, scale configuration and ornamentation, morphometrics, and dewlap and body colour patterns); and (2) we inferred the evolutionary relationships within this species group. Our analyses confirmed the formerly reported differentiation between populations of those high-altitude lizards, and we identified several unknown evolutionary lineages. Our results provided evidence for the existence of nine distinct, independently evolving evolutionary lineages in the heterodermus group. As a result, we described two morphologically and genetically highly distinct lineages as species new to science (A. quimbaya sp. nov. and A. tequendama sp. nov.). We redescribed A. heterodermus and erected as a valid species Anolis richteri, a previously described synonym of A. heterodermus. A taxonomic key for the identification of species of the Phenacosaurus clade was presented. The identification of two additional poorly-known lineages suggested that the diversity of this group of lizards is still unknown; therefore, it is necessary to establish measurements for the group´s conservation, as well as to perform fieldwork and revision of herpetological collections to identify possible hidden diversity within the group.

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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.73.e94004

Authors: Abraham Lozano, Jack W. Sites Jr, Aurelio Ramírez-Bautista, Jonathon C. Marshall, Numa P. Pavón, Raciel Cruz-Elizalde

Abstract: Sexual dimorphism is a widespread feature in the Animal Kingdom. In lizards of the Sceloporus grammicus complex, studies of sexual dimorphism that analyze the allometric trajectories of body traits remain unexplored. Here we investigate sexual dimorphism in key phenotypic traits, including body size (snout-vent length, SVL) as well as head length (HL), head width (HW), and forearm length (FL). We use an allometric approach to detect differences in scale relationships among body parts in the S. grammicus complex in Mexico. We focus on two chromosomal races within this complex, F5 (2n = 34) and FM2 (2n = 46). In the complex, we found that males are larger than females in all morphological variables, and this pattern was confirmed in both races. We determined negative allometric trajectories (SVL vs. HL and HW), isometry (SVL vs. FL) and intersexual differences in the slopes of the SVL vs. HL and HW; the males showed steeper slopes. Thus, the growth of the head is more pronounced in males than females. Additionally, we found between-race differences in these trajectories (SVL vs. FL) and in all morphological variables (F5 lizards are larger than those of the FM2 race), which correlate with their chromosomal divergence. We discuss biological implications of our findings in relation to sexual selection and natural selection.

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DOI: 10.3897/vz.72.e94799

Authors: Ishan Agarwal, Tejas Thackeray, Akshay Khandekar

Abstract: South Asian Cnemaspis are one of the most diverse clades of gekkonids in South Asia with their highest diversity in the Western Ghats and Sri Lanka. These geckos include only a few nocturnal species and are largely diurnal or cathemeral and restricted to relatively cool habitats. One of the prominently diurnal subgroups in South Asian Cnemaspis is the bangara clade, which includes six species distributed in southern India on the eastern slopes of the Western Ghats, the southern Eastern Ghats and Palghat Gap. In this paper, we describe five more species of the bangara clade from the Shevaroyan landscape, including three from Kollimalai and one each from Yercaud and Pachaimalai, all in Tamil Nadu. These new species show 4.6–19.7 % uncorrected sequence divergence on the mitochondrial ND2 gene from each other and known species of the bangara clade and are morphologically diagnosable in body size, the number of paravertebral tubercles between limb insertions, the number of dorsal tubercle rows, the number of ventral scale rows across the belly, the number of femoral and precloacal pores and poreless scales separating these series, and aspects of colouration. The discovery of these five new species adds to the growing discoveries of cool-adapted species in southern India outside the Western Ghats and highlights the role of sky-islands in diversification. The Shevaroyan landscape shows high levels of microendemism with eight species distributed in an area of < 2000 km2, and all these species restricted to much smaller areas of actual distribution. With an area of < 500 km2 respectively, the massif of Pachaimalai has a single endemic and the massifs of Yercaud and Kollimalai have three endemic Cnemaspis species each.

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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.e89324

Authors: Surya Narayanan, Aravind NA

Abstract: We describe a new species of small-sized (SVL 24.6–28.8 mm) rupicolous Cnemaspis from the Biligirirangan Hills (BR Hills), Karnataka based on morphology and molecular (mitochondrial 16S and ND2) data. The new species is nested within the monticola, mysoriensis and gracilis clades, where it is basal to monticola and gracilis clades in the Maximum Likelihood analysis and is basal to the monticola clade in the Bayesian phylogenetic analysis.

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DOI: 10.3897/vz.72.e85269

Authors: Mariana P. Marques, Luis M. P. Ceríaco, Matthew P. Heinicke, Rachal M. Chehouri, Werner Conradie, Krystal A. Tolley, Aaron M. Bauer

Abstract: The genus Heliobolus comprises four recognized species, all endemic to sub-Saharan Africa. Of these, only Heliobolus lugubris occurs in southern Africa, its distribution extending from Angola in the west to Mozambique in the east and reaching as far south as parts of northern South Africa. Like many of the reptile species that occur in southern Africa, Heliobolus lugubris is poorly studied, and preliminary investigation suggested that it may contain cryptic diversity. The present work focusses on the Angolan population of H. lugubris and uses an integrative taxonomic approach based on morphological, coloration and DNA sequence data. The results indicate that some of the current and historical specimens of H. lugubris from Angola do not correspond to the nominotypical form, and that differences between specimens suggest the presence of two additional species, described here as Heliobolus bivari sp. nov. from the southernmost xeric/desertic regions and plateau of Namibe Province, southwestern Angola and H. crawfordi sp. nov. from the Serra da Neve inselberg north through the sub-desert coastal regions of northern Namibe, Benguela, and Kwanza Sul provinces. Nominotypical Heliobolus lugubris is confirmed to occur in Cuando Cubango Province, southeastern Angola.

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DOI: 10.3897/vz.72.e89660

Authors: Surya Narayanan, Sandeep Das, Amirtha Balan, Roshin Tom, Nitin Divakar, Rajkumar Kp, P. Hopeland, V. Deepak

Abstract: A new species of Cyrtodactylus is described from the southern Western Ghats of India. It is distinguished from all species of the ­Cyrtodactylus collegalensis species complex in colour pattern and resembles the Srilankan endemic C. yakhuna in overall colouration. Phylogenetic analysis of mitochondrial ND2 DNA sequences indicates that the new species is most closely related to species in the C. collegalensis complex and differs from them by an uncorrected pairwise genetic distance of 10.5–12.9%.

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DOI: 10.3897/vz.72.e78828

Authors: Lucas Rafael Uchôa, Fagner Ribeiro Delfim, Daniel Oliveira Mesquita, Guarino Rinaldi Colli, Adrian Antonio Garda, Thaís B. Guedes

Abstract: Abstract The Caatinga is the largest seasonal dry tropical forest in South America and it has been historically neglected in terms of its biodiversity. Regarding lizards, different studies led to the current knowledge of diversity and endemism in Caatinga, but detailed syntheses are scarce in the literature. We present the most detailed and up-to-date synthesis of knowledge about Caatinga lizards by providing a detailed (i) list of species; (ii) taxonomic richness patterns; (iii) knowledge gaps and spatial biases; and (iv) detailed distribution maps of all species that contain at least one occurrence record within Caatinga limits. We created a distribution database using occurrences of lizards in Caatinga based on scientific collections, field collection, and literature. We produce up-to-date distribution maps, calculate the Extent of Occurrence and provide the environmental and bioclimatic profile for each species recorded. We draw taxonomic richness and sampling gap maps. Our database has 20,538 records of occurrence of lizards of the Caatinga. We recorded 93 lizard species (13 families), 52.7% of which are endemic. Forthy-four percent of the species present restricted distributions. We identified that 53% of the Caatinga area (or 70% of the municipalities) has no record of occurrence of lizards. The data presented are an important step towards synthesizing in detail the accumulated knowledge about Caatinga lizards and is crucial for accurate strategies for the conservation planning. It directs actions to advance our knowledge on Caatinga lizards: to concentrate inventories in sample void areas; continuous update of the species occurrence database, advance in the generation of autoecology data for species.

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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.72.e82343

Authors: Akshay Khandekar, Tejas Thackeray, Ishan Agarwal

Abstract: We describe three distinct, small-bodied, scansorial species of south Asian Cnemaspis from Kalakad Mundanthurai Tiger Reserve, Tirunelveli district, Tamil Nadu, India—Cnemaspis azhagu sp. nov. from Thirukurungudi forest range, Cnemaspis mundanthuraiensis sp. nov. from Mundanthurai forest range and Cnemaspis kalakadensis sp. nov. from Kalakad forest range. Phylogenetic analyses using a partial sequence of the mitochondrial ND2 gene and general morphology places each of the three new species in the beddomei, gracilis and littoralis clades, respectively. The three new species are diagnosed from all other described members of their respective clades by a suite of differing morphological characters including snout vent length, number of dorsal tubercle rows at mid-body, number of paravertebral tubercles, presence or absence of spine-like scales on flanks, number of ventral scales across belly at mid-body, number of ventral scales from mental to anterior border of cloaca, number of lamellae under digit IV of pes, number of femoral and/or precloacal pores and poreless scales separating these series, as well as subtle colouration differences. We also provide some novel characters of tail tuberculation of the three new species described herein. With the discovery of these three new species, eight species of geckos including five Cnemaspis are now known to be endemic to KMTR.

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DOI: 10.3897/vz.72.e82201

Authors: Anh The Nguyen, Tang Van Duong, Perry L. Wood Jr., L. Lee Grismer

Abstract: An integrative taxonomic analysis of species in the colubrid genus Lycodon Fitzinger, 1826 recovered two new syntopic species of the L. rufozonatus complex from the imperiled Song Giang River valley in Khan Hoa Province, of Southern Vietnam. Although L. truongi sp. nov. and L. anakradaya sp. nov. are syntopic, they are not particularly closely related and can be differentiated from each other and all other species in the L. rufozonatus complex on the basis of meristics, morphometrics, color pattern, and uncorrected pairwise genetic distance based on the mitochondrial gene cytochrome b. The discovery of these two new range-restricted species and a previously described range-restricted gekkonid in the genus Cyrtodactylus Gray, 1828 from the same valley, underscores the necessity of continued field work in the Song Giang River valley so as to catalog the unrealized herpetological diversity in this area and establish research-based conservation programs.

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DOI: 10.3897/vz.72.e80615

Authors: L. Lee Grismer, Attapol Rujirawan, Siriporn Yodthong, Bryan L. Stuart, Minh Duc Le, Dzung Trung Le, Yodchaiy Chuaynkern, Perry L. Wood, Jr., Anchalee Aowphol

Abstract: Abstract Convergent morphological specializations for an arboreal lifestyle in most species of the Cyrtodactylus brevipalmatus group have been a confounding factor for establishing a stable taxonomy among its species. Recent references to C. interdigitalis from throughout Thailand and Laos were made without comparisons to the type material from Tham Yai Nam Nao, Nam Nao National Park, Phetchabun Province, Thailand, but instead, were based on general morphological similarity and distribution. The taxonomy of C. interdigitalis is stabilized here by comparing the paratypes to other specimens from Thailand and Laos and recovering their phylogenetic relationships based on newly acquired genetic data, including those from the type locality. The phylogeny recovered all specimens outside the type locality to be either C. ngati from Vietnam or new species closely related to C. ngati. Cyrtodactylus interdigitalis is shown here to be a range-restricted upland endemic on the Phetchabun massif of northern Thailand. The phylogeny also indicates that C. ngati extends hundreds of kilometers farther south into northern Thailand and central Laos. We hypothesize that the significant morphological divergence in body shape of the types of C. ngati, compared to that of the Lao and Thai populations, may be due to local adaptions for utilizing karst (C. ngati) rather than vegetation (Lao and Thai populations). Additionally, phylogenetic and multivariate analyses identified a potentially new species from Phu Hin Rong Kla National Park, Phitsanulok Province, in northern Thailand and another from the Khlong Naka Wildlife Sanctuary, Ranong Province, in southern Thailand. A series of newly examined specimens from Kaeng Krachan National Park, Phetchaburi Province, Thailand represents a possible ~82 km range extension to the southeast of C. rukhadeva. This research continues to underscore the high diversity of range-restricted upland endemics in Thailand and the importance of examining type material (if possible) in the context of a phylogeny so as to construct proper taxonomies that reveal, rather than obscure, diversity.

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DOI: 10.3897/vz.72.e76308

Authors: Akshay Khandekar, Tejas Thackeray, Ishan Agarwal

Abstract: We describe three new small-bodied, cryptic species of south Asian Cnemaspis belonging to the mysoriensis and goaensis clades from the Mysore Plateau and the Western Ghats biodiversity hotspot in Karnataka, peninsular India; and provide a key to members of each clade, besides providing the first ND2 sequence data for C. ranganaensis. Cnemaspis tigris sp. nov. from Kaiwara in Karnataka is a member of the mysoriensis clade and can be morphologically distinguished from all six described members of the clade in a number of meristic characters and subtle differences in colouration, beside ≥ 12.1–23.4 % uncorrected pairwise ND2 sequence divergence. Cnemaspis sakleshpurensis sp. nov. from Sakleshpur and C. vijayae sp. nov. from Coorg, both in the Western Ghats of Karnataka, are members of the goaensis clade and are easily diagnosed from all three described members of the clade in meristic characters beside 5.2–14.8 % divergence from described members of the clade and 14.6 % from each other in uncorrected pairwise ND2 sequence divergence. The discovery of these new species from two different clades and biogeographic regions is not surprising, given the steep rise in the number of species of Cnemaspis known from peninsular India. Comprehensive geographic sampling in conjunction with molecular and morphological data is essential to understand the true diversity and distributional ranges of species within this ancient clade of gekkonid lizards.

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DOI: 10.3897/vz.72.e76046

Authors: Sandeep Das, Saunak Pal, Sasidharan Siddharth, Muhamed Jafer Palot, Veerappan Deepak, Surya Narayanan

Abstract: Abstract A new large (94–107mm SVL) species of gecko of the genus Hemidactylus is described from the drier parts of the Western Ghats of India. The new species is closely related to H. graniticolus, from which it can be distinguished based on dorsal pholidosis at mid-body, the structure of tubercles on the dorsum, dorsal pholidosis on the tail. The new species is also 6.6–7.2% divergent from H. graniticolus in the ND2 mitochondrial gene.

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DOI: 10.3897/vz.72.e77702

Authors: L. Lee Grismer, Lelani del Pinto, Evan S. H. Quah, Shahrul Anuar, Michael Cota, Jimmy A. McGuire, Djoko T. Iskandar, Perry L. Wood Jr, Jesse L. Grismer

Abstract: Abstract An integrative taxonomic analysis of Sundaic populations of Gekko smithii from the Thai-Malaya Peninsula, Sumatra, and Borneo recovered four deeply divergent mitochondrial lineages that are separated by major geographic barriers (mountains and seaways). Furthermore, they bear a number of concordant statistically significant differences in meristic and morphometric features, morphospatial separation in multivariate space, and discrete differences in color pattern. Gekko smithii sensu stricto is restricted to southern Thailand south of the Isthmus of Kra and Peninsular Malaysia west of the Banjaran (mountain range) Titiwangsa, being that the type locality is on Penang Island, Penang. Gekko hulk sp. nov. is a new species from extreme southern Thailand and Peninsular Malaysia east of the Banjaran Titiwangsa and five east coast islands—the type locality being Pulau (island) Tioman, Pahang. Gekko cf. albofasciolatus is tentatively used to include Bornean populations west of the Iran Mountains in Sabah and Sarawak which, in the absence of molecular data, cannot unequivocally be separated morphologically from G. albofasciolatus from the type locality at Banjarmasin, Kalimantan, Indonesia east of the Iran Mountains. In the absence of molecular data, G. albomaculatus is resurrected to include mainland Sumatran, Nias Island, and Banyak Islands populations which, based on their morphology, cannot be separated from descriptions of G. albomaculatus from the type locality of Bangka Island, 15 km off the southeast coast of mainland Sumatra. Further integrative analyses of all Sumatran and Bornean populations are currently underway as well as the enigmatic Wallacean populations from Sulawesi. Data are presented that strongly suggest all references to G. smithii from Java stem from a 151 year-old misidentification of a specimen of G. gecko of unknown provenance. Additionally, there are no vouchered records of G. smithii from Myanmar. The phylogeographic patterns of Sundaic populations of the G. smithii complex are concordant with those of a plethora of other Sundaic lineages.

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DOI: 10.3897/vz.72.e78092

Authors: L. Lee Grismer, Kin Onn Chan, Robert E. Lovich, Jesse L. Grismer

Abstract: This study re-analyzes morphometric and meristic data among island night lizards, Xantusia riversiana, from the California Channel Islands of San Clemente, Santa Barbara, and San Nicolas in order to ascertain whether the implementation of different statistical methods can recover different results that could potentially alter biological interpretations. Our results concur with a recent previous study demonstrating that the three island populations differ morphologically from one another and that the San Nicolas Island population is the most divergent. Several important aspects, however, of the previous study depart significantly from those recovered here. Our analyses found sexual dimorphism within each population for both morphometric and meristic characters to be relatively uncommon whereas the previous study found nearly all characters to be sexually dimorphic for all island populations. The previous study also recovered significant differences among the three island populations for all morphometric characters whereas far fewer differences were recovered in the present study. Both studies found few significant inter-island differences among the meristic characters. The discordances between these two studies stem from differences in the a priori treatment of the raw character data and the different downstream statistical analyses and visualization techniques used on those data. This was particularly relevant with the use here of an allometric growth algorithm for size-correcting the morphometric data not used in the previous study and by treating all three populations as independently evolving groups. We did not conduct analyses where data from the San Clemente and Santa Barbara island populations were conflated based on their subspecific designation (X. r. reticulata) and then compared to data from the independently evolving San Nicolas Island population. This imprudent use of taxonomy violates the assumptions of statistical independence. We emphasize that explicit justification for the use of particular statistical analyses should occur in all studies—especially if the results bear on the implementation of effective and efficient resource management programs.

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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.e71995

Authors: Gustavo Campillo-García, Oscar Flores-Villela, Brett Oliver Butler, Julián Andrés Velasco Vinasco, Fabiola Ramírez Corona

Abstract: The spiny lizard genus Sceloporus was described by Wiegmann in 1828, with S. torquatus posteriorly designated as the type species. The taxonomic history of S. torquatus is complicated, as it has been confused with other taxa by numerous authors. Many modern systematics works have been published on Sceloporus, but none have included all five recognized S. torquatus subspecies: S. t. torquatus, S. t. melanogaster, S. t. binocularis, S. t. mikeprestoni, and S. t. madrensis. Additionally, there is previous evidence for at least one unnamed taxon. The present study is the first taxonomic revision of the enigmatic S. torquatus based on molecular phylogenies using combined molecular data from 12S, ND4 and RAG1 genes, and Maximum Likelihood and Bayesian inference phylogenetic methods. This work includes the most extensive sampling across the entire distribution, as well as divergence time estimates and environmental niche modelling, which combined offer a spatio-temporal framework for understanding the evolution of the species. Additionally, a series of morphological characters are analyzed to identify significant differences between lineages consistently recovered in the molecular phylogenies. Using this integrative approach, evidence is presented for eight lineages within the S. torquatus complex, five of which correspond to previously recognized subspecies and three represent unnamed taxa masked by morphological conservatism. Finally, to maintain taxonomic stability a lectotype and paralectoype are designated for S. torquatus, and certain taxonomic changes are suggested in order to reflect the phylogenetic relationships within the S. torquatus complex.

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DOI: 10.3897/vz.71.e75967

Authors: V. Deepak, Samuel Lalronunga, Esther Lalhmingliani, Abhijit Das, Surya Narayanan, Indraneil Das, David J. Gower

Abstract: Xenodermidae is a generally poorly known lineage of caenophidian snakes found in South, East and Southeast Asia. We report molecular phylogenetic analyses for a multilocus data set comprising all five currently recognised genera and including new mitochondrial and nuclear gene sequence data for the recently described Stoliczkia vanhnuailianai. Our phylogenetic results provide very strong support for the non-monophyly of Stoliczkia, as presently constituted, with S. borneensis being more closely related to Xenodermus than to the Northeast Indian S. vanhnuailianai. Based on phylogenetic relationships and morphological distinctiveness, we transfer Stoliczkia borneensis to a new monotypic genus endemic to Borneo, Paraxenodermus gen. nov. We also present new morphological data for P. borneensis.

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DOI: 10.3897/vz.71.e76069

Authors: L. Lee Grismer, Chatmongkon Suwannapoom, Parinya Pawangkhanant, Roman A. Nazarov, Platon V. Yushchenko, Mali Naiduangchan, Minh Duc Le, Vinh Quang Luu, Nikolay A. Poyarkov

Abstract: The first integrative taxonomic analysis of the Cyrtodactylus brevipalmatus group of Southeast Asia recovered two newly discovered populations from the Tenasserim Mountains in Suan Phueng District, Ratchaburi Province, Thailand as a new species described here as C. rukhadeva sp. nov. Based on 1397 base pairs of the mitochondrial gene NADH dehydrogenase subunit 2 (ND2), C. rukhadeva sp. nov. is the well-supported sister species to a clade containing three undescribed species, C. ngati, and C. cf. interdigitalis with a large uncorrected pairwise sequence divergence from other species in the brevipalmatus group ranging from 15.4–22.1%. Cyrtodactylus elok and C. brevipalmatus are recovered as poorly supported sister species and the well-supported sister lineage to the remainder of the brevipalmatus group. Cyrtodactylus rukhadeva sp. nov. is putatively diagnosable on the basis of a number of meristic characters and easily separated from the remaining species of the brevipalmatus group by a number of discrete morphological characters as well as its statistically significant wide separation in multivariate morphospace. The discovery of C. rukhadeva sp. nov. continues to underscore the unrealized herpetological diversity in the upland forests of the Tenasserim Mountains and that additional field work will undoubtedly result in the discovery of additional new species.

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DOI: 10.3897/vz.71.e62787

Authors: Gaurang Gowande, Saunak Pal, Daniel Jablonski, Rafaqat Masroor, Pushkar U. Phansalkar, Princia Dsouza, Aditi Jayarajan, Kartik Shanker

Abstract: Abstract The genus Calotes Cuvier, 1817 (Agamidae: Draconinae) is highly diverse, with species occurring in South and Southeast Asia, and Oceania. Most species of the subfamily except C. versicolor have narrow geographic distributions. Calotes versicolor is distributed from western Iran in the west to south China and Indonesia in the east and has been introduced to parts of Africa and North America. The species has had a complicated taxonomic history; multiple species and subspecies related to C. versicolor were described from India and adjoining regions, which were synonymized in subsequent revisions. However, a study of Burmese C. versicolor yielded two new species, C. htunwini and C. irawadi, indicating that C. versicolor is a species complex. Such integrative taxonomic studies have not been carried out in India, the supposed type locality of C. versicolor. Hence, we studied C. versicolor sensu lato from the Indian subcontinent and generated sequences of mitochondrial 16S and COI fragments from tissues sampled from multiple localities in the region, including the type localities of its synonyms. Phylogenetic analyses revealed four well-supported, deeply-divergent lineages, supported by morphological data. These lineages represent (i) C. versicolor sensu stricto, from South India and parts of the east coast, (ii) C. irawadi sensu lato from northeast India and Southeast Asia, (iii) a synonym from the eastern Indo-Gangetic Plains which we resurrect here, and (iv) a subspecies from Pakistan which we elevate to species level. We provide re-descriptions for the resurrected or elevated species, and a diagnostic key to the species of the C. versicolor complex. The study shows that C. versicolor sensu stricto is endemic to parts of southern and eastern India, and not widely distributed, though it may have been introduced to other parts of the world.

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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.e64781

Authors: Javier Lobón-Rovira, Werner Conradie, David Buckley Iglesias, Raffael Ernst, Luis Veríssimo, Ninda Baptista, Pedro Vaz Pinto

Abstract: Abstract The taxonomy of Angolan Hemidactylus has recently been revised. However, the lack of fresh material for some groups and regions, has led to the misidentification of some taxa and an underestimation of actual diversity in others. To shed light on the evolutionary history and systematics of Angolan Hemidactylus, we generated a new phylogenetic hypothesis for the group, and updated the taxonomy following an integrative approach. This resulted in the description of four new species (H. pfindaensis sp. nov., H. faustus sp. nov., H. carivoensis sp. nov. and H. cinganji sp. nov.), the reevaluation of two recently described species (H. vernayi and H. paivae) and the synonymization of a recently described species (H. hannahsabinnae). We estimate divergence times for these lineages, providing a preliminary interpretation of their speciation process. Moreover, we suggest and outline 13 Angolan Main Biogeographic Units (AMBUs) in the area, defining a new biogeographic context for future works on Angolan herpetofauna. We now recognize eleven Hemidactylus species in Angola, and we provide here a new morphological key for Angolan Hemidactylus to help with identifications and species assignments in this group.

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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.e69214

Authors: L. Lee Grismer

Abstract: Abstract It is hypothesized that shape differences between the closely related sandstone night lizard (Xantusia gracilis) and the granite night lizard (X. henshawi) may be correlated with structual differences in their respective microhabitats. Multivariate and univariate analyses of 22 morphometric characters taken from the head, body, and limbs of both saxicolus specialists recovered statistically significant differences between them with X. gracilis having a wider head, longer snout, larger eyes, wider sternum, higher and wider pelvis, thinner limbs, longer forearms and thighs, and longer hind limbs. Many of these same proportional differences have been reported among very closely related saxicolus species in other lizard families (i.e. Eublepharidae, Gekkonidae, Phrynosomatidae) that also live on different rocky substrates. This supports the inference that morphometric differences between X. gracilis and X. henshawi are ecomorphological adaptations for navigating the substantially different substrates of their respective microhabitats. Xantusia gracilis is restricted to a loose, heterogeneous, sandstone microhabitat composed of large boulders, small rocks, and cliff faces where cracks, crevices, holes, and exfoliations are used as retreats versus the compact, more homogeneous, granite boulder microhabitat of X. henshawi where narrow spaces beneath exfoliations and cap-rocks are the preferred retreats.

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DOI: 10.3897/vz.71.e66871

Authors: Amanda Kaatz, Jesse L. Grismer, L. Lee Grismer

Abstract: Abstract By studying ecomorophology in the context of phylogeny, researchers can parse out similarity due to common ancestry versus that due to convergence. This is especially true among relatively closely related species where both phylogenetic and environmental constraints may be operating simultaneously. We explored these issues among three karst-associated species from two lineages of Cyrtodactylus—the sworderi group from Peninsular Malaysia and the swamp clade from Peninsular Malaysia and western Indonesia of the agamensis group. A stochastic character mapping analysis using five different habitat preferences corroborated a larger previous analysis in recovering a general habitat preference as an ancestral condition for all habitat preferences and a karst habitat preference in C. guakanthanensis and C. gunungsenyumensis of the sworderi group and C. metropolis of the swamp clade as convergently evolved. Multivariate and univariate analyses of 10 morphometric characters revealed that the ecomorphological similarity between C. guakanthanensis and C. gunungsenyumensis of the sworderi group was also convergent. The ecomorphology of C. metropolis of the swamp clade was intermediate between a karst-adapted ecomorphology and a swamp-generalists ecomorphology. Of the 10 morphometric characters employed in this analysis, only three—head length, head width, and forelimb width—showed any signs of phylogenetic signal. Cyrtodactylus metropolis is hypothesized to be a recently refuged swamp-dwelling species that frequented the Batu Caves environments prior to urbanization of the surrounding swamp habitat to which it is now confined.

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DOI: 10.3897/vz.71.e65969

Authors: L. Lee Grismer, Hai Ngoc Ngo, Shuo Qi, Ying-Yong Wang, Minh Duc Le, Thomas Ziegler

Abstract: Abstract Maximum likelihood (ML) and Bayesian inference (BI) analyses using two mitochondrial (16S and cyt b) and two nuclear (CMOS and RAG1) genes and 103 specimens recovered the first phylogenies of all 23 extant species of Goniurosaurus. The analyses strongly supported the recognition of four monophyletic species groups with identical inter-specific relationships within the kuroiwae, lichtenfelderi, and yingdeensis groups but discordant topologies at some nodes within the luii group. Both analyses recovered a polyphyletic G. luii with respect to G. kadoorieorum, and owing to the lack of diagnostic characters in the latter, it is considered a junior synonym of G. luii. A stochastic character mapping analysis of karst versus non-karst habitat preference suggested that karstic landscapes may have played a major role in the evolution and diversification of Goniurosaurus. A karst habitat preference is marginally supported as the most probable ancestral condition for Goniurosaurus as well as for the kuroiwae, luii, and yingdeensis groups. However, a non-karst habitat preference is marginally supported as the most probable ancestral condition for the lichtenfelderi group. Multivariate and univariate ecomorphological analyses of the karst-adapted G. catbaensis, G. huuliensis, and G. luii of the luii group and the granite-stream-adapted G. lichtenfelderii of the lichtenfelderi group demonstrated that their markedly statistically different body shapes may be an adaptive response that contributes to habitat partitioning in areas of northern Vietnam where they are nearly sympatric.

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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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DOI: 10.3897/vz.71.e60800

Authors: Daniel Jablonski, Marco Antônio Ribeiro-Júnior, Shai Meiri, Erez Maza, Oleg V. Kukushkin, Marina Chirikova, Angelika Pirosová, Dušan Jelić, Peter Mikulíček, David Jandzik

Abstract: The Levant represents one of the most important reptile diversity hotspots and centers of endemism in the Western Palearctic. The region harbored numerous taxa in glacial refugia during the Pleistocene climatic oscillations. Due to the hostile arid conditions in the warmer periods they were not always able to spread or come into contact with populations from more distant regions. One large and conspicuous member of the Levantine herpetofauna is the legless anguid lizard Pseudopus apodus. This species is distributed from the Balkans to Central Asia with a portion of its range running along the eastern Mediterranean coast. Mitochondrial and nuclear DNA sequences, microsatellite genotypes, and morphology show that populations in this region differ from the two named subspecies and presumably had a long independent evolutionary history during the Quaternary. Here we describe the Levantine population as a new subspecies and present biogeographic scenarios for its origin and diversification. The new subspecies is genetically highly diverse, and it forms a sister lineage to Pseudopus from the remaining parts of the range according to mtDNA. It is the largest-bodied of the three subspecies, but occupies the smallest range.

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DOI: 10.3897/vz.71.e60225

Authors: Anh The Nguyen, Tang Van Duong, L. Lee Grismer, Nikolay A. Poyarkov

Abstract: An integrative analysis of genetic, morphological, and ecological data recovered a new granite cave-adapted species, Cyrtodactylus raglai sp. nov., from the Song Giang River Valley, Khanh Hoa Province, Vietnam. Cyrtodactylus raglai sp. nov. is nested within one of two major clades within the irregularis species group where it forms a monophyletic group with C. cryptus and its sister species C. kingsadai. It differs from its sister species by an uncorrected pairwise sequence divergence of 16.5% and 16.8% based on the mitochondrial genes NADH dehydrogenase subunit 2 and its flanking tRNAs and the first subunit of cytochrome oxidase c (CO1), respectively. Cyrtodactylus raglai sp. nov. is a narrow-range endemic restricted to a riparian, granite cave microhabitat and its overall morphology bears that of other granite cave-dwelling ecomorphs in the genus. The Song Giang River Valley and its associated waterways are currently threatened by the construction of a hydropower station which will negatively impact the surrounding ecosystem. Urgent field surveys in this region are necessary in order to obtain critical data on its biodiversity and importance to conservation efforts in southern Vietnam.

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DOI: 10.3897/vz.71.e59307

Authors: L. Lee Grismer, Perry L. Wood, Jr., Nikolay A. Poyarkov, Minh D. Le, Fred Kraus, Ishan Agarwal, Paul M. Oliver, Sang N. Nguyen, Truong Q. Nguyen, Suranjan Karunarathna, Luke J. Welton, Bryan L. Stuart, Vinh Q. Luu, Aaron M. Bauer, Kyle A. O’Connell, Evan S. H. Quah, Kin O. Chan, Thomas Ziegler, Hanh Ngo, Roman A. Nazarov, Anchalee Aowphol, Siriwadee Chomdej, Chatmongkon Suwannapoom, Cameron D. Siler, Shahrul Anuar, Ngo V. Tri, Jesse L. Grismer

Abstract: The gekkonid genus Cyrtodactylus is the third most speciose vertebrate genus in the world, containing well over 300 species that collectively range from South Asia to Melanesia across some of the most diverse landscapes and imperiled habitats on the planet. A genus-wide phylogeny of the group has never been presented because researchers working on different groups were using different genetic markers to construct phylogenies that could not be integrated. We present here Maximum likelihood and Bayesian inference mitochondrial and mito-nuclear phylogenies incorporating of 310 species that include dozens of species that had never been included in a genus-wide analysis. Based on the mitochondrial phylogeny, we partition Cyrtodactylus into 31 well-supported monophyletic species groups which, if used as recommended herein, will increase the information content of future integrative taxonomic analyses that continue to add new species to this genus at an ever-increasing annual rate. Data presented here reiterate the outcome of several previous studies indicating that Cyrtodactylus comprises an unprecedented number of narrow-range endemics restricted to single mountain tops, small islands, or karst formations that still remain unprotected. This phylogeny can provide a platform for various comparative ecological studies that can be integrated with conservation management programs across the broad diversity of landscapes and habitats occupied by this genus. Additionally, these data indicate that the true number of Cyrtodactylus remains substantially underrepresented.

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DOI: 10.3897/vz.71.e59461

Authors: Juan José Torres-Ramírez, Teddy Angarita-Sierra, Mario Vargas-Ramírez

Abstract: In northern South America, amphisbaenians are rarely seen among the herpetofauna.Thus, general knowledge about them is very poor. During a herpetological survey in 2012 at Casanare, Colombia, we found two specimens of an unusual Amphisbaena. A third specimen sharing the same morphotype labeled Amphisbaena sp. from Vichada department was found deposided in an Colombian reptile collection. Based on morphological analyses together with phylogenetic analyses of 1029 base pairs of the mitochondrial DNA (mtDNA), we describe a new species of Amphisbaena that inhabits in the Orinoquian region of Colombia. The new species is part of a phylogenetic clade together with A. mertensii and A. cunhai (central-southern Brazil), exhibiting a great genetic distance (26.1–28.9%) between the newly identified lineage versus those taxa, and versus the sympatric taxa A. alba and A. fuliginosa. Morphologically, this new Amphisbaena can be distinguished from their congeners by characters combination of number of preocloacal pores, absence of malar scale, postgenial scales and body and caudal annuli counts. Amphisbaena gracilis is on morphology grounds the most similar species. However, the new species can be distinguished from it by having higher body annuli counts, angulus ories aliegned with the edges of the ocular scales and center of frontal scales, less number of large middorsal segments of the first and second body annulus, and rostral scale visible from above. The description of this new Amphisbaena species points out the urgent need to increase the knowledge of worm lizards in Colombia

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DOI: 10.3897/vz.71.e59495

Authors: Aurélien Miralles, Teddy Bruy, Angelica Crottini, Andolalao Rakotoarison, Fanomezana M. Ratsoavina, Mark D. Scherz, Robin Schmidt, Jörn Köhler, Frank Glaw, Miguel Vences

Abstract: The Paroedura bastardi clade, a subgroup of the Madagascan gecko genus Paroedura, currently comprises four nominal species: P. bastardi, supposedly widely distributed in southern and western Madagascar, P. ibityensis, a montane endemic, and P. tanjaka and P. neglecta, both restricted to the central west region of the island. Previous work has shown that Paroedura bastardi is a species complex with several strongly divergent mitochondrial lineages. Based on one mitochondrial and two nuclear markers, plus detailed morphological data, we undertake an integrative revision of this species complex. Using a representative sampling for seven nuclear and five mitochondrial genes we furthermore propose a phylogenetic hypothesis of relationships among the species in this clade. Our analyses reveal at least three distinct and independent evolutionary lineages currently referred to P. bastardi. Conclusive evidence for the species status of these lineages comes from multiple cases of syntopic occurrence without genetic admixture or morphological intermediates, suggesting reproductive isolation. We discuss the relevance of this line of evidence and the conditions under which concordant differentiation in unlinked loci under sympatry provides a powerful approach to species delimitation, and taxonomically implement our findings by (1) designating a lectotype for Paroedura bastardi, now restricted to the extreme South-East of Madagascar, (2) resurrecting of the binomen Paroedura guibeae Dixon & Kroll, 1974, which is applied to the species predominantly distributed in the South-West, and (3) describing a third species, Paroedura rennerae sp. nov., which has the northernmost distribution within the species complex.

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