Slug-eating snakes of the subfamily Pareinae are an insufficiently studied group of snakes specialized in feeding on terrestrial mollusks. Currently Pareinae encompass three genera with 34 species distributed across the Oriental biogeographic region. Despite the recent significant progress in understanding of Pareinae diversity, the subfamily remains taxonomically challenging. Here we present an updated phylogeny of the subfamily with a comprehensive taxon sampling including 30 currently recognized Pareinae species and several previously unknown candidate species and lineages. Phylogenetic analyses of mtDNA and nuDNA data supported the monophyly of the three genera Asthenodipsas, Aplopeltura, and Pareas. Within both Asthenodipsas and Pareas our analyses recovered deep differentiation with each genus being represented by two morphologically diagnosable clades, which we treat as subgenera. We further apply an integrative taxonomic approach, including analyses of molecular and morphological data, along with examination of available type materials, to address the longstanding taxonomic questions of the subgenus Pareas, and reveal the high level of hidden diversity of these snakes in Indochina. We restrict the distribution of P. carinatus to southern Southeast Asia, and recognize two subspecies within it, including one new subspecies proposed for the populations from Thailand and Myanmar. We further revalidate P. berdmorei, synonymize P. menglaensis with P. berdmorei, and recognize three subspecies within this taxon, including the new subspecies erected for the populations from Laos and Vietnam. Furthermore, we describe two new species of Pareas from Vietnam: one belonging to the P. carinatus group from southern Vietnam, and a new member of the P. nuchalis group from the central Vietnam. We provide new data on P. temporalis, and report on a significant range extension for P. nuchalis. Our phylogeny, along with molecular clock and ancestral area analyses, reveal a complex diversification pattern of Pareinae involving a high degree of sympatry of widespread and endemic species. Our analyses support the “upstream” colonization hypothesis and, thus, the Pareinae appears to have originated in Sundaland during the middle Eocene and then colonized mainland Asia in early Oligocene. Sundaland and Eastern Indochina appear to have played the key roles as the centers of Pareinae diversification. Our results reveal that both vicariance and dispersal are responsible for current distribution patterns of Pareinae, with tectonic movements, orogeny and paleoclimatic shifts being the probable drivers of diversification. Our study brings the total number of Pareidae species to 41 and further highlights the importance of comprehensive taxonomic revisions not only for the better understanding of biodiversity and its evolution, but also for the elaboration of adequate conservation actions.
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.
We describe a new species of Oligodon from the highlands of the Langbian Plateau, southern Truong Son Mountains, Vietnam, based on morphological and molecular phylogenetic analyses. The new species, Oligodon rostralis sp. nov is distinguished from its congeners by the following morphological characters: medium size in adults (male TL = 582 mm); small and broad head with long protruding snout; dorsal scale row formula 15-15-13; 167 ventrals, 47 subcaudals; single preocular, single postocular; loreal and presubocular absent; six supralabials, third and fourth entering orbit; six infralabials, anterior four contacting first pair of chin shields; internasals separate from prefrontals; nasal divided; single anterior and two posterior temporals; cloacal plate undivided; hemipenes short, bilobed, bifurcating in anterior one third of their length, extending to 8th subcaudal, lacking spines and papillae, with a prominent transverse flounces and distal calyces; six maxillary teeth, the posterior three enlarged; dorsal pattern consisting of 14+4 large dark-brown blotches and a bright-orange vertebral stripe on tail and dorsum; and ventral surfaces in life cream laterally with dark quadrangular spots; dark temporal streak present, edged with white. We also provide additional information on O. annamensis, including a morphological dataset of all specimens known from natural history collections and confirmation of an earlier record of O. annamensis from Cambodia. We also provide the first record of O. annamensis for Dak Lak Province. Phylogenetic analyses of mtDNA genes (3,131 bp of 12S rRNA, 16S rRNA and cyt b) suggest sister relationships of Oligodon rostralis sp. nov. and O. annamensis and place them in one clade with the O. cyclurus and O. taeniatus species groups, which is concordant with previous studies on the phylogenetic relationships of Oligodon. Our study demonstrates high level of herpetofaunal diversity and endemism of Langbian Plateau and further supports the importance of this area for conservation herpetofaunal diversity in Indochina.
An integrative taxonomic analysis recovered the new species Cnemaspis selenolagus sp. nov. of the C. siamensis group as the sister species to C. punctatonuchalis. The new species was discovered in mountain evergreen tropical forests of in Suan Phueng District of Ratchaburi Province, western Thailand. Additionally, the analysis recovered a deep genetic divergence between northern and southern clades within the C. siamensis group that occur on opposite sides of the Isthmus of Kra—a well-known biogeographic region of cladogenic turnover. The description of C. selenolagus sp. nov. brings the total number of species of Cnemaspis in Thailand to 16, 11 of which compose the C. siamensis group—a lineage endemic to the Thai-Malay Peninsula. This underscores the physiographic complexity of this narrow peninsula in that it can support a large number of closely related species in only the northern two-thirds of its length.
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 cattienensisVassilieva 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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