The evolutionary history of an accidental model organism, the leopard gecko Eublepharis macularius (Squamata: Eublepharidae).

The leopard gecko, Eublepharis macularius, is a widely used model organism in laboratory and experimental studies. The high phenotypic diversity in the pet trade, the fact that the provenance of different breeding lines is unknown, and that distinct Eublepharis species are known to hybridize, implies that the continued use of E. macularius as a model requires clarity on the origin of the lineages in the pet trade. We combine multi-locus sequence data and the first range-wide sampling of the genus Eublepharis to reconstruct the evolutionary history of the Eublepharidae and Eublepharis, with an updated time-tree for the Eublepharidae. Our sampling includes five of the six recognized species and additional nominal taxa of uncertain status comprising 43 samples from 34 localities plus 48 pet-trade samples. The Eublepharidae began diversifying in the Cretaceous. Eublepharis split from its sister genera in Africa in the Palaeocene-Eocene, and began diversifying in the Oligocene-Miocene, with late Miocene-Pliocene cladogenesis giving rise to extant species. The current species diversity within this group is moderately underestimated. Our species delimitation suggests 10 species with four potentially unnamed divergent lineages in Iran, India and Pakistan. All 30 individuals of E. macularius that we sampled from the pet trade, which include diverse morphotypes, come from a few shallow E. macularius clades, confirming that lab and pet trade strains are part of a single taxon. One of the wild-caught haplotypes of E. macularius, from near Karachi, Pakistan, is identical to (10) pet-trade samples and all other captive populations are closely related to wild-caught animals from central/southern Pakistan (0.1-0.5 % minimum pairwise uncorrected ND2 sequence divergence).

Two new species of yellow-tailed Hemidactylus Goldfuss, 1820 (Squamata: Gekkonidae) from rocky outcrops on the Telangana Plateau, India.

Two distinct species of the gekkonid genus Hemidactylus are described from the dry zone of peninsular India from the state of Telangana. The two sister species, Hemidactylus flavicaudus sp. nov. and H. xericolus sp. nov., are nested within the morphologically cryptic H. brookii group, but are clearly distinguishable from all known species in having a bright yellow tail and yellow markings on the head, besides unique combinations of meristic characters and small body size (< 45 mm snout to vent length). The two new species are also deeply divergent from each other and other members of the H. brookii group in mitochondrial NADH dehydrogenase 2 and cytochrome b sequences. Hemidactylus flavicaudus sp. nov. can be diagnosed from H. xericolus sp. nov. based on the number of dorsal tubercle rows at midbody (11-14 versus 6-8). These descriptions of evolutionarily and morphologically distinct species highlight the dearth of studies carried out in the dry zone of peninsular India and the urgent need to assess biodiversity in the face of rapid land-use changes in this landscape.

An expanded ND2 phylogeny of the brookii and prashadi groups with the description of three new Indian Hemidactylus Oken (Squamata: Gekkonidae).

We present an expanded ND2 phylogeny of the brookii and prashadi groups of Indian Hemidactylus. While relationships within the clades are not well resolved, we recover many well supported subclades and a number of deeply divergent lineages within each clade. We use morphological data to describe three of these divergent lineages as new species; Hemidactylus kolliensis sp. nov., a member of the prashadi group from high elevation in the Kollimalai Massif in Tamil Nadu, Hemidactylus chikhaldaraensis sp. nov. and Hemidactylus sankariensis sp. nov., both members of the brookii group, are described from high elevation of the Satpuras in Maharashtra and low elevation of Tamil Nadu, respectively. The three new species exhibit >15% divergence for the ND2 gene from their phylogenetic sisters and differ in a number of morphological characters from all Indian congeners. The discovery of three more endemic Hemidactylus species from rocky habitats suggests that many more remain to be discovered across India.

On the systematic status of the genus Oriocalotes Günther, 1864 (Squamata: Agamidae: Draconinae) with the description of a new species from Mizoram state, Northeast India.

The montane agamid lizard genus Oriocalotes is currently considered monotypic, represented by the species, O. paulus. The systematic status of this taxon has remained questionable since its initial descriptions in the mid-1800s. A detailed molecular and morphological study was carried out to assess the validity of this genus, and its systematic position within the Asian agamid subfamily, Draconinae. Freshly collected and historical museum specimens from the type locality of O. paulus were examined morphologically, along with additional samples collected from localities in Mizoram state, Northeast India. Utilising newly generated molecular sequences (two mitochondrial and three nuclear genes), combined with those previously published for representative genera from the subfamilies Draconinae and Agaminae, Maximum Likelihood and Bayesian phylogenetic trees were constructed. Phylogenetic results suggest that Oriocalotes is part of the widespread South and Southeast Asian radiation of Calotes. Comparative morphological studies (including external morphology, hemipenis and osteology) between Oriocalotes and related genera further support this systematic placement. Oriocalotes is herein regarded as a junior subjective synonym of Calotes. Calotes paulus comb. nov. is also assigned a lectotype and given a detailed redescription based on the lectotype, paralectotypes and additional topotypic material. Furthermore, the specimens collected from Mizoram populations are found to be morphologically and genetically distinct from Calotes paulus comb. nov., and are described herein as a new species, Calotes zolaiking sp. nov.

Diversification in the mountains: a generic reappraisal of the Western Ghats endemic gecko genus Dravidogecko Smith, 1933 (Squamata: Gekkonidae) with descriptions of six new species.

The monotypic genus Dravidogecko, represented by its type-species D. anamallensis, is singular amongst peninsular Indian gekkonid lineages in its endemism to the Western Ghats. Molecular species delimitation approaches reveal at least seven species-level lineages within the genus from its distribution range across the mid-high elevations of the southern Western Ghats of India. These lineages, albeit superficially cryptic, are patently diagnosable from each other by employing a limited but precise set of morphological characters. Six of these lineages that were obscured under the nomen D. anamallensis are herein recognized as distinct species. A reappraisal of the genus Dravidogecko is provided based on external morphology and osteological characters, along with a detailed redescription of the holotype of D. anamallensis. A key to the species based on diagnostic characters is presented. Gene-trees based on mitochondrial and nuclear DNA data recovered marginally disparate topologies and were consequently coalesced into a species-tree for phylogenetic inference. Timetree analysis reveals late Miocene cladogenesis in this group and establishes late Palaeocene divergence from its sister genus, Hemidactylus, making Dravidogecko one of the earliest, extant lizard lineages to have colonized peninsular India.

A new genus and species of natricine snake from northeast India.

Based on the first molecular phylogenetic analyses of samples from northeast India, specimens referred to Rhabdops from this region are more closely related to the southeast and east Asian natricine genera Opisthotropis Günther, 1872 and Sinonatrix Rossman Eberle, 1977 (as well as to New World and western Palearctic natricines) than to peninsular Indian (true) Rhabdops. Morphologically, these northeast Indian populations differ from other natricines by having a single ('fused' or unpaired) internasal shield and a single prefrontal shield. Given the morphological and phylogenetic distinctiveness of these northeast Indian populations, we refer them to a new genus, Smithophis gen. nov., and transfer Rhabdops bicolor (Blyth, 1854) to Smithophis bicolor comb. nov. Based on morphological and molecular variation within our northeast Indian sample, we additionally describe Smithophis atemporalis sp. nov. from the state of Mizoram.

Six new Cyrtodactylus (Squamata: Gekkonidae) from northeast India.

We use mitochondrial sequence data to identify divergent lineages within the gekkonid genus Cyrtodactylus in northeast India and use morphological data to describe six new species from within the Indo-Burma clade of Cyrtodactylus. The new species share an irregular colour pattern but differ from described species from the region in morphology and mitochondrial sequence data (>11 % uncorrected pairwise sequence divergence). Three new species are from along the Brahmaputra River and three are from mountains south of the Brahmaputra, including the largest Cyrtodactylus from India and the fifth gecko to be described from a major Indian city, Guwahati.

Two new species of bent toed geckos, Cyrtodactylus Gray, 1827 (Squamata: Gekkonidae) from Northeast India with comments on name-bearing types from the region.

Two new species of the genus Cyrtodactylus are described based on collections from Northeast India, and descriptions and diagnoses of previously described species and synonyms from the region are provided. One of the new species from lowland habitats in Tripura is medium sized, with a continuous series of precloacal pores, and is allied to C. ayeyarwadyensis from Myanmar; while the other new species from West Bengal is smaller, with both femoral and precloacal pores, and is related to C. gubernatoris from West Bengal and Sikkim. We designate a lectotype for Cyrtodactylus khasiensis and recognize the C. khasiensis synonym Gymnodactylus himalayicus as a valid species of Cyrtodactylus based on morphological evidence, and suggest the recognition of the last remaining subspecies of C. khasiensis, C. k. cayuensis, as a valid species based on geography and morphology. The stabilization of the taxa Cyrtodactylus khasiensis and C. gubernatoris with a limited subset of characters and specific distributional data, as well as the elevation of Cyrtodactylus himalayicus comb. nov. from synonymy, will allow the descriptions of many more Cyrtodactylus species from the region.

A new cryptic, rupicolous species of Hemidactylus Oken, 1817 (Squamata: Gekkonidae) from Meghamalai, Tamil Nadu, India.

A distinct new gecko of the genus Hemidactylus is described from the Meghamalai massif in Tamil Nadu, India. This large sized (average SVL 88.2±16.0 to at least 112.2 mm), cryptic, rock-dwelling species is superficially similar to its tuberculate South Asian congeners of comparable size, but can be distinguished from them in having 17-19 longitudinal rows of fairly regularly arranged, strongly keeled, striated tubercles with the two most medial parasagittal rows smaller, flatter, strongly keeled and rounded, increasing in size laterally and becoming strongly conical at the flanks; 8-10 and 10-12 sub-digital lamellae under the first and fourth digits respectively of both manus and pes; 17-22 femoral pores on each side separated medially by a diastema of 10-11 poreless scales; 11-15 supralabials and 9-14 infralabials. Molecular data supports the distinctiveness of this species and helps ascertain its phylogenetic position within the larger Indian Hemidactylus clade.

A new species of large-bodied, tuberculate Hemidactylus Oken (Squamata: Gekkonidae) from the Eastern Ghats, India.

A distinct new gecko of the genus Hemidactylus is described from Andhra Pradesh, India. This large-sized (snout to vent length up to at least 105 mm), scansorial Hemidactylus is characterized by dorsal scalation of small granules intermixed with large, pointed, trihedral tubercles that form 16-17 fairly regularly arranged longitudinal rows at midbody; 9-11 subdigital lamellae below the first and 11-13 below the fourth digit; 6-8 strongly pointed and keeled enlarged tubercles on the original tail; 20-23 femoral pores separated by 4 poreless scales in males; 11-13 supralabials and 9-11 infralabials. This is the third vertebrate endemic to the Mahendragiri Range, highlighting the significance of this topographically complex region.

A unique mating strategy without physical contact during fertilization in Bombay Night Frogs (Nyctibatrachus humayuni) with the description of a new form of amplexus and female call.

Anurans show the highest diversity in reproductive modes of all vertebrate taxa, with a variety of associated breeding behaviours. One striking feature of anuran reproduction is amplexus. During this process, in which the male clasps the female, both individuals' cloacae are juxtaposed to ensure successful external fertilization. Several types of amplexus have evolved with the diversification of anurans, and secondary loss of amplexus has been reported in a few distantly related taxa. Within Nyctibatrachus, a genus endemic to the Western Ghats of India, normal axillary amplexus, a complete loss of amplexus, and intermediate forms of amplexus have all been suggested to occur, but many species remain unstudied. Here, we describe the reproductive behaviour of N. humayuni, including a new type of amplexus. The dorsal straddle, here defined as a loose form of contact in which the male sits on the dorsum of the female prior to oviposition but without clasping her, is previously unreported for anurans. When compared to known amplexus types, it most closely resembles the form of amplexus observed in Mantellinae. Furthermore, we prove that, opposed to the situation in most anurans, male semen release happens before egg deposition. We hypothesize that the male ejaculates on the female's dorsum and that sperm subsequently runs from her back and hind legs before fertilizing the eggs. A second feature characterizing anuran breeding is the advertisement call, mostly produced solely by males. Despite recent descriptions of several new Nyctibatrachus species, few studies have explored their vocal repertoire. We describe both the male advertisement call and a female call for N. humayuni. The presence of a female call has not been reported within Nyctibatrachidae, and has been reported in less than 0.5% of anuran species. Altogether, our results highlight a striking diversity and several unique aspects of Nyctibatrachus breeding behaviour.

A new rock dwelling Hemidactylus (Squamata: Gekkonidae) from Chhattisgarh, India.

A distinct new species of gecko of the genus Hemidactylus is described from the Kanker district of Chhattisgarh State, east-central India. This large-sized (SVL average 81.33±13.40 to at least 98.0 mm) Hemidactylus is characterized by a dorsum with small granules, intermixed with 10-12 rows of irregularly arranged, slightly larger, rounded, weakly-keeled tubercles at midbody; 10-12 and 13-15 subdigital lamellae on the first and fourth digits, respectively, of both manus and pes; a single enlarged postcloacal tubercle on either side of the tail; 10-12 femoral pores on each thigh separated by 5-8 poreless scales; 12-14 supralabials and 10-12 infralabials.

The first teresomatan caecilian (Amphibia: Gymnophiona) from the Eastern Ghats of India--a new species of Gegeneophis Peters, 1880.

A new species of caecilian amphibian, Gegeneophis orientalis sp. nov., is described based on a series of nine specimens from high elevation (ca. 1,200 m) habitats in the Eastern Ghats in the states of Andhra Pradesh and Odisha, India. This species differs from all other congeners in having only bicuspid teeth in the outer as well as inner rows. The new species is the first caecilian reported from the state of Odisha, the first teresomatan caecilian from the Eastern Ghats, and is the only Indian indotyphlid known from outside the Western Ghats region.

A long-lost relic from the Eastern Ghats: Morphology, distribution and habitat of Sepsophis punctatus Beddome, 1870 (Squamata: Scincidae).

Sepsophis punctatus Beddome 1870, the only species of a monotypic genus, was described based on a single specimen from the Eastern Ghats of India. We rediscovered the species based on specimens from Odisha and Andhra Pradesh state, India, after a gap of 137 years, including four specimens from close to the type locality. The holotype was studied in detail, and we present additional morphological characters of the species with details on natural history, habitat and diet. The morphological characters of the holotype along with two additional specimens collected by Beddome are compared with the specimens collected by us. We also briefly discuss the distribution of other members of the subfamily Scincinae and their evolutionary affinities.

Mountain-associated clade endemism in an ancient frog family (Nyctibatrachidae) on the Indian subcontinent.

Night frogs (Nyctibatrachidae) form a family endemic to the Western Ghats, a hill chain along the west coast of southern India. Extant members of this family are descendants of a lineage that originated on the subcontinent during its longtime isolation in the Late Cretaceous. Because the evolutionary history of Nyctibatrachidae has always been tightly connected to the subcontinent, these tropically-adapted frogs are an ideal group for studying how patterns of endemism originated and evolved during the Cenozoic in the Western Ghats. We used a combined set of mitochondrial and nuclear DNA fragments to investigate the phylogenetic relationships of 120 ingroup specimens of all known species of Nyctibatrachidae. Our analyses indicate that, although this family had an early origin on the Indian subcontinent, the early diversification of extant nyctibatrachids happened only in the Eocene. Biogeographic analyses show that dispersal across the Palghat gap and Shencottah gap was limited, which led to clade endemism within mountain ranges of the Western Ghats. It is likely that multiple biota have been affected simultaneously by these prominent geographical barriers. Our study therefore further highlights the importance of considering the Western Ghats-Sri Lanka biodiversity hotspot as an assemblage of distinct mountain regions, each containing endemism and deserving attention in future conservation planning.

South Asia supports a major endemic radiation of Hemidactylus geckos.

Hemidactylus geckos are a species-rich component of many tropical lizard assemblages. We sampled deeply among tropical Asian, and especially South Asian, taxa and used a multi-gene approach to establish the affinities of Indian and Sri Lankan Hemidactylus and to evaluate the monophyly of previously proposed taxa within the genus. There is only weak support for the monophyly of tropical Asian Hemidactylus as a whole, but two strongly supported subclades were retrieved: the bowringii group is a predominantly East Asian clade that reaches South Asia only peripherally; the brookii group is a morphologically diverse clade that represents a previously unrecognized, species-rich (25 species), chiefly South Asian radiation. Deep genetic divergences support the specific recognition of three Sri Lankan taxa previously regarded as subspecies of mainland forms and validate H. subtriedrus as a species distinct from H. triedrus. Unlike several other vertebrate groups, Sri Lankan Hemidactylus do not represent a single insular radiation. Rather, each of six Sri Lankan species reached the island independently from different continental sources. There have been extensive Quaternary land connections between India and Sri Lanka but sister-species pairs of Hemidactylus on the two land masses diverged from one another much earlier, probably in the mid-Miocene.

Two new endemic genera and a new species of toad (Anura: Bufonidae) from the Western Ghats of India.

BACKGROUND: Bufonidae are a large family of toads with a subcosmopolitan distribution. Recent molecular phylogenetic analyses have revealed a radiation of toads (Adenominae) with distinct adult and larval ecomorphs on the Southern parts of the Indian subcontinent. The Indian torrential species "Ansonia" ornata has a basal position in this clade and does not group with South-East Asian Ansonia. Additionally, the nested position of "Bufo" koynayensis and an undescribed sister species, and their distinct ecologies including a non-typical egg-laying strategy within bufonids, support the recognition of a second distinct genus. In this paper we describe two new genera and one new species from the Adenominae clade. FINDINGS: Ansonia ornata Günther, 1876 "1875" is transferred to Ghatophryne gen. nov., a genus of torrentially adapted toads that are endemic to the Western Ghats of India. On the basis of close morphological resemblance and distribution, Ansonia rubigina Pillai and Pattabiraman, 1981 is provisionally transferred to this new genus. The Western Ghats endemic toad Bufo koynayensis Soman, 1963 is transferred to a new genus Xanthophryne gen. nov. Based on molecular and morphological evidence, we additionally describe a new species, Xanthophryne tigerinus sp. nov., from Amboli in the Western Ghats. CONCLUSION: The descriptions and subsequent taxonomic changes we propose result in three genera of bufonids recognised as being endemic to the Western Ghats (Ghatophryne gen. nov., Xanthophryne gen. nov. and Pedostibes), and one to Sri Lanka (Adenomus). The spatial distribution, and arrangement of these lineages at the base of Adenominae diversification, reflects their Early Neogene isolation in the Western Ghats-Sri Lanka hotspot.