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Unlike other snakes, most species of Rhabdophis possess glands in their dorsal skin, sometimes limited to the neck, known as nuchodorsal and nuchal glands, respectively. Those glands contain powerful cardiotonic steroids known as bufadienolides, which can be deployed as a defense against predators. Bufadienolides otherwise occur only in toads (Bufonidae) and some fireflies (Lampyrinae), which are known or believed to synthesize the toxins. The ancestral diet of Rhabdophis consists of anuran amphibians, and we have shown previously that the bufadienolide toxins of frog-eating species are sequestered from toads consumed as prey. However, one derived clade, the Rhabdophis nuchalis Group, has shifted its primary diet from frogs to earthworms. Here we confirm that the worm-eating snakes possess bufadienolides in their nucho-dorsal glands, although the worms themselves lack such toxins. In addition, we show that the bufadienolides of R. nuchalis Group species are obtained primarily from fireflies. Although few snakes feed on insects, we document through feeding experiments, chemosensory preference tests, and gut contents that lampyrine firefly larvae are regularly consumed by these snakes. Furthermore, members of the R. nuchalis Group contain compounds that resemble the distinctive bufadienolides of fireflies, but not those of toads, in stereochemistry, glycosylation, acetylation, and molecular weight. Thus, the evolutionary shift in primary prey among members of the R. nuchalis Group has been accompanied by a dramatic shift in the source of the species’ sequestered defensive toxins.

A large body of evidence indicates that evolutionary innovations of novel organs have facilitated the subsequent diversification of species. Investigation of the evolutionary history of such organs should provide important clues for understanding the basis for species diversification. An Asian natricine snake, Rhabdophis tigrinus, possesses a series of unusual organs, called nuchal glands, which contain cardiotonic steroid toxins known as bufadienolides. Rhabdophis tigrinus sequesters bufadienolides from its toad prey and stores them in the nuchal glands as a defensive mechanism. Among more than 3,500 species of snakes, only 17 Asian natricine species are known to possess nuchal glands or their homologues. These 17 species belong to three nominal genera, Balanophis, Macropisthodon, and Rhabdophis. In Macropisthodon and Rhabdophis, however, species without nuchal glands also exist. To infer the evolutionary history of the nuchal glands, we investigated the molecular phylogenetic relationships among Asian natricine species with and without nuchal glands, based on variations in partial sequences of Mt‐CYB, Cmos, and RAG1 (total 2,767 bp). Results show that all species with nuchal glands belong to a single clade (NGC). Therefore, we infer that the common ancestor of this clade possessed nuchal glands with no independent origins of the glands within the members. Our results also imply that some species have secondarily lost the glands. Given the estimated divergence time of related species, the ancestor of the nuchal gland clade emerged 19.18 mya. Our study shows that nuchal glands are fruitful subjects for exploring the evolution of novel organs. In addition, our analysis indicates that reevaluation of the taxonomic status of the genera Balanophis and Macropisthodon is required. We propose to assign all species belonging to the NGC to the genus Rhabdophis, pending further study. Some snakes have novel organs. To infer the evolutionary pathway of the organs, we investigated molecular phylogeny of snakes.

Frogs of the genus Microhyla include some of the world’s smallest amphibians and represent the largest radiation of Asian microhylids, currently encompassing 50 species, distributed across the Oriental biogeographic region. The genus Microhyla remains one of the taxonomically most challenging groups of Asian frogs and was found to be paraphyletic with respect to large-sized fossorial Glyphoglossus . In this study we present a time-calibrated phylogeny for frogs in the genus Microhyla , and discuss taxonomy, historical biogeography, and morphological evolution of these frogs. Our updated phylogeny of the genus with nearly complete taxon sampling includes 48 nominal Microhyla species and several undescribed candidate species. Phylogenetic analyses of 3,207 bp of combined mtDNA and nuDNA data recovered three well-supported groups: the Glyphoglossus clade, Southeast Asian Microhyla II clade (includes M. annectens species group), and a diverse Microhyla I clade including all other species. Within the largest major clade of Microhyla are seven well-supported subclades that we identify as the M. achatina , M. fissipes , M. berdmorei , M. superciliaris , M. ornata , M. butleri , and M. palmipes species groups. The phylogenetic position of 12 poorly known Microhyla species is clarified for the first time. These phylogenetic results, along with molecular clock and ancestral area analyses, show the Microhyla—Glyphoglossus assemblage to have originated in Southeast Asia in the middle Eocene just after the first hypothesized land connections between the Indian Plate and the Asian mainland. While Glyphoglossus and Microhyla II remained within their ancestral ranges, Microhyla I expanded its distribution generally east to west, colonizing and diversifying through the Cenozoic. The Indian Subcontinent was colonized by members of five Microhyla species groups independently, starting with the end Oligocene—early Miocene that coincides with an onset of seasonally dry climates in South Asia. Body size evolution modeling suggests that four groups of Microhyla have independently achieved extreme miniaturization with adult body size below 15 mm. Three of the five smallest Microhyla species are obligate phytotelm-breeders and we argue that their peculiar reproductive biology may be a factor involved in miniaturization. Body size increases in Microhyla—Glyphoglossus seem to be associated with a burrowing adaptation to seasonally dry habitats. Species delimitation analyses suggest a vast underestimation of species richness and diversity in Microhyla and reveal 15–33 undescribed species. We revalidate M. nepenthicola , synonymize M. pulverata with M. marmorata , and provide insights on taxonomic statuses of a number of poorly known species. Further integrative studies, combining evidence from phylogeny, morphology, advertisement calls, and behavior will result in a better systematic understanding of this morphologically cryptic radiation of Asian frogs.

We describe a large gecko of genus Hemidactylus from the southern face peneplain foothills of the Central Highlands of Sri Lanka. It closely resembles Hemidactylus hunaeDeraniyagala 1937 but is distinguished by adult males reaching 121.2 mm snout–vent length; presence of 11 or 12 supralabials at the midorbit position; dorsal scalation of homogeneous granules intermixed with large, conical, carinate tubercles that form 12–14 irregularly arranged longitudinal rows at midbody; dorsal furrow distinct with a narrow space between medial parasagittal rows; 3 or 4 pairs of postmentals; no spine-like tubercles on nape; ventrals in 36–39 rows at midbody; 21–24 femoral pores on each side separated medially by 5–7 nonpored enlarged scales; scales on posterior thigh granular, not enlarged; lamellae divided, 12 or 13 below the fourth toe; tail segmented with whorls of lateral tubercles, with each whorl consisting of 6 enlarged, conical, carinate tubercles; median row enlarged and broad; single postcloacal tubercle (spur) on each side; and body dorsum with a series of black edged bright saddles from occiput to tail tip. Additionally, we provide a redescription for H. hunae based on its adult female holotype.

Based on phylogenetic and morphological characters, we revise the systematics of the natricid genus Fowlea in Sri Lanka, comprising two morphospecies. The taxonomy of the Sri Lankan populations has long been controversial, and one of the species has, for more than a decade, been listed as Xenochrophis cf. piscator. Although the Sri Lankan populations are morphologically allied to Fowlea piscator in India, they are genetically highly divergent from the eastern Indian F. piscator sensu lato with a p-distance of 9.9–12.3%, and from southwestern Indian Fowlea species with a p-distance of 4.9–11.1% in the mitochondrial cytochrome b gene. Here, we resurrect Müller's (1887) variety, Tropidonotus quincunciatus var. unicolor, as a distinct taxon, elevate it to the species level, and assign it to the genus Fowlea. Therefore, the population so far recognized as X. cf. piscator will be treated hereafter as F. unicolor and we redescribe it and its holotype (by monotypy). We tentatively restrict this species to Sri Lanka and state the possibility of a population in southern India too. The second distinct species, Fowlea asperrima, which is endemic to Sri Lanka, has long been confused with its sympatric congener, F. unicolor comb. nov., and we designate a lectotype and redescribe it herein. Currently, nine species of the genus Fowlea are now recognized, but it is likely that further species (including those regarded as subjective synonyms) remain unrecognized.

AIM: There are several competing hypotheses to explain the high species richness of the Indo‐Australian Archipelago (IAA) marine biodiversity hotspot centred within Southeast (SE) Asia. We use phylogenetic methods to provide a novel perspective on this problem using viviparous sea snakes, a group with high species richness in the IAA that is highly distinct from other taxa previously studied, both phylogenetically (Reptilia, Amniota) and biologically (e.g. viviparity and direct development). LOCATION: Indian Ocean and the West Pacific. METHODS: We used likelihood and Bayesian methods to reconstruct a multi‐locus time‐calibrated phylogeny for c. 70% of viviparous sea snake species, many sampled from multiple localities in Australasia, Southeast Asia and the Indian Ocean. We then compared rates and temporal concordance of inferred vicariance and dispersal events between marine basins using several approaches including new Bayesian analyses that allow for clade‐specific and event‐specific dispersal rates. RESULTS: Phylogenetic analyses and novel Bayesian biogeographical reconstructions indicate that viviparous sea snakes underwent rapid speciation after colonizing SE Asia c. 3 million years ago. Most of the SE Asian sea snake diversity is the result of in situ speciation, most consistent with the ‘centre of origin’ and ‘centre of refuge’ models for biodiversity hotspots. There is also speciation at the periphery, or entirely outside SE Asia; however, contrary to predictions of the ‘accumulation’ and ‘overlap’ models, these new outlying taxa do not preferentially disperse back into SE Asia. Instead, lineages are equally likely to disperse either into or away from SE Asia. MAIN CONCLUSION: The high diversity of sea snakes in SE Asia (and hence the IAA) is mostly explained by in situ speciation rather than accumulation or overlap. Most speciation events are contemporaneous with sea level changes that generated and dissolved barriers between marine basins during the last 2.5 million years.

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 mmSVL) 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.

Comprehensive assessments of species’ extinction risks have documented the extinction crisis 1 and underpinned strategies for reducing those risks 2 . Global assessments reveal that, among tetrapods, 40.7% of amphibians, 25.4% of mammals and 13.6% of birds are threatened with extinction 3 . Because global assessments have been lacking, reptiles have been omitted from conservation-prioritization analyses that encompass other tetrapods 4 – 7 . Reptiles are unusually diverse in arid regions, suggesting that they may have different conservation needs 6 . Here we provide a comprehensive extinction-risk assessment of reptiles and show that at least 1,829 out of 10,196 species (21.1%) are threatened—confirming a previous extrapolation 8 and representing 15.6 billion years of phylogenetic diversity. Reptiles are threatened by the same major factors that threaten other tetrapods—agriculture, logging, urban development and invasive species—although the threat posed by climate change remains uncertain. Reptiles inhabiting forests, where these threats are strongest, are more threatened than those in arid habitats, contrary to our prediction. Birds, mammals and amphibians are unexpectedly good surrogates for the conservation of reptiles, although threatened reptiles with the smallest ranges tend to be isolated from other threatened tetrapods. Although some reptiles—including most species of crocodiles and turtles—require urgent, targeted action to prevent extinctions, efforts to protect other tetrapods, such as habitat preservation and control of trade and invasive species, will probably also benefit many reptiles. An extinction-risk assessment of reptiles shows that at least 21.1% of species are threatened by factors such as agriculture, logging, urban development and invasive species, and that efforts to protect birds, mammals and amphibians probably also benefit many reptiles.

Colour polymorphism is a pervasive phenomenon in both animal and plant kingdoms and understanding its evolution and maintenance is of great interest. Among the lizards of Sri Lanka, the endemic skink Lankascincus fallax shows throat colour polymorphism in which, the underlying basis is not clearly known. In this study, we examined the relationship of the three different throat colour morphs observed in this species with the sex, body size and the geographic location of L. fallax. Live skinks were sampled from two locations in Sri Lanka and sex and the throat colour was categorized visually and the snout to vent length (SVL) was measured. Tail tips of some selected individuals from the two locations were taken and a fragment of the 12S rRNA gene was sequenced in representative individuals having the different throat colour morphs. Pairwise genetic distance of the three colour morphs ranged between 0.4-0.5% confirming that the three colour morphs were the same species. Three colour morphs (red, black and white) were observed in males in both locations, while only the white morph was observed in females, suggesting that the throat color polymorphism was confined to males. There was a significant difference between the mean SVL of males with red and black throat colours (39.35 mm) and males with white throat colours (30.31 mm). Thus, the study suggests that the throat colour in these skinks is highly associated with sex and the body size in males. The study further suggests that L. fallax is sexually dichromatic and that the males show throat colour polymorphism. However, future studies are necessary to understand the underlying drivers for the presence and maintenance of sexual dichromatism and throat colour polymorphism in L. fallax. Competing Interest Statement The authors have declared no competing interest.

Hemipenial characteristics have historically provided a wealth of comparative morphological characters for the systematic classification of snakes. However, the organs remain poorly known in many groups, particularly tropical and burrowing lineages. Here, we report on hemipenial morphology for 12 species from five genera from the family Uropeltidae: Melanophidium punctatum, M. cf. wynaudense, Plectrurus perrotetii, Rhinophis karinthandani, R. melanoleucus, R. saffragamus, R. sanguineus, Teretrurus cf. hewstoni, Uropeltis bhupathyi, U. cf. ceylanica, U. macrolepis, and U. rajendrani. Many are photographed or illustrated here for the first time. In Melanophidium, the organ is bulbous and mushroom-shaped, with the sulcus spermaticus winding through numerous convoluted folds. In Plectrurus and Teretrurus, it is simple, smooth, and conical. In Sri Lankan Rhinophis and some Uropeltis, the organ generally resembles previously described hemipenes from other species in those genera in being simple, subcylindrical, and covered in fine spines. However, a median lobular process is observed in the Indian species R. karinthandani, R. melanoleucus, and R. sanguineus, seemingly representing a novel bilobate morphology. One species, U. bhupathyi, exhibits a novel, bulbous morphology, but this may be an artifact of preservation. The hemipenes of the Uropeltidae and their sister group Cylindrophiidae resemble some typhlopoid blindsnakes more than their henophidian relatives such as pythons and boas. Whether this is due to convergence related to microhabitat, a form of sexual selection unrelated to ecomorphology, or symplesiomorphy from an ancestral snake morphology is unclear. Gross hemipenial morphology can now serve to diagnose uropeltids to the genus level or species group, though more data and comparative series are needed to determine whether other characters, such as the number and location of spines, can potentially differentiate taxa at finer scales.