The macroevolutionary singularity of snakes.

Snakes and lizards (Squamata) represent a third of terrestrial vertebrates and exhibit spectacular innovations in locomotion, feeding, and sensory processing. However, the evolutionary drivers of this radiation remain poorly known. We infer potential causes and ultimate consequences of squamate macroevolution by combining individual-based natural history observations (>60,000 animals) with a comprehensive time-calibrated phylogeny that we anchored with genomic data (5400 loci) from 1018 species. Due to shifts in the dynamics of speciation and phenotypic evolution, snakes have transformed the trophic structure of animal communities through the recurrent origin and diversification of specialized predatory strategies. Squamate biodiversity reflects a legacy of singular events that occurred during the early history of snakes and reveals the impact of historical contingency on vertebrate biodiversity.

Anti-HIV Activity of Snake Venom Phospholipase A2s: Updates for New Enzymes and Different Virus Strains.

Since the beginning of the HIV epidemic, lasting more than 30 years, the main goal of scientists was to develop effective methods for the prevention and treatment of HIV infection. Modern medicines have reduced the death rate from AIDS by 80%. However, they still have side effects and are very expensive, dictating the need to search for new drugs. Earlier, it was shown that phospholipases A2 (PLA2s) from bee and snake venoms block HIV replication, the effect being independent on catalytic PLA2 activity. However, the antiviral activity of human PLA2s against Lentiviruses depended on catalytic function and was mediated through the destruction of the viral membrane. To clarify the role of phospholipolytic activity in antiviral effects, we analyzed the anti-HIV activity of several snake PLA2s and found that the mechanisms of their antiviral activity were similar to that of mammalian PLA2. Our results indicate that snake PLA2s are capable of inhibiting syncytium formation between chronically HIV-infected cells and healthy CD4-positive cells and block HIV binding to cells. However, only dimeric PLA2s had pronounced virucidal and anti-HIV activity, which depended on their catalytic activity. The ability of snake PLA2s to inactivate the virus may provide an additional barrier to HIV infection. Thus, snake PLA2s might be considered as candidates for lead molecules in anti-HIV drug development.

No link between population isolation and speciation rate in squamate reptiles.

Rates of species formation vary widely across the tree of life and contribute to massive disparities in species richness among clades. This variation can emerge from differences in metapopulation-level processes that affect the rates at which lineages diverge, persist, and evolve reproductive barriers and ecological differentiation. For example, populations that evolve reproductive barriers quickly should form new species at faster rates than populations that acquire reproductive barriers more slowly. This expectation implicitly links microevolutionary processes (the evolution of populations) and macroevolutionary patterns (the profound disparity in speciation rate across taxa). Here, leveraging extensive field sampling from the Neotropical Cerrado biome in a biogeographically controlled natural experiment, we test the role of an important microevolutionary process-the propensity for population isolation-as a control on speciation rate in lizards and snakes. By quantifying population genomic structure across a set of codistributed taxa with extensive and phylogenetically independent variation in speciation rate, we show that broad-scale patterns of species formation are decoupled from demographic and genetic processes that promote the formation of population isolates. Population isolation is likely a critical stage of speciation for many taxa, but our results suggest that interspecific variability in the propensity for isolation has little influence on speciation rates. These results suggest that other stages of speciation-including the rate at which reproductive barriers evolve and the extent to which newly formed populations persist-are likely to play a larger role than population isolation in controlling speciation rate variation in squamates.

Epidemiological study of snakebite cases in Sikkim: Risk modeling with regard to the habitat suitability of common venomous snakes.

BACKGROUND: Snakebite envenoming is listed as category 'A' Neglected Tropical Disease. To achieve the target of WHO (World Health Organization) 2019, it becomes necessary to understand various attributes associated with snakebite including community awareness, improvisation of medical facilities and to map the potential distribution of venomous snakes responsible for the bite. Hence this study is conducted in Sikkim, India to understand the epidemiology of snakebite in Sikkim. The potential distribution and risk mapping of five common venomous snakes are done for effective management of snakebite cases. METHODS AND FINDINGS: The snakebite cases registered in six district hospitals and four PHCs (Primary Health Centers) of Sikkim were collected from the year 2011 to 2018. Community survey was also conducted to supplement the data. Ecological Niche Modeling (ENM) was performed to predict the potential habitat of five common venomous snakes of Sikkim. The risk modeling of snakebite cases was done at the level of Gram Panchayat Unit (GPU) using Geographically Weighted Regression (GWR) and Ordinary Linear Square (OLS) model. We found higher number of male victims inflicted with snakebite envenomation. The potential distribution of the five venomous snakes showed satisfactory mean AUC (Area under Curve) value. Both the models showed significant positive association of snakebite cases with habitat suitability of the venomous snakes. Hospital data revealed no death cases whereas community data reported 24 deaths. CONCLUSIONS: Death from snakebite reflected in community data but not in hospital data strongly indicates the people's belief in traditional medicine. Though people of Sikkim have rich traditional knowledge, in case of snakebite traditional practices may be ineffective leading to loss of life. Sensitizing people and improving medical facilities along with proper transport facilities in rural areas might significantly reduce the snakebite casualties in the state.

A new species of Gerrhopilus (family: Gerrhopilidae), with comments on the taxonomic status of Gerrhopilus ater suturalis (Brongersma).

The genus Gerrhopilus is briefly reviewed with a reevaluation of Gerrhopilus ater suturalis and its elevation to full species. In addition, a new species from Sumatra is described based on a single specimen, one of the three species of Gerrhopilus having both an inferior preocular and inferior ocular, and further distinguishable by the supralabial overlap pattern.

Evolution and dispersal of snakes across the Cretaceous-Paleogene mass extinction.

Mass extinctions have repeatedly shaped global biodiversity. The Cretaceous-Paleogene (K-Pg) mass extinction caused the demise of numerous vertebrate groups, and its aftermath saw the rapid diversification of surviving mammals, birds, frogs, and teleost fishes. However, the effects of the K-Pg extinction on the evolution of snakes-a major clade of predators comprising over 3,700 living species-remains poorly understood. Here, we combine an extensive molecular dataset with phylogenetically and stratigraphically constrained fossil calibrations to infer an evolutionary timescale for Serpentes. We reveal a potential diversification among crown snakes associated with the K-Pg mass extinction, led by the successful colonisation of Asia by the major extant clade Afrophidia. Vertebral morphometrics suggest increasing morphological specialisation among marine snakes through the Paleogene. The dispersal patterns of snakes following the K-Pg underscore the importance of this mass extinction event in shaping Earth's extant vertebrate faunas.

The status of the anomalepidid snake Liotyphlops albirostris and the revalidation of three taxa based on morphology and ecological niche models.

Liotyphlops is a genus of blindsnakes distributed in Central and South America. We reviewed specimens of Liotyphlops albirostris along its current distribution range and, based on morphological data and ecological niche modeling analyses, we restrict the geographical range of L. albirostris and validate three previously described species. In this revision, we describe the morphological variation in the populations from Panamá, Colombia, Ecuador, and Venezuela, and propose a new taxonomic arrangement. We revalidate three previous synonyms of L. albirostris to full species status, while dividing the populations from Colombia in two subspecies-one attributed to a previously recognized species from the Caribbean region, and a new one from the Andean region. The new species differs from L. albirostris from Panamá in cephalic scale arrangements that effectively reduces the previously reported variability of these scales in L. albirostris. We also explore some osteological differences that are congruent with the variation observed. We hope that the recognition of these new species better represents the diversity within Liotyphlops, helping to bring these new species out of their cryptic status so that they will be considered in future conservation efforts.

Impact of Repetitive DNA Elements on Snake Genome Biology and Evolution.

The distinctive biology and unique evolutionary features of snakes make them fascinating model systems to elucidate how genomes evolve and how variation at the genomic level is interlinked with phenotypic-level evolution. Similar to other eukaryotic genomes, large proportions of snake genomes contain repetitive DNA, including transposable elements (TEs) and satellite repeats. The importance of repetitive DNA and its structural and functional role in the snake genome, remain unclear. This review highlights the major types of repeats and their proportions in snake genomes, reflecting the high diversity and composition of snake repeats. We present snakes as an emerging and important model system for the study of repetitive DNA under the impact of sex and microchromosome evolution. We assemble evidence to show that certain repetitive elements in snakes are transcriptionally active and demonstrate highly dynamic lineage-specific patterns as repeat sequences. We hypothesize that particular TEs can trigger different genomic mechanisms that might contribute to driving adaptive evolution in snakes. Finally, we review emerging approaches that may be used to study the expression of repetitive elements in complex genomes, such as snakes. The specific aspects presented here will stimulate further discussion on the role of genomic repeats in shaping snake evolution.

Burrowing in blindsnakes: A preliminary analysis of burrowing forces and consequences for the evolution of morphology.

Burrowing is a common behavior in vertebrates. An underground life-style offers many advantages but also poses important challenges including the high energetic cost of burrowing. Scolecophidians are a group of morphologically derived subterranean snakes that show great diversity in form and function. Although it has been suggested that leptotyphlopids and anomalepidids mostly use existing underground passageways, typhlopids are thought to create their own burrows. However, the mechanisms used to create burrows and the associated forces that animals may be able to generate remain unknown. Here, we provide the first data on push forces in scolecophidians and compare them with those in some burrowing alethinophidian snakes. Our results show that typhlopids are capable of generating higher forces for a given size than other snakes. The observed differences are not due to variation in body diameter or length, suggesting fundamental differences in the mechanics of burrowing or the way in which axial muscles are used. Qualitative observations of skull and vertebral shape suggest that the higher forces exerted by typhlopids may have impacted the evolution of their anatomy. Our results provide the basis for future studies exploring the diversity of form and function in this fascinating group of animals. Quantitative comparisons of the cranial and vertebral shape in addition to collecting functional and ecological data on a wider array of species would be particularly important to test the patterns described here.

On the identity, nomenclatural status and authorship of Coluber monticolus Cantor, 1839 (Reptilia: Serpentes).

Currently, Coluber monticolus Cantor is considered a junior synonym of the Common kukri snake, Oligodon arnensis Shaw. In this work, we analyzed long time disregarded, unpublished original drawings and manuscripts concerning South Asian snakes. Additionally we recall the hitherto little-noticed contact and exchange between B. H. Hodgson and T. E. Cantor. We correct erroneous assumptions regarding the authorship and identity of C. monticolus and, in accordance with Art. 23.9 of the "Code" (ICZN 1999), we declare Dipsas multifasciata Blyth a nomen protectum with priority over Coluber monticolus Hodgson in Cantor (nomen oblitum).

A new species of Psammophis (Serpentes: Psammophiidae) from the Turpan Basin in northwest China.

An adult sand snake specimen was collected during a herpetofaunal survey conducted in the Turpan Basin in northwest China. Phylogenetic analyses revealed that this specimen, along with other snake sloughs and skins collected from different localities in the Turpan Basin formed a clade that is sister to Psammophis lineolatus. This taxon exhibited substantial divergence from its congeners (P. lineolatus and P. condanarus) with uncorrelated p-distances ranging from 11.9 ± 0.9% to 15.8 ± 1.6% for the ND4 gene and from 10.2 ± 0.8% to 13.8 ± 1.1% for the Cytb gene. Given the genetic differences along with morphological differences, we describe the specimen from the Turpan Basin as Psammophis turpanensis sp. nov. We provide detailed morphological descriptions, and compare this specimen with five Asian sand snakes and the Afro-Asian Sand Snake, P. schokari. In addition, we provide brief comments on the biogeography of Psammophis in China.

Contributions to ophiology in Zootaxa 20012020: patterns and trends.

There are currently 3,900 recognized, extant snake species belonging to 529 genera globally (Uetz et al. 2021; this study), making snakes one of the most diverse major groups of squamates. Of the 665 currently recognized species that were described between 2001 and 2020 (a ~17% increase in total species), ~34% of these (226 species) were described in Zootaxa. This number does not include species resurrected from synonymy. The other ~66% (439) species were described in 105 other journals, bulletins or books (Fig.1a). Overall, the number of new snake species described every year is gradually increasing, and 40% of the new species described since 2011 were published in Zootaxa. Following Zootaxa, the second ranked journal, with 37 described species since 2001, is Herpetologica (Fig. 2). Anecdotally, the choice of Zootaxa as a publication outlet for new species descriptions by most authors is based on speed of publication post-acceptance, publication free of charge, relatively unconstrained number of papers published per year, relatively unconstrained manuscript length, expert section editors and reviewers, and consolidated scientometric parameters.

A return-on-investment approach for prioritization of rigorous taxonomic research needed to inform responses to the biodiversity crisis.

Global biodiversity loss is a profound consequence of human activity. Disturbingly, biodiversity loss is greater than realized because of the unknown number of undocumented species. Conservation fundamentally relies on taxonomic recognition of species, but only a fraction of biodiversity is described. Here, we provide a new quantitative approach for prioritizing rigorous taxonomic research for conservation. We implement this approach in a highly diverse vertebrate group-Australian lizards and snakes. Of 870 species assessed, we identified 282 (32.4%) with taxonomic uncertainty, of which 17.6% likely comprise undescribed species of conservation concern. We identify 24 species in need of immediate taxonomic attention to facilitate conservation. Using a broadly applicable return-on-investment framework, we demonstrate the importance of prioritizing the fundamental work of identifying species before they are lost.

Evaluating spatiotemporal dynamics of snakebite in Sri Lanka: Monthly incidence mapping from a national representative survey sample.

BACKGROUND: Snakebite incidence shows both spatial and temporal variation. However, no study has evaluated spatiotemporal patterns of snakebites across a country or region in detail. We used a nationally representative population sample to evaluate spatiotemporal patterns of snakebite in Sri Lanka. METHODOLOGY: We conducted a community-based cross-sectional survey representing all nine provinces of Sri Lanka. We interviewed 165 665 people (0.8% of the national population), and snakebite events reported by the respondents were recorded. Sri Lanka is an agricultural country; its central, southern and western parts receive rain mainly from Southwest monsoon (May to September) and northern and eastern parts receive rain mainly from Northeast monsoon (November to February). We developed spatiotemporal models using multivariate Poisson process modelling to explain monthly snakebite and envenoming incidences in the country. These models were developed at the provincial level to explain local spatiotemporal patterns. PRINCIPAL FINDINGS: Snakebites and envenomings showed clear spatiotemporal patterns. Snakebite hotspots were found in North-Central, North-West, South-West and Eastern Sri Lanka. They exhibited biannual seasonal patterns except in South-Western inlands, which showed triannual seasonality. Envenoming hotspots were confined to North-Central, East and South-West parts of the country. Hotspots in North-Central regions showed triannual seasonal patterns and South-West regions had annual patterns. Hotspots remained persistent throughout the year in Eastern regions. The overall monthly snakebite and envenoming incidences in Sri Lanka were 39 (95%CI: 38-40) and 19 (95%CI: 13-30) per 100 000, respectively, translating into 110 000 (95%CI: 107 500-112 500) snakebites and 45 000 (95%CI: 32 000-73 000) envenomings in a calendar year. CONCLUSIONS/SIGNIFICANCE: This study provides information on community-based monthly incidence of snakebites and envenomings over the whole country. Thus, it provides useful insights into healthcare decision-making, such as, prioritizing locations to establish specialized centres for snakebite management and allocating resources based on risk assessments which take into account both location and season.

Snakebite Envenoming Diagnosis and Diagnostics.

Snakebite envenoming is predominantly an occupational disease of the rural tropics, causing death or permanent disability to hundreds of thousands of victims annually. The diagnosis of snakebite envenoming is commonly based on a combination of patient history and a syndromic approach. However, the availability of auxiliary diagnostic tests at the disposal of the clinicians vary from country to country, and the level of experience within snakebite diagnosis and intervention may be quite different for clinicians from different hospitals. As such, achieving timely diagnosis, and thus treatment, is a challenge faced by treating personnel around the globe. For years, much effort has gone into developing novel diagnostics to support diagnosis of snakebite victims, especially in rural areas of the tropics. Gaining access to affordable and rapid diagnostics could potentially facilitate more favorable patient outcomes due to early and appropriate treatment. This review aims to highlight regional differences in epidemiology and clinical snakebite management on a global scale, including an overview of the past and ongoing research efforts within snakebite diagnostics. Finally, the review is rounded off with a discussion on design considerations and potential benefits of novel snakebite diagnostics.

ToxCodAn: a new toxin annotator and guide to venom gland transcriptomics.

MOTIVATION: Next-generation sequencing has become exceedingly common and has transformed our ability to explore nonmodel systems. In particular, transcriptomics has facilitated the study of venom and evolution of toxins in venomous lineages; however, many challenges remain. Primarily, annotation of toxins in the transcriptome is a laborious and time-consuming task. Current annotation software often fails to predict the correct coding sequence and overestimates the number of toxins present in the transcriptome. Here, we present ToxCodAn, a python script designed to perform precise annotation of snake venom gland transcriptomes. We test ToxCodAn with a set of previously curated transcriptomes and compare the results to other annotators. In addition, we provide a guide for venom gland transcriptomics to facilitate future research and use Bothrops alternatus as a case study for ToxCodAn and our guide. RESULTS: Our analysis reveals that ToxCodAn provides precise annotation of toxins present in the transcriptome of venom glands of snakes. Comparison with other annotators demonstrates that ToxCodAn has better performance with regard to run time ($>20x$ faster), coding sequence prediction ($>3x$ more accurate) and the number of toxins predicted (generating $>4x$ less false positives). In this sense, ToxCodAn is a valuable resource for toxin annotation. The ToxCodAn framework can be expanded in the future to work with other venomous lineages and detect novel toxins.

The original description and date of publication of emCylindrophis/em emlineatus/em and remarks on its type locality (Reptilia: Squamata: Cylindrophiidae).

The snake genus Cylindrophis Wagler, 1828 belongs to the  monogeneric family Cylindrophiidae comprising 15 species distributed predominately throughout SE Asia, with one extralimital species occurring in Sri Lanka (Bernstein et al. 2020: 535). Cylindrophis lineatus is a rare species known from only eight museum specimens (discussed herein), and a photograph of one live individual from Kuching, Sarawak, East Malaysia (Stuebing et al. 2014: 63). Despite being originally described with Singapore as the type-locality (Blanford 1881: 217, 218), it is currently understood that C. lineatus is endemic to western Sarawak, East Malaysia (Stuebing et al. 2014: 63). Wallach et al. (2014: 204) stated that C. lineatus is also found in Kalimantan, but did not provide any references. This appears to have been followed by Bernstein et al. (2020: 537), who provide a map indicating C. lineatus occurrences in Kalimantan. The original description of Cylindrophis lineatus is conventionally cited as Blanford (1881: 217, 218, pl. 20). Herein, we demonstrate that the authorship and date of publication of this taxon should correctly be Cylindrophis lineatus Dennys, 1880b, and discuss that the type locality should be changed to "Borneo".

Two new species and a new country record of the genus emAchalinus/em (Reptilia: Squamata: Xenodermidae) from China.

Combining the results from morphological and molecular analyses, we explore the taxonomy of the genus Achalinus from Southwest China. As a result, we describe two new species, A. panzhihuaensis sp. nov. and A. yangdatongi sp. nov. from southern Sichuan and southern Yunnan provinces, respectively, and we record a new country record, A. emilyae, from Guangxi Zhuang A. R.. The mitochondrial genealogy suggests that A. panzhihuaensis sp. nov. is sister to A. meiguensis, while A. yangdatongi sp. nov. clusters with the sister species A. juliani and A. ater. Both new species show considerable genetic divergence from their recognized congeners (uncorrected p-distance 6.2 % in COI gene). Furthermore, both new species can be diagnosed from closely related congeners by a combination of pholidosis characters. With our discovery, we provide a revised key to the 13 species from China and discuss some of the remaining issues regarding the taxonomy of the genus in China.

A new species of emThamnodynastes/em Wagler, 1830 from western Amazonia, with notes on morphology for members of the emThamnodynastes/em empallidus/em group (Serpentes, Dipsadidae, Tachymenini).

The genus Thamnodynastes is the most diverse within the tribe Tachymenini, with an extensive and complex taxonomic history. The brief descriptions and lack of robust diagnostic characters are the main sources for identification errors and for the difficulty to assess the diversity estimates of the genus. The Thamnodynastes pallidus group was briefly designated to encompass the most arboreal species of the genus, with thinner bodies and longer tails: T. pallidus, T. longicaudus, T. sertanejo, and a fourth undescribed species. After its designation, no other paper addressed this group and its morphological variation, especially for the hemipenis, is still undetermined. After the analysis of all species of Thamnodynastes we were able to corroborate the distinctiveness of the T. pallidus group and to accurately diagnose its fourth species from the western portion of the Amazonia lowlands. The new species is distinguishable from all congeners, except T. sertanejo, by the absence of ventral longitudinal stripes, 17/17/11 dorsal scale rows, and dorsal dark brown blotches on the anterior third of the body. The new species is distinguished from T. sertanejo by the higher number of subcaudals, lower number of ventrals, and smaller body and head sizes. We also provide additional diagnostic features for the Thamnodynastes pallidus group, including new data on hemipenial variation. Finally, we briefly discuss the defensive behavior and morphological characters associated with arboreality in members of the T. pallidus species group.

India's biogeographic history through the eyes of blindsnakes- filling the gaps in the global typhlopoid phylogeny.

The Indian subcontinent's unique geological history is reflected in the diverse assemblage of its biota. The blindsnake superfamily Typhlopoidea, with its unique mix of ancient as well as younger lineages in Asia, provides an opportunity to understand the various biotic exchange scenarios proposed for the Indian landmass. In this study, we aim to understand the biogeographic origins of the four genera of typhlopoids found in India and to decipher their times and modes of arrival in the subcontinent. Five nuclear markers were sequenced for 12 samples collected from across India, encompassing all four genera under study. Published sequences of typhlopoid genera were compiled and combined with Indian sequences to generate a global dataset. Phylogenetic relationships were reconstructed using maximum likelihood and Bayesian inference methods. Divergence times were estimated using BEAST 1.8.2. Ancestral geographical ranges were estimated using DEC + J, implemented in BioGeoBEARS. Divergence time estimates suggest that Gerrhopilus is an ancient lineage, and the lineage leading to it was present on the Indian landmass since the last 100 million years. The other three genera are more recent dispersals into India, possibly trans-oceanic. Biogeographic reconstructions suggest an East Gondwanan origin for Typhlopoidea, an African origin for Grypotyphlops and an Asian origin for Indotyphlops and Argyrophis. It appears that India harbours a combination of ancient and more recently dispersed lineages of typhlopoids. The genus Gerrhopilus is of Gondwanan origin that likely dispersed out of India into Southeast Asia. The other genera are intrusive elements that dispersed into India from Africa (Grypotyphlops) and Asia (Indotyphlops and possibly Argyrophis) post break-up of Gondwana. Thus, our study provides further evidence on the ability of blindsnakes to undergo long distance trans-oceanic dispersal. Results also suggest an Asian origin for typhlopoids from Australasia, Philippines and Wallacea.