Comparative skull osteology of Amphisbaena arda and Amphisbaena vermicularis (Squamata: Amphisbaenidae).

The skull anatomy of amphisbaenians directly influences their capacity to burrow and is crucial for the study of their systematics, which ultimately contributes to our comprehension of their evolution and ecology. In this study, we employed three-dimensional X-ray computed tomography to provide a detailed description and comprehensive comparison of the skull anatomy of two amphisbaenian species with similar external morphology, Amphisbaena arda and Amphisbaena vermicularis. Our findings revealed some differences between the species, especially in the sagittal crest of the parietal bone, the ascendant process, and the transverse occipital crest of the occipital complex. We also found intraspecific variation within A. vermicularis, with some specimens displaying morphology that differed from their conspecifics but not from A. arda. The observed intraspecific variation within A. vermicularis cannot be attributed to soil features because all specimens came from the same locality. Specimen size and soil type may play a role in the observed differences between A. arda and A. vermicularis, as the single A. arda specimen is the largest of our sample and soil type and texture differ between the collection sites of the two species.

Comparing morphological and secretory aspects of cephalic glands among the New World coral snakes brings novel insights on their biological roles.

Oral and other cephalic glands have been surveyed by several studies with distinct purposes. Despite the wide diversity and medical relevance of the New World coral snakes, studies focusing on understanding the biological roles of the glands within this group are still scarce. Specifically, the venom glands of some coral snakes were previously investigated but all other cephalic glands remain uncharacterized. In this sense, performing morphological and molecular analysis of these glands may help better understand their biological role. Here, we studied the morphology of the venom, infralabial, rictal, and harderian glands of thirteen species of Micrurus and Micruroides euryxanthus. We also performed a molecular characterization of these glands from selected species of Micrurus using transcriptomic and proteomic approaches. We described substantial morphological variation in the cephalic glands of New World coral snakes and structural evidence for protein-secreting cells in the inferior rictal glands. Our molecular analysis revealed that the venom glands, as expected, are majorly devoted to toxin production, however, the infralabial and inferior rictal glands also expressed some toxin genes at low to medium levels, despite the marked morphological differences. On the other hand, the harderian glands were dominated by the expression of lipocalins, but do not produce toxins. Our integrative analysis, including the prediction of biological processes and pathways, helped decipher some important traits of cephalic glands and better understand their biology.

Genomic evidence reveals intraspecific divergence of the hot-spring snake (Thermophis baileyi), an endangered reptile endemic to the Qinghai-Tibet plateau.

Understanding how and why species evolve requires knowledge on intraspecific divergence. In this study, we examined intraspecific divergence in the endangered hot-spring snake (Thermophis baileyi), an endemic species on the Qinghai-Tibet Plateau (QTP). Whole-genome resequencing of 58 sampled individuals from 15 populations was performed to identify the drivers of intraspecific divergence and explore the potential roles of genes under selection. Our analyses resolved three groups, with major intergroup admixture occurring in regions of group contact. Divergence probably occurred during the Pleistocene as a result of glacial climatic oscillations, Yadong-Gulu rift, and geothermal fields differentiation, while complex gene flow between group pairs reflected a unique intraspecific divergence pattern on the QTP. Intergroup fixed loci involved selected genes functionally related to divergence and local adaptation, especially adaptation to hot spring microenvironments in different geothermal fields. Analysis of structural variants, genetic diversity, inbreeding, and genetic load indicated that the hot-spring snake population has declined to a low level with decreased diversity, which is important for the conservation management of this endangered species. Our study demonstrated that the integration of demographic history, gene flow, genomic divergence genes, and other information is necessary to distinguish the evolutionary processes involved in speciation.

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 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.

A morphological and molecular study of Hydrodynastes gigas (Serpentes, Dipsadidae), a widespread species from South America.

BACKGROUND: Studies with integrative approaches (based on different lines of evidence) are fundamental for understanding the diversity of organisms. Different data sources can improve the understanding of the taxonomy and evolution of snakes. We used this integrative approach to verify the taxonomic status of Hydrodynastes gigas (Duméril, Bibron & Duméril, 1854), given its wide distribution throughout South America, including the validity of the recently described Hydrodynastes melanogigas Franco, Fernandes & Bentim, 2007. METHODS: We performed a phylogenetic analysis of Bayesian Inference with mtDNA 16S and Cytb, and nuDNA Cmos and NT3 concatenated (1,902 bp). In addition, we performed traditional morphometric analyses, meristic, hemipenis morphology and coloration pattern of H. gigas and H. melanogigas. RESULTS: According to molecular and morphological characters, H. gigas is widely distributed throughout South America. We found no evidence to support that H. gigas and H. melanogigas species are distinct lineages, therefore, H. melanogigas is a junior synonym of H. gigas. Thus, the melanic pattern of H. melanogigas is the result of a polymorphism of H. gigas. Melanic populations of H. gigas can be found in the Tocantins-Araguaia basin.

Pythons in the Eocene of Europe reveal a much older divergence of the group in sympatry with boas.

Extant large constrictors, pythons and boas, have a wholly allopatric distribution that has been interpreted largely in terms of vicariance in Gondwana. Here, we describe a stem pythonid based on complete skeletons from the early-middle Eocene of Messel, Germany. The new species is close in age to the divergence of Pythonidae from North American Loxocemus and corroborates a Laurasian origin and dispersal of pythons. Remarkably, it existed in sympatry with the stem boid Eoconstrictor. These occurrences demonstrate that neither dispersal limitation nor strong competitive interactions were decisive in structuring biogeographic patterns early in the history of large, hyper-macrostomatan constrictors and exemplify the synergy between phylogenomic and palaeontological approaches in reconstructing past distributions.

Multilocus phylogeny of Paratelmatobiinae (Anura: Leptodactylidae) reveals strong spatial structure and previously unknown diversity in the Atlantic Forest hotspot.

The Brazilian Atlantic Forest harbors high levels of anuran diversity and endemism, including several taxa restricted to small geographic ranges. Here, we provide a multilocus phylogeny for Paratelmatobiinae, a leptodactylid subfamily composed of small-ranged species distributed in the Brazilian Atlantic Forest and in the campo rupestre ecosystem. We performed Bayesian inference and maximum likelihood analyses using three mitochondrial and five nuclear markers, and a matrix comprising a broad taxonomic sampling. We then delimitated independently evolving lineages within the group. We recovered Paratelmatobiinae and each of its four genera as monophyletic and robustly supported. Five putatively new species included in our analyses were unambiguously supported in the phylogenetic trees and delimitation analyses. We also recovered other deeply divergent and geographically structured lineages within the four genera of Paratelmatobiinae. Our estimation of divergence times indicates that diversification in the subfamily began in the Eocene and continued until the Pleistocene. We discuss possible scenarios of diversification for the four genera of Paratelmatobiinae, and outline the implications of our findings for taxonomy and conservation.

Phylogenomics, Biogeography, and Morphometrics Reveal Rapid Phenotypic Evolution in Pythons After Crossing Wallace's Line.

Ecological opportunities can be provided to organisms that cross stringent biogeographic barriers towards environments with new ecological niches. Wallace's and Lyddeker's lines are arguably the most famous biogeographic barriers, separating the Asian and Australo-Papuan biotas. One of the most ecomorphologically diverse groups of reptiles, the pythons, is distributed across these lines, and are remarkably more diverse in phenotype and ecology east of Lydekker's line in Australo-Papua. We used an anchored hybrid enrichment approach, with near complete taxon sampling, to extract mitochondrial genomes and 376 nuclear loci to resolve and date their phylogenetic history. Biogeographic reconstruction demonstrates that they originated in Asia around 38-45 Ma and then invaded Australo-Papua around 23 Ma. Australo-Papuan pythons display a sizeable expansion in morphological space, with shifts towards numerous new adaptive optima in head and body shape, coupled with the evolution of new micro-habitat preferences. We provide an updated taxonomy of pythons and our study also demonstrates how ecological opportunity following colonization of novel environments can promote morphological diversification in a formerly ecomorphologically conservative group. [Adaptive radiation; anchored hybrid enrichment; biogeography; morphometrics; snakes.].

Interrogating Genomic-Scale Data for Squamata (Lizards, Snakes, and Amphisbaenians) Shows no Support for Key Traditional Morphological Relationships.

Genomics is narrowing uncertainty in the phylogenetic structure for many amniote groups. For one of the most diverse and species-rich groups, the squamate reptiles (lizards, snakes, and amphisbaenians), an inverse correlation between the number of taxa and loci sampled still persists across all publications using DNA sequence data and reaching a consensus on the relationships among them has been highly problematic. In this study, we use high-throughput sequence data from 289 samples covering 75 families of squamates to address phylogenetic affinities, estimate divergence times, and characterize residual topological uncertainty in the presence of genome-scale data. Importantly, we address genomic support for the traditional taxonomic groupings Scleroglossa and Macrostomata using novel machine-learning techniques. We interrogate genes using various metrics inherent to these loci, including parsimony-informative sites (PIS), phylogenetic informativeness, length, gaps, number of substitutions, and site concordance to understand why certain loci fail to find previously well-supported molecular clades and how they fail to support species-tree estimates. We show that both incomplete lineage sorting and poor gene-tree estimation (due to a few undesirable gene properties, such as an insufficient number of PIS), may account for most gene and species-tree discordance. We find overwhelming signal for Toxicofera, and also show that none of the loci included in this study supports Scleroglossa or Macrostomata. We comment on the origins and diversification of Squamata throughout the Mesozoic and underscore remaining uncertainties that persist in both deeper parts of the tree (e.g., relationships between Dibamia, Gekkota, and remaining squamates; among the three toxicoferan clades Iguania, Serpentes, and Anguiformes) and within specific clades (e.g., affinities among gekkotan, pleurodont iguanians, and colubroid families).

The unique traits of the subgenus Unicus within Chiasmocleis Méhely, 1094 (Anura: Microhylidae).

Chiasmocleis is the most speciose genus of Neotropical microhylids. The genus consists of three monophyletic clades that were recently recognized as subgenera. Within Chiasmocleis, the subgenus Unicus has a basal position in the phylogeny and contains a single species that occurs in the North Atlantic Forest of Brazil, isolated from the ranges of other Chiasmocleis. However, the subgenus Unicus lacked a formal description and consequently so far was a nomen nudum. Herein, we provide a diagnosis (i.e., morphological and molecular apomorphies) of the subgenus Unicus de Sá, Tonini, van Huss, Long, Cuddy, Forlani, Peloso, Zaher and Haddad, 2019b; for convenience, the name has also been registered in ZooBank.

Correction: Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes).

[This corrects the article DOI: 10.1371/journal.pone.0216148.].

Large-scale molecular phylogeny, morphology, divergence-time estimation, and the fossil record of advanced caenophidian snakes (Squamata: Serpentes).

Caenophidian snakes include the file snake genus Acrochordus and advanced colubroidean snakes that radiated mainly during the Neogene. Although caenophidian snakes are a well-supported clade, their inferred affinities, based either on molecular or morphological data, remain poorly known or controversial. Here, we provide an expanded molecular phylogenetic analysis of Caenophidia and use three non-parametric measures of support-Shimodaira-Hasegawa-Like test (SHL), Felsentein (FBP) and transfer (TBE) bootstrap measures-to evaluate the robustness of each clade in the molecular tree. That very different alternative support values are common suggests that results based on only one support value should be viewed with caution. Using a scheme to combine support values, we find 20.9% of the 1265 clades comprising the inferred caenophidian tree are unambiguously supported by both SHL and FBP values, while almost 37% are unsupported or ambiguously supported, revealing the substantial extent of phylogenetic problems within Caenophidia. Combined FBP/TBE support values show similar results, while SHL/TBE result in slightly higher combined values. We consider key morphological attributes of colubroidean cranial, vertebral and hemipenial anatomy and provide additional morphological evidence supporting the clades Colubroides, Colubriformes, and Endoglyptodonta. We review and revise the relevant caenophidian fossil record and provide a time-calibrated tree derived from our molecular data to discuss the main cladogenetic events that resulted in present-day patterns of caenophidian diversification. Our results suggest that all extant families of Colubroidea and Elapoidea composing the present-day endoglyptodont fauna originated rapidly within the early Oligocene-between approximately 33 and 28 Mya-following the major terrestrial faunal turnover known as the "Grande Coupure" and associated with the overall climate shift at the Eocene-Oligocene boundary. Our results further suggest that the caenophidian radiation originated within the Caenozoic, with the divergence between Colubroides and Acrochordidae occurring in the early Eocene, at ~ 56 Mya.

A new species of the Amphisbaena (Squamata, Amphisbaenidae) from the Brazilian Cerrado with a key for the two-pored species.

Here, we describe a new species of Amphisbaena with two precloacal pores from open Cerrado areas of the municipality of Arenópolis, in the Brazilian state of Goiás. The new species differs from other South American amphisbaenids by the folllowing combination of characters: (1) snout rounded in dorsal view and slightly convex in lateral view; (2) two precloacal pores; (3) 161-176 dorsal half-annuli; and (4) 12-15 tail annuli. Our molecular phylogenetic analysis retrieved a monophyletic Amphisbaena silvestrii group, with A. silvestrii positioned as the sister-group of a clade formed by Amphisbaena anaemariae and the new species described herein. Members of the A. silvestrii group including A. neglecta and A. crisae not added in our phylogenetic analysis are characterized by a relatively small body, two precloacal pores, body coloration with dark and light areas, and lack of specializations on the cephalic or caudal shields. We present a key for two-pored species of Amphisbaena.

Amphisbaena mensae Catro-Mello, 2000 is a synonym of Amphisbaena talisiae Vanzolini, 1995 (Squamata: Amphisbaenia: Amphisbaenidae).

Amphisbaena talisiae and A. mensae, two worm lizard species endemic to the Cerrado ecoregion, in central Brazil, are considered synonyms based on morphological characters. With the proposed synonymy, the name A. talisiae has priority of use over A. mensae. Amphisbaena talisiae can be distinguished from its congeners by a series of morphological characters, including a round head, three supralabial and three infralabial scales, postmalar row absent, four precloacal pores without a median hiatus, 205-234 body annuli, 17-29 caudal annuli, 10-14 dorsal and 14-18 ventral segments in a midbody annulus. The species is recorded from southeastern Mato Grosso, eastern Minas Gerais and central Tocantins states in central Brazil, and its conservation status should be changed from Data Deficient to Least Concern.

A new species of Ischnocnema Reinhardt and Lütken, 1862 (Anura: Brachycephalidae) of the I. lactea species series from southeastern Brazil.

We describe a new species of Ischnocnema from the Serra da Bocaina mountain range, state of São Paulo, southeastern Brazil, based on morphological, bioacoustic, and mtDNA data. The new species is retrieved with high support values within the I. lactea species series as the sister species of I. spanios. Ischnocnema bocaina sp. nov. is characterized by its medium size (18.6-19.0 mm), a smooth venter, a rounded snout in dorsal view and acuminate in lateral view, a slightly expanded subgular, single vocal sac, a round and whitish, poorly-developed glandular-appearing nuptial pad on the dorsal surface of the thumb, and a nonpulsed advertisement call with 9 to 18 notes. We raise to 38 the number of Ischnocnema species, the 12th described in the past 10 years.

Multiple connections between Amazonia and Atlantic Forest shaped the phylogenetic and morphological diversity of Chiasmocleis Mehely, 1904 (Anura: Microhylidae: Gastrophryninae).

Chiasmocleis is the most species-rich genus of Neotropical microhylids. Herein, we provide the first comprehensive multilocus phylogeny for the genus, including all but 3 of the 34 recognized species and multiple individuals per species. We discuss cryptic speciation, species discovery, patterns of morphological evolution, and provide a historical biogeographic analysis to account for the current distribution of the genus. Diversification of Chiasmocleis from other New World microhylids began during the Eocene, app. 40 mya, in forested areas, and current diversity seems to be a product of recurrent connections between the Atlantic Forest and Amazonia. Small-sized species evolved independently three times in Chiasmocleis. Furthermore, the extremely small-bodied (i.e. miniaturized) species with associated loss of digits, phalanges, and pectoral girdle cartilages evolved only once and are restricted to Amazonia. Using the phylogeny, we recognized three subgenera within Chiasmocleis: Chiasmocleis Méhely, 1904, Relictus subg. nov., and Syncope Walker, 1973. The recognition of the subgenus Syncope informs future research on patterns of miniaturization in the genus, and the subgenus Relictus highlights isolation of an endemic and species-poor lineage to the Atlantic Forest, early (about 40 mya) in the history of Chiasmocleis.

A new species of Apostolepis (Serpentes, Dipsadidae, Elapomorphini) from the Cerrado of Central Brazil.

We describe a new species of the genus Apostolepis found near São Salvador Hydroelectric Power Plant, municipality of São Salvador do Tocantins, in the central Cerrado ecoregion. The new species is distinguished from its congeners by the following combination of characters: presence of eleven dorsolateral stripes, five infralabial scales and the lack of preoculars scales. The new species shares some external characteristics mainly with A. arenaria, A. gaboi and A. nelsonjorgei, with which it is compared.