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Pupil constriction has important functional consequences for animal vision, yet the evolutionary mechanisms underlying diverse pupil sizes and shapes are poorly understood. We aimed to quantify the diversity and evolution of pupil shapes among amphibians and to test for potential correlations to ecology based on functional hypotheses. Using photographs, we surveyed pupil shape across adults of 1294 amphibian species, 74 families and three orders, and additionally for larval stages for all families of frogs and salamanders with a biphasic ontogeny. For amphibians with a biphasic life history, pupil shape changed in many species that occupy distinct habitats before and after metamorphosis. In addition, non-elongated (circular or diamond) constricted pupils were associated with species inhabiting aquatic or underground environments, and elongated pupils (with vertical or horizontal long axes) were more common in species with larger absolute eye sizes. We propose that amphibians provide a valuable group within which to explore the anatomical, physiological, optical and ecological mechanisms underlying the evolution of pupil shape.

We report the rediscovery of Oligodon melaneus 112 years after its original description and document the third, and only non-type, specimen for the species. The new specimen was found 267 km east of the type locality (Tindharia, West Bengal state) from Assam state, India. We designate a lectotype for the species, and provide an extended description of a freshly collected male specimen. Phylogenetic analyses of 16s and cytb mitochondrial genes provide support for O. melaneus being closely related to the widespread South Asian endemic O. arnensis.

Made available in DSpace on 2022-04-29T08:39:58Z (GMT). No. of bitstreams: 0 Previous issue date: 2022-01-01 The spectral characteristics of vertebrate ocular lenses affect the image of the world that is projected onto the retina, and thus help shape diverse visual capabilities. Here, we tested whether amphibian lens transmission is driven by adaptation to diurnal activity (bright light) and/or scansorial habits (complex visual environments). Spectral transmission through the lenses of 79 species of frogs and six species of salamanders was measured, and data for 29 additional frog species compiled from published literature. Phylogenetic comparative methods were used to test ecological explanations of variation in lens transmission and to test for selection across traits. Lenses of diurnal (day-active) and scansorial (climbing) frogs transmitted significantly less shortwave light than those of non-diurnal or non-scansorial amphibians, and evolutionary modelling suggested that these differences have resulted from differential selection. The presence of shortwave-transparent lenses was common among the sampled amphibians, which implies that many are sensitive to shortwave light to some degree even in the absence of visual pigments maximally sensitive in the UV. This suggests that shortwave light, including UV, could play an important role in amphibian behaviour and ecology. Shortwave-absorbing lens pigments likely provide higher visual acuity to diurnally active frogs of multiple ecologies and to nocturnally active scansorial frogs. This new mechanistic understanding of amphibian visual systems suggests that shortwave-filtering lenses are adaptive not only in daylight conditions but also in those scotopic conditions where high acuity is advantageous. Read the free Plain Language Summary for this article on the Journal blog. Department of Life Sciences The Natural History Museum Department of Herpetology California Academy of Sciences Department of Vertebrate Zoology National Museum of Natural History Smithsonian Institution Department of Biology Amphibian and Reptile Diversity Research Center The University of Texas at Arlington Department of Biology York University Ecology Evolution and Development Group Department of Wetland Ecology Estación Biológica de Doñana (CSIC) Departamento de Biodiversidade and Centro de Aquicultura (CAUNESP) I.B. Universidade Estadual Paulista Department of Biology The Pennsylvania State University Department of Biological Sciences Institute for the Environment Florida International University Programa de Pós-graduação em Conservação da Fauna Universidade Federal de São Carlos Division of Optometry & Visual Science School of Health Sciences City University of London Departamento de Biodiversidade and Centro de Aquicultura (CAUNESP) I.B. Universidade Estadual Paulista

The stylommatophoran land-snail genus Corilla is endemic to Sri Lanka and India's Western Ghats. On the basis of habitat distribution and shell morphology, the 10 extant Sri Lankan species fall into two distinct groups, lowland and montane. Here, we use phylogenetic analyses of restriction-site-associated DNA sequencing (RADseq) data and ancestral-state reconstructions of habitat association and shell morphology to clarify the systematics and evolution of Sri Lankan Corilla. Our dataset consists of 9 species of Corilla. Phylogenetic analyses were based on 88 assemblies (9,604-4,132,850 bp) generated by the RADseq assembler ipyrad, using four parameter combinations and different levels of missing data. Trees were inferred using a maximum likelihood (ML) approach. Ancestral states were reconstructed using maximum parsimony (MP) and ML approaches, with 1 binary state character analysed for habitat association (lowland vs montane) and 6 binary state characters analysed for shell morphology (shape, colour, lip width, length of upper palatal folds, orientation of upper palatal folds and collabral sculpture). Over a wide range of missing data (40-87 % missing individuals per locus) and assembly sizes (62,279-4,132,850 bp), nearly all trees conformed to one of two topologies (A and B), most re-lationships were strongly supported and total branch support approached the maximal value. Apart from the position of Corilla odontophora 'south', topologies A and B showed similar, well-resolved relationships at and above the species level. Our study agrees with the shell-based taxonomy of C. adamsi, C. beddomeae, C. carabinata, C. colletti and C. humberti (all maximally supported as monophyletic species). It shows that C. erronea and C. fryae constitute a single relatively widespread species (for which the valid name is C. erronea) and that the names C. gudei and C. odontophora each apply to at least two distinct, yet conchologically-cryptic species. The MP and ML ancestral-state reconstructions yielded broadly similar results and provide firm evi-dence that diversification in Sri Lankan Corilla has involved evolutionary convergence in the shell morphology of lowland lineages, with a pale shell and wide lip having evolved on at least two separate occasions (in C. carabinata and C. colletti) from montane ancestors having a dark, narrow-lipped shell.

Natricine snakes are geographically widespread, species rich (with ~250 extant species) and both morphologically and ecologically diverse. We present a multilocus DNA sequence phylogeny for 249 natricine specimens representing 189 named species, including 69 specimens and 21 species not previously sampled. Our inferred Bayesian and maximum likelihood trees form the basis for evaluations of genus-level classification, historical biogeography, lineage diversification, and dietary, habit and reproductive-mode diversity and evolution, although several, mostly deeper, relationships remain poorly resolved. The optimal trees support natricine origins in Asia, with dispersals to Australo-Melanesia, sub-Saharan Africa (including Seychelles Archipelago, excluding Aldabra), Europe and North Africa and into North and Central America. Viviparity appears to have evolved independently three times in Natricinae but was not significantly associated with an aquatic habit. We found limited associations between habit and diet categories. We propose generic reallocations for four natricine species and highlight other points of uncertainty in natricine classification.

The shape and relative size of an ocular lens affect the focal length of the eye, with consequences for visual acuity and sensitivity. Lenses are typically spherical in aquatic animals with camera-type eyes and axially flattened in terrestrial species to facilitate vision in optical media with different refractive indices. Frogs and toads (Amphibia: Anura) are ecologically diverse, with many species shifting from aquatic to terrestrial ecologies during metamorphosis. We quantified lens shape and relative size using 179 micro X-ray computed tomography scans of 126 biphasic anuran species and tested for correlations with life stage, environmental transitions, adult habits and adult activity patterns. Across broad phylogenetic diversity, tadpole lenses are more spherical than those of adults. Biphasic species with aquatic larvae and terrestrial adults typically undergo ontogenetic changes in lens shape, whereas species that remain aquatic as adults tend to retain more spherical lenses after metamorphosis. Further, adult lens shape is influenced by adult habit; notably, fossorial adults tend to retain spherical lenses following metamorphosis. Finally, lens size relative to eye size is smaller in aquatic and semiaquatic species than other adult ecologies. Our study demonstrates how ecology shapes visual systems, and the power of non-invasive imaging of museum specimens for studying sensory evolution.

Uropeltis dindigalensis (Beddome, 1877) is a poorly known uropeltid (shieldtail) snake from peninsular India. Here we report morphological data for 14 preserved and nine uncollected specimens, most of which have not been previously reported. We designate a lectotype from the type series, describe it, and present the first published photographs of some of the type material. Recharacterization of U. dindigalensis clarifies its taxonomic distinctiveness and its geographical distribution. We report new natural-history observations from museum specimens and of individuals in life, including viviparity and feeding behaviour. The species is known only from the higher elevations of Sirumalai hills, in the southern end of the Eastern Ghats. Uropeltis dindigalensis appears somewhat capable of tolerating low-intensity agriculture. The main immediate conservation threats are probably greater intensity human activities, including road traffic, and possibly climate change given that the species is restricted to the uppermost elevations of a small hill range. The very small extent of occurrence within a single threat-defined location (none of which lies within a protected area), combined with decreasing quality and quantity of habitat, likely qualifies U. dindigalensis as Critically Endangered (or, at best, Endangered) under IUCN Red List criteria.

Non-visual opsins are transmembrane proteins expressed in the eyes, skin, and brain of many animals. When paired with a light-sensitive chromophore, non-visual opsins form photopigment systems involved in various non-visual, light-detection functions, including circadian rhythm regulation, light-seeking behavior, and detection of seasonality. Previous research has primarily explored the diversity and function of non-visual opsins in model organisms, with few studies investigating their molecular evolution in non-model species. Here we explored molecular evolution of non-visual opsin genes in anuran amphibians (frogs and toads). With diverse lifestyles ranging from fossorial to aquatic, anurans inhabit a diverse array of light environments, which makes them a compelling system for studying the evolution of light detection mechanisms. Using whole-eye transcriptomes from 79 anuran species, as well as genomes and multi-tissue transcriptomes from an additional 15 species, we 1) identify which non-visual opsin genes are expressed in the eyes of anurans; 2) compare selective constraint (ω, or dN/dS) among non-visual opsin genes; and 3) test for potential adaptive evolution by comparing selection between discrete ecological classes in anurans (e.g. diurnal versus non-diurnal). We consistently recovered 14 non-visual opsin genes from anuran eye transcriptomes, compared to 18 genes that we recovered genome wide, and detected positive selection in a subset of these genes. We also found variation in selective constraint between discrete ecological and life-history classes, which may reflect functional adaptation in non-visual opsin genes. Although non-visual opsins remain poorly understood, these findings provide insight into their molecular evolution and set the stage for future research on their potential function across taxa with diverse environmental, developmental, and morphological adaptations.

The concept of ecomorphs, whereby species with similar ecologies have similar phenotypes regardless of their phylogenetic relatedness, is often central to discussions regarding the relationship between ecology and phenotype. However, some aspects of the concept have been questioned, and sometimes species have been grouped as ecomorphs based on phenotypic similarity without demonstrating ecological similarity. Within snakes, similar head shapes have convergently evolved in species living in comparable environments and/or with similar diets. Therefore, ecomorphs could exist in some snake lineages, but this assertion has rarely been tested for a wide-ranging group within a single framework. Natricine snakes (Natricinae) are ecomorphologically diverse and currently distributed in Asia, Africa, Europe and north-central America. They are primarily semiaquatic or ground-dwelling terrestrial snakes, but some are aquatic, burrowing or aquatic and burrowing in habit and may be generalist or specialist in diet. Thus, natricines present an interesting system to test whether snakes from different major habit categories represent ecomorphs. We quantify morphological similarity and disparity in head shape among 191 of the ca. 250 currently recognized natricine species and apply phylogenetic comparative methods to test for convergence. Natricine head shape is largely correlated with habit, but in some burrowers is better explained by dietary specialism. Convergence in head shape is especially strong for aquatic burrowing, semiaquatic and terrestrial ecomorphs and less strong for aquatic and burrowing ecomorphs. The ecomorph concept is useful for understanding natricine diversity and evolution, though would benefit from further refinement, especially for aquatic and burrowing taxa.

We describe a new species of Stoliczkia from Mizoram, India. Stoliczkia vanhnuailianai sp. nov. is identified as a member of the genus Stoliczkia by distinct scale arrangements on the posterior of the head, and by scales on the body being separated by scaleless skin, and it differs from the two known congeners in body and head scalation. This is only the third specimen of Stoliczkia collected from India, and the sixth reported specimen for the genus. A revised key to the identification of the species of Stoliczkia is provided.

The world currently recognizes 214 species of Caecilians (Amphibia: Gymnophiona) most of which occur in the wet tropics and some adjacent subtropical regions. Of the ten-family classification known, three occur in Asia, viz. Chikilidae (endemic to northeast India, Indotyphlidae (India’s Western and Eastern Ghats) and Ichthyophiidae. However, until this report, there were no confirmed reports of any caecilian species from Bhutan, although their presence has been assumed likely given their occurrence in adjacent countries. This report provides the first confirmed report of caecilians in Bhutan with work to identify the species to be carried on later with further research.

Animals with biphasic lifecycles often inhabit different visual environments across ontogeny. Many frogs and toads (Amphibia: Anura) have free-living aquatic larvae (tadpoles) that metamorphose into adults that inhabit a range of aquatic and terrestrial environments. Ecological differences influence eye size across species, but these relationships have not yet been explored across life stages in an ontogenetic allometric context. We examined eye-body size scaling in a species with aquatic larvae and terrestrial adults, the common frog Rana temporaria, using a well-sampled developmental series. We found a shift in ontogenetic allometric trajectory near metamorphosis indicating prioritized growth in tadpole eyes. To explore the effects of different tadpole and adult ecologies on eye-body scaling, we expanded our taxonomic sampling to include developmental series of eleven additional anuran species. Intraspecific eye-body scaling was variable among species, with 8/12 species exhibiting a significant change in allometric slope between tadpoles and adults. Traits categorizing both tadpole ecology (microhabitat, eye position, mouth position) and adult ecology (habitat, activity pattern) across species had significant effects on allometric slopes among tadpoles, but only tadpole eye position had a significant effect among adults. Our study suggests that relative eye growth in the preliminary stages of biphasic anuran ontogenies is somewhat decoupled and may be shaped by both immediate ecological need (i.e. tadpole visual requirements) and what will be advantageous during later adult stages.

Urocotyledon inexpectata, a small gecko endemic to the granitic islands of the Seychelles, has previously been demonstrated to comprise two highly distinct clades based on mitochondrial and nuclear DNA sequences, with one lineage exclusive to a northern group of islands, and the second lineage exclusive to the more southerly islands. Here we complement the genetic data with additional analyses to determine if the clades should be considered distinct species. We present and analyse new morphological data, including skull and jaw osteology, and supplement the available genetic data with DNA sequences for individuals from the previously unsampled island of Felicité, which cluster with the other northern island samples. Despite the high morphological similarity between Urocotyledon populations from northern and southern islands, diagnostic characters were identified, with the northern lineage having a more completely ossified skull, more tooth loci on maxilla and dentary, shorter hindlimbs, relatively smaller eye diameter and relatively wider head. We therefore describe the northern lineage as a distinct species, Urocotyledon norzilensis sp. nov., and discuss the biological and conservation management implications of this taxonomic action.

Uropeltidae is a clade of small fossorial snakes (ca. 64 extant species) endemic to peninsular India and Sri Lanka. Uropeltid taxonomy has been confusing, and the status of some species has not been revised for over a century. Attempts to revise uropeltid systematics and undertake evolutionary studies have been hampered by incompletely sampled and incompletely resolved phylogenies. To address this issue, we take advantage of historical museum collections, including type specimens, and apply genome-wide shotgun (GWS) sequencing, along with recent field sampling (using Sanger sequencing) to establish a near-complete multilocus species-level phylogeny (ca. 87% complete at species level). This results in a phylogeny that supports the monophyly of all genera (if Brachyophidium is considered a junior synonym of Teretrurus), and provides a firm platform for future taxonomic revision. Sri Lankan uropeltids are probably monophyletic, indicating a single colonisation event of this island from Indian ancestors. However, the position of Rhinophis goweri (endemic to Eastern Ghats, southern India) is unclear and warrants further investigation, and evidence that it may nest within the Sri Lankan radiation indicates a possible recolonisation event. DNA sequence data and morphology suggest that currently recognised uropeltid species diversity is substantially underestimated. Our study highlights the benefits of integrating museum collections in molecular genetic analyses and their role in understanding the systematics and evolutionary history of understudied organismal groups.

The African natricine genus Limnophis is represented by two species: Limnophis bicolor Gunther, 1865 and Limnophis bangweolicus (Mertens, 1936). They are stout-bodied, semi-aquatic snakes that most...

The Manda Beds of southwest Tanzania have yielded key insights into the early evolutionary radiation of archosaurian reptiles. Many key archosaur specimens were collected from the Manda Beds in the 1930s and 1960s, but until recently, few of these had been formally published. Here, we describe an archosaur specimen collected in 1963 which has previously been referred to informally as Pallisteria angustimentum . We recognize this specimen as the type of a new taxon, Mambawakale ruhuhu gen. et sp. nov. The holotype and only known specimen of M. ruhuhu comprises a partial skull of large size (greater than 75 cm inferred length), lower jaws and fragments of the postcranium, including three anterior cervical vertebrae and a nearly complete left manus. Mambawakale ruhuhu is characterized by several cranial autapomorphies that allow it to be distinguished with confidence from all other Manda Beds archosaurs, with the possible exception of Stagonosuchus nyassicus for which comparisons are highly constrained due to very limited overlapping material. Phylogenetic analysis suggests that M. ruhuhu is an early diverging pseudosuchian, but more precise resolution is hampered by missing data. Mambawakale ruhuhu is one of the largest known pseudosuchians recovered to date from the Middle Triassic.

Molecular genetic data have recently been incorporated in attempts to reconstruct the ecology of the ancestral snake, though this has been limited by a paucity of data for one of the two main extant snake taxa, the highly fossorial Scolecophidia. Here we present and analyze vision genes from the first eye-transcriptomic and genome-wide data for Scolecophidia, for Anilios bicolor, and A. bituberculatus, respectively. We also present immunohistochemistry data for retinal anatomy and visual opsin-gene expression in Anilios. Analyzed in the context of 19 lepidosaurian genomes and 12 eye transcriptomes, the new genome-wide and transcriptomic data provide evidence for a much more reduced visual system in Anilios than in non-scolecophidian (=alethinophidian) snakes and in lizards. In Anilios, there is no evidence of the presence of 7 of the 12 genes associated with alethinophidian photopic (cone) phototransduction. This indicates extensive gene loss and many of these candidate gene losses occur also in highly fossorial mammals with reduced vision. Although recent phylogenetic studies have found evidence for scolecophidian paraphyly, the loss in Anilios of visual genes that are present in alethinophidians implies that the ancestral snake had a better-developed visual system than is known for any extant scolecophidian.

The Manda Beds of southwest Tanzania have yielded key insights into the early evolutionary radiation of archosaurian reptiles. Many key archosaur specimens were collected from the Manda Beds in the 1930s and 1960s, but until recently, few of these had been formally published. Here, we describe an archosaur specimen collected in 1963 which has previously been referred to informally as

A correction to this paper has been published: https://doi.org/10.1007/s10682-021-10109-w

Pupil constriction has important functional consequences for animal vision, yet the evolutionary mechanisms underlying diverse pupil sizes and shapes, often among animals that occupy optically similar environments, are poorly understood. We aimed to quantify the diversity and evolution of pupil shapes among amphibians and test for potential correlations to ecology based on functional hypotheses. Using photographs, we surveyed pupil shape and the orientation of the constricted pupil across adults of 1293 amphibian species, 72 families, and 3 orders, and additionally for larval life stages for all families of frogs and salamanders with a biphasic ontogeny. Pupil shape is exceptionally diverse in amphibians with evolutionary transitions throughout the amphibian tree of life. For amphibians with a biphasic life history, we found that pupils change in many species that occupy distinct habitats before and after metamorphosis. Finally, we found that non-elongated (round or diamond) constricted pupils were correlated with species inhabiting consistently dim light environments (burrowing and aquatic species) and that elongated pupils (vertical and horizontal) were more common in species with larger absolute eye sizes. We propose that amphibians provide a valuable group within which to explore the anatomical, physiological, optical, and ecological mechanisms underlying the evolution of pupil shape.

In the present contribution we revise, figure, and redescribe several isolated braincases of the iconic aetosaur Desmatosuchus from the Placerias Quarry locality, Chinle Formation, Arizona, USA. The detailed study of the isolated braincases from the UCMP collection allowed us to assign them at the species-level and recognize two species of Desmatosuchus for the Placerias Quarry: D. spurensis and D. smalli. The former can be distinguished from the latter by the presence of a transverse sulcus on the parietals, deep median pharyngeal recess on the basisphenoid, almost no gap between the basal tubera and the basipterygoid processes, and the exoccipitals meeting at the midline. The presence of D. smalli at the Placerias Quarry has not been previously reported. Based on the braincases UCMP 27408, 27410, 27407, three new brain endocasts were developed through CT scan images, reconstructing the most complete endocranial casts known for an aetosaur, including the encephalon, cranial nerves, inner ear, and endocranial vasculature. The cranial endocasts also exhibited some differences between both species of Desmatosuchus, with D. spurensis having a distinguishable dural expansion and markedly asymmetric anterior and posterior semicircular canals of the labyrinth. Additionally, the combination of osteological features and the endocranial casts allowed us to identify and discuss the presence of an ossified orbitosphenoid on the anteriormost region of the braincase among aetosaurs. Furthermore, we were able to reinterpret some of the observations made by previous authors on the endocast of the holotype of Desmatosuchus spurensis (UMMP VP 7476) and provide some insight into their neurosensory capabilities. Fil: Von Baczko, Belen. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Desojo, Julia Brenda. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Paleontología Vertebrados; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata; Argentina Fil: Gower, David J.. Natural History Museum; Reino Unido Fil: Ridgely, Ryan. Ohio University; Estados Unidos Fil: Bona, Paula. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. División Paleontología Vertebrados; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata; Argentina Fil: Witmer, Lawrence. Ohio University; Estados Unidos

Background Differences in morphology, ecology, and behavior through ontogeny can result in opposing selective pressures at different life stages. Most animals, however, transition through two or more distinct phenotypic phases, which is hypothesized to allow each life stage to adapt more freely to its ecological niche. How this applies to sensory systems, and in particular how sensory systems adapt across life stages at the molecular level, is not well understood. Here, we used whole-eye transcriptomes to investigate differences in gene expression between tadpole and juvenile southern leopard frogs (Lithobates sphenocephalus), which rely on vision in aquatic and terrestrial light environments, respectively. Because visual physiology changes with light levels, we also tested the effect of light and dark exposure. Results We found 42% of genes were differentially expressed in the eyes of tadpoles versus juveniles and 5% for light/dark exposure. Analyses targeting a curated subset of visual genes revealed significant differential expression of genes that control aspects of visual function and development, including spectral sensitivity and lens composition. Finally, microspectrophotometry of photoreceptors confirmed shifts in spectral sensitivity predicted by the expression results, consistent with adaptation to distinct light environments. Conclusions Overall, we identified extensive expression-level differences in the eyes of tadpoles and juveniles related to observed morphological and physiological changes through metamorphosis and corresponding adaptive shifts to improve vision in the distinct aquatic and terrestrial light environments these frogs inhabit during their life cycle. More broadly, these results suggest that decoupling of gene expression can mediate the opposing selection pressures experienced by organisms with complex life cycles that inhabit different environmental conditions throughout ontogeny.

Identifying individual animals is important for studying populations and for the optimal management of individual animals in captivity. In the absence of natural markings that discriminate individuals, such identification may require animals to be marked by researchers. Amphibians are challenging subjects to mark due to their small size and sensitive, permeable and frequently shed skin. Visible Implant Elastomer (VIE) has been widely used to mark amphibians, but no long-term study has validated this technique in caecilian amphibians. We anaesthetised and attempted to VIE mark seven Herpele squalostoma and one Microcaecilia unicolor held at ZSL London Zoo. No specimens suffered ill effects of anaesthesia or VIE injection, but mean persistence of marks was 191 days in H. squalostoma suggesting that this marking technique is not suitable for identifying individuals of this species in the long-term. We were unable to inject VIE into the M. unicolor and/or the elastomer was not visible through the darkly pigmented skin. Further research is required to develop methods for long-term marking of a diversity of caecilians.

Erythrosuchidae is a clade of early archosauriform reptiles that were large-bodied, hypercarnivorous, possibly apex predators in late Early and Middle Triassic ecosystems following the Permo-Triassic mass extinction. Chalishevia cothurnata from the late Middle Triassic (Ladinian) of Russia, is the stratigraphically youngest known erythrosuchid species, but the holotype and referred material of this taxon has received little study. Here, we provide the first detailed anatomical description of C. cothurnata, including comparisons to other erythrosuchids. Although known from relatively fragmentary material, the anatomy of C. cothurnata is distinctive, including an autapomorphic strongly slanted ventral border of the antorbital fossa. The presence of a large accessory opening (the “accessory antorbital fenestra”) in the skull between the premaxilla, nasal and maxilla, together with the inferred presence of a narrow postnarial process of the premaxilla that articulated with a slot on the nasal, provides strong evidence for a sister taxon relationship between C. cothurnata and the erythrosuchid Shansisuchus shansisuchus from the early Middle Triassic (Anisian) of China. The inferred basal skull length of C. cothurnata was approximately 80 cm, making it one of the largest erythrosuchids known. Fil: Butler, Richard. University of Birmingham; Reino Unido Fil: Sennikov, Andrey. Kazan Federal University. Institute of Geology and Petroleum Technologies; Rusia Fil: Ezcurra, Martin Daniel. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; Argentina Fil: Gower, David. Natural History Museum; Reino Unido