The oldest known snakes from the Middle Jurassic-Lower Cretaceous provide insights on snake evolution

The previous oldest known fossil snakes date from ∼100 million year old sediments (Upper Cretaceous) and are both morphologically and phylogenetically diverse, indicating that snakes underwent a much earlier origin and adaptive radiation. We report here on snake fossils that extend the record backwards in time by an additional ∼70 million years (Middle Jurassic-Lower Cretaceous). These ancient snakes share features with fossil and modern snakes (for example, recurved teeth with labial and lingual carinae, long toothed suborbital ramus of maxillae) and with lizards (for example, pronounced subdental shelf/gutter). The paleobiogeography of these early snakes is diverse and complex, suggesting that snakes had undergone habitat differentiation and geographic radiation by the mid-Jurassic. Phylogenetic analysis of squamates recovers these early snakes in a basal polytomy with other fossil and modern snakes, where Najash rionegrina is sister to this clade. Ingroup analysis finds them in a basal position to all other snakes including Najash.Fil: Caldwell, Michael Wayne. University of Alberta; CanadáFil: Nydam, Randall L.. Department Of Anatomy, Midwestern University, Glendale; Estados UnidosFil: Palci, Alessandro. South Australian Museum. Earth Sciences Section; AustraliaFil: Apesteguía, Sebastián. Fundación de Historia Natural Félix de Azara; Argentina. Universidad Maimónides. Área de Investigaciones Biomédicas y Biotecnológicas. Centro de Estudios Biomédicos, Biotecnológicos, Ambientales y de Diagnóstico; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentin

The morphology of the inner ear of squamate reptiles and its bearing on the origin of snakes

The inner ear morphology of 80 snake and lizard species, representative of a range of ecologies, is here analysed and compared to that of the fossil stem snake Dinilysia patagonica, using three-dimensional geometric morphometrics. Inner ear morphology is linked to phylogeny (we find here a strong phylogenetic signal in the data that can complicate ecological correlations), but also correlated with ecology, with Dinilysia resembling certain semi-fossorial forms (Xenopeltis and Cylindrophis), consistent with previous reports. We here also find striking resemblances between Dinilysia and some semi-aquatic snakes, such as Myron (Caenophidia, Homalopsidae). Therefore, the inner ear morphology of Dinilysia is consistent with semi-aquatic as well as semi-fossorial habits: the most similar forms are either semi-fossorial burrowers with a strong affinity to water (Xenopeltis and Cylindrophis) or amphibious, intertidal forms which shelter in burrows (Myron). Notably, Dinilysia does not cluster as closely with snakes with exclusively terrestrial or obligate burrowing habits (e.g. scolecophidians and uropeltids). Moreover, despite the above similarities, Dinilysia also occupies a totally unique morphospace, raising issues with linking it with any particular ecological category.Alessandro Palci, Mark N. Hutchinson, Michael W. Caldwell and Michael S.Y. Le

On the Origin and Evolution of the Ophidia

With well over 3,400 described species, snakes undoubtedly represent one of the most successful groups of reptiles. Much has been written about their ecology, behavior, anatomy, relationships and evolution. However, despite the debate about the origin of this taxonomic group dating back to the second half of the XIX century, no consensus has been reached, yet. Scenarios that portray the first snakes as evolving from aquatic lacertilian ancestors are countered with others that see the first snakes as the result of long-term adaptations to a burrowing, cryptic lifestyle. The supporters of the first type of scenario found their evidence mostly in osteological comparisons of non-burrowing snakes with extinct aquatic lizards (e.g., mosasaurs, dolichosaurs and adriosaurs), while the supporters of the second type of scenario base their conclusions mostly on anatomical comparisons between modern legless squamates and burrowing snakes. The debate is further complicated by the scarcity of well-preserved fossil remains that may help elucidate the origin of the group, and by the contradictory interpretations that different authors have provided after examination of the same fossil specimens. Therefore, the goal of this work was that of analyzing all the evidence that has been put forward so far in support of the two main origin scenarios, critically evaluate the contradictory evidence provided by different researchers, examine first-hand all the pivotal extant and fossil taxa that have been used in the formulation and support of each scenario, and provide a phylogenetic analysis of snakes within squamate reptiles that is based both on molecular and morphological data. The examination of over 400 specimens of squamates, including both extant and fossil species, lead to the following results: (1) the redescription of several important fossil taxa (Pachyrhachis, Eupodophis, Haasiophis, Najash, and Dinilysia), and the identification, in some of them, of anatomical features never reported before (e.g., chevron bones in Haasiophis, mental foramina and a sacral vertebra in Pachyrhachis); (2) the discovery of new material (a pelvic girdle) attributable to the fossil species Wonambi naracoortensis, a snake whose pelvic anatomy was previously unknown; (3) the retrieval of evidence that supports a reinterpretation of the circumorbital bones of snakes, with particular regard to the “postorbital” and the “supraorbital”, here reinterpreted as primary homologues of the jugal and postfrontal, respectively; (4) a detailed assessment of what constitutes the “crista circumfenestralis” of snakes and how this anatomical feature varies within the Ophidia; (5) new hypotheses regarding the ingroup relationships of snakes, which imply a possible convergent evolution of the macrostomatan skull condition, and the possibility that scolecophidians may represent an aberrant lineage of alethinophidian snakes

Shape and size variation in elapid snake fangs, and the effects of phylogeny and diet

Published online: 9 October 2023Recent studies have found correlations between the shape of snake teeth/fangs and diet. These studies were done at a very broad phylogenetic scale, making it desirable to test if correlations are still detectable at a narrower evolutionary scale, specifically within the family Elapidae. To this end, we studied fang shape in a dense selection of elapids representing most genera worldwide (74%). We used three-dimensional geometric morphometrics to analyse fang diversity and evaluate possible correlations between fang shape, fang size, and diet. We detected only weak phylogenetic signal in our dataset for both shape and size, and no significant evolutionary allometry when correcting for phylogeny. Overall, the distribution of elapid fangs in morphospace was found to be surprisingly conservative, with only a few outliers. The only two dietary categories that were found to have a significant effect on fang shape are fish and snakes, while mammals have a significant effect on absolute but not relative fang size. Our results show that there are disparate patterns in fang-diet relationships at different evolutionary scales. Across all venomous snakes, previous work found that fangs are strongly influenced by diet, but within elapids our study shows these same associations are weaker and often non-significant. This could result from limitations in these types of studies, or could reflect the fact that elapids are a relatively young clade, where recent extensive divergences in diet have yet to be mirrored in fang shape, suggesting a lag between changes in ecology and dental morphology.Alessandro Palci, Michael S. Y. Lee, Jenna M. Crowe, Riddell, Emma Sherrat

Palaeoecological inferences for the fossil Australian snakes yurlunggur and wonambi (Serpentes, madtsoiidae)

Madtsoiids are among the most basal snakes, with a fossil record dating back to the Upper Cretaceous (Cenomanian). Most representatives went extinct by the end of the Eocene, but some survived in Australia until the Late Cenozoic. Yurlunggur and Wonambi are two of these late forms, and also the best-known madtsoiids to date. A better understanding of the anatomy and palaeoecology of these taxa may shed light on the evolution and extinction of this poorly known group of snakes and on early snake evolution in general. A digital endocast of the inner ear of Yurlunggur was compared to those of 81 species of snakes and lizards with known ecological preferences using three-dimensional geometric morphometrics. The inner ear of Yurlunggur most closely resembles both that of certain semiaquatic snakes and that of some semifossorial snakes. Other cranial and postcranial features of this snake support the semifossorial interpretation. While the digital endocast of the inner ear of Wonambi is too incomplete to be included in a geometric morphometrics study, its preserved morphology is very different from that of Yurlunggur and suggests a more generalist ecology. Osteology, palaeoclimatic data and the palaeobiogeographic distribution of these two snakes are all consistent with these inferred ecological differences.Alessandro Palci, Mark N. Hutchinson, Michael W. Caldwell, John D. Scanlon, Michael S. Y. Le

Novel vascular plexus in the head of a sea snake (Elapidae, Hydrophiinae) revealed by high-resolution computed tomography and histology

Novel phenotypes are often linked to major ecological transitions during evolution. Here, we describe for the first time an unusual network of large blood vessels in the head of the sea snake Hydrophis cyanocinctus. MicroCT imaging and histology reveal an intricate modified cephalic vascular network (MCVN) that underlies a broad area of skin between the snout and the roof of the head. It is mostly composed of large veins and sinuses and converges posterodorsally into a large vein (sometimes paired) that penetrates the skull through the parietal bone. Endocranially, this blood vessel leads into the dorsal cerebral sinus, and from there, a pair of large veins depart ventrally to enter the brain. We compare the condition observed in H. cyanocinctus with that of other elapids and discuss the possible functions of this unusual vascular network. Sea snakes have low oxygen partial pressure in their arterial blood that facilitates cutaneous respiration, potentially limiting the availability of oxygen to the brain. We conclude that this novel vascular structure draining directly to the brain is a further elaboration of the sea snakes' cutaneous respiratory anatomy, the most likely function of which is to provide the brain with an additional supply of oxygen.Alessandro Palci, Roger S. Seymour, Cao Van Nguyen, Mark N. Hutchinson, Michael S. Y. Lee and Kate L. Sander

Cretaceous Blind Snake from Brazil Fills Major Gap in Snake Evolution

Blind snakes (Scolecophidia) are minute cryptic snakes that diverged at the base of the evolutionary radiation of modern snakes. They have a scant fossil record, which dates back to the Upper Paleocene-Lower Eocene ( 56 Ma); this late appearance conflicts with molecular evidence, which suggests a much older origin for the group (during the Mesozoic: 160–125 Ma). Here we report a typhlopoid blind snake from the Late Cretaceous of Brazil, Boipeba tayasuensis gen. et sp. nov, which extends the scolecophidian fossil record into the Mesozoic and reduces the fossil gap predicted by molecular data. The new species is estimated to have been over 1 m long, much larger than typical modern scolecophidians (<30 cm). This finding sheds light on the early evolution of blind snakes, supports the hypothesis of a Gondwanan origin for the Typhlopoidea, and indicates that early scolecophidians had large body size, and only later underwent miniaturization.Thiago Schineider Fachini, Silvio Onary, Alessandro Palci, Michael S.Y. Lee, Mario Bronzati, and Annie Schmaltz Hsio

The morphology of the inner ear of squamate reptiles and its bearing on the origin of snakes

Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited.The inner ear morphology of 80 snake and lizard species, representative of a range of ecologies, is here analysed and compared to that of the fossil stem snake Dinilysia patagonica, using three-dimensional geometric morphometrics. Inner ear morphology is linked to phylogeny (we find here a strong phylogenetic signal in the data that can complicate ecological correlations), but also correlated with ecology, with Dinilysia resembling certain semi-fossorial forms (Xenopeltis and Cylindrophis), consistent with previous reports. We here also find striking resemblances between Dinilysia and some semi-aquatic snakes, such as Myron (Caenophidia, Homalopsidae). Therefore, the inner ear morphology of Dinilysia is consistent with semi-aquatic as well as semi-fossorial habits: the most similar forms are either semi-fossorial burrowers with a strong affinity to water (Xenopeltis and Cylindrophis) or amphibious, intertidal forms which shelter in burrows (Myron). Notably, Dinilysia does not cluster as closely with snakes with exclusively terrestrial or obligate burrowing habits (e.g. scolecophidians and uropeltids). Moreover, despite the above similarities, Dinilysia also occupies a totally unique morphospace, raising issues with linking it with any particular ecological category

Reacquisition of the lower temporal bar in sexually dimorphic fossil lizards provides a rare case of convergent evolution

Temporal fenestration has long been considered a key character to understand relationships amongst reptiles. In particular, the absence of the lower temporal bar (LTB) is considered one of the defining features of squamates (lizards and snakes). In a re-assessment of the borioteiioid lizard Polyglyphanodon sternbergi (Cretaceous, North America), we detected a heretofore unrecognized ontogenetic series, sexual dimorphism (a rare instance for Mesozoic reptiles), and a complete LTB, a feature only recently recognized for another borioteiioid, Tianyusaurus zhengi (Cretaceous, China). A new phylogenetic analysis (with updates on a quarter of the scorings for P. sternbergi) indicates not only that the LTB was reacquired in squamates, but it happened independently at least twice. An analysis of the functional significance of the LTB using proxies indicates that, unlike for T. zhengi, this structure had no apparent functional advantage in P. sternbergi, and it is better explained as the result of structural constraint release. The observed canalization against a LTB in squamates was broken at some point in the evolution of borioteiioids, whereas never re-occuring in other squamate lineages. This case of convergent evolution involves a mix of both adaptationist and structuralist causes, which is unusual for both living and extinct vertebrates