Xenodens calminechari gen. et sp. nov., a bizarre mosasaurid (Mosasauridae, Squamata) with shark-like cutting teeth from the upper Maastrichtian of Morocco, North Africa

The mosasaurids (Mosasauridae) were a group of lizards that became highly specialized for marine life in the mid-Cretaceous. By the end of the Cretaceous, they had undergone an adaptive radiation, and showed a wide range of body sizes, locomotor styles, and diets. Their ranks included piscivores, apex predators, and durophages. Here, we report a new taxon, Xenodens calminechari gen. et sp. nov., from the upper Maastrichtian phosphates of Morocco, with dental specializations unlike those of any known reptile. Teeth form a unique dental battery in which short, laterally compressed and hooked teeth formed a saw-like blade. Unique features of tooth structure and implantation suggest affinities with the durophagous Carinodens. The tooth arrangement seen in Xenodens not only expands known disparity of mosasaurids, but is unique among Squamata, or even Tetrapoda. The specialized dentition implies a previously unknown feeding strategy, likely involving a cutting motion used to carve pieces out of large prey, or in scavenging. This novel dental specialization adds to the already considerable disparity and functional diversity of the late Maastrichtian mosasaurids and marine reptiles. This provides further evidence for a diverse marine fauna just prior to the K-Pg extinction

Stelladens mysteriosus: A Strange New Mosasaurid (Squamata) from the Maastrichtian (Late Cretaceous) of Morocco

Mosasaurids, a clade of specialized marine squamates, saw a major adaptive radiation in the Late Cretaceous, evolving a wide range of body sizes, shapes, and specialized tooth morphologies. The most diverse known mosasaurid faunas come from the late Maastrichtian phosphates of Morocco. Here, we report an unusual new mosasaurid, Stelladens mysteriosus, based on a partial jaw and associated tooth crowns from lower Couche III phosphatic deposits at Sidi Chennane, Oulad Abdoun Basin, Morocco. Stelladens is characterized by short, triangular tooth crowns with a series of strong, elaborate, and serrated ridges on the lingual surface of the tooth, functioning as accessory carinae. Morphology of the teeth and associated jaw fragment suggest affinities with Mosasaurinae. No close analogues to the unique tooth morphology of Stelladens are known, either extant or extinct. It may have had an unusual and highly specialized diet, a specialized prey-capture strategy, or both. The diversity of mosasaurid teeth is much higher than that of plesiosaurs, ichthyosaurs, or extant marine mammals, and likely reflects both the ecological diversity of mosasaurids and complex developmental mechanisms responsible for tooth formation in mosasaurines. Mosasaurid diversity continued to increase up to the Cretaceous–Paleogene boundary.Research of XPS is financed by the Spanish Ministry of Science and Innovation (MCIN) and the European Regional Development Fund (FEDER) (research project PID2021-122612OB-I00), and by the Basque Country Gouvernment (research group IT1485-22). This study is carried out within the framework of the agreement between the universities of Bath and Cadi Ayyad

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

An effect size statistical framework for investigating sexual dimorphism in non-avian dinosaurs and other extinct taxa

Despite reports of sexual dimorphism in extinct taxa, such claims in non-avian dinosaurs have been underrepresented recently (~the last decade) and often criticized. Since dimorphism is widespread in sexually reproducing organisms today, underrepresentation might suggest either methodological shortcomings or that this diverse group exhibited highly unusual reproductive biology. Univariate significance testing, especially for bimodality, is ineffective and prone to false negatives. Species recognition and mutual sexual selection hypotheses, therefore, may not be required to explain supposed absence of sexual dimorphism across the grade, likely a type II error. Instead, multiple lines of evidence support sexual selection and variation of structures consistent with secondary sexual characteristics, strongly suggesting sexual dimorphism in non-avian dinosaurs. We propose a framework for studying sexual dimorphism in fossils, focusing on likely secondarily sexual traits and testing against all alternate hypotheses for variation in them using multiple lines of evidence. We use effect size statistics appropriate for low sample sizes, rather than significance testing, to analyze potential divergence of growth curves in traits and constrain estimates for dimorphism magnitude. In many cases, estimates of sexual variation can be reasonably accurate, and further developments in methods to improve sex assignments and account for intrasexual variation (e.g., mixture modelling) will improve accuracy. It is better to compare estimates for the magnitude of and support for dimorphism between datasets than to dichotomously reject or fail to reject monomorphism in a single species, enabling the study of sexual selection across phylogenies and time. We defend our approach with simulated and empirical data, including dinosaur data, showing that even simple approaches can yield fairly accurate estimates of sexual variation in many cases, allowing for comparison of species with high and low support for sexual variation.Funding provided by: National Science FoundationCrossref Funder Registry ID: http://dx.doi.org/10.13039/100000001Award Number: PLR 1341645 and FRES 192588

Torosaurus Is Not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy

Background: In horned dinosaurs, taxonomy is complicated by the fact that the cranial ornament that distinguishes species changes with age. Based on this observation, it has been proposed that the genera Triceratops and Torosaurus are in fact synonymous, with specimens identified as Torosaurus representing the adult form of Triceratops. The hypothesis of synonymy makes three testable predictions: 1) the species in question should have similar geographic and stratigraphic distributions, 2) specimens assigned to Torosaurus should be more mature than those assigned to Triceratops, and 3) intermediates should exist that combine features of Triceratops and Torosaurus. The first condition appears to be met, but it remains unclear whether the other predictions are borne out by the fossil evidence. Methodology/Principal Findings: We assessed the relative maturity of Torosaurus and Triceratops specimens by coding skulls for characters that vary with maturity, and then using a clustering analysis to arrange them into a growth series. We found that a well-defined sequence of changes exists in horned dinosaurs: development of cranial ornament occurs in juveniles, followed by fusion of the skull roof in subadults, and finally, the epoccipitals, epijugals, and rostral fuse to the skull in adults. Using this scheme, we identified mature and immature individuals of both Torosaurus and Triceratops. Furthermore, we describe the ventral depressions on the frill of Triceratops, and show that they differ in shape and position from the parietal fenestrae of Torosaurus. Thus, we conclude that these structures are not intermediates between the soli

Cranial Ontogeny in Stegoceras validum (Dinosauria: Pachycephalosauria): A Quantitative Model of Pachycephalosaur Dome Growth and Variation

Historically, studies of pachycephalosaurs have recognized plesiomorphically flat-headed taxa and apomorphically domed taxa. More recently, it has been suggested that the expression of the frontoparietal dome is ontogenetic and derived from a flat-headed juvenile morphology. However, strong evidence to support this hypothesis has been lacking. Here we test this hypothesis in a large, stratigraphically constrained sample of specimens assigned to Stegoceras validum, the best known pachycephalosaur, using multiple independent lines of evidence including conserved morphology of ornamentation, landmark-based allometric analyses of frontoparietal shape, and cranial bone histology. New specimens show that the diagnostic ornamentation of the parietosquamosal bar is conserved throughout the size range of the sample, which links flat-headed specimens to domed S. validum. High-resolution CT scans of three frontoparietals reveal that vascularity decreases with size and document a pattern that is consistent with previously proposed histological changes during growth. Furthermore, aspects of dome shape and size are strongly correlated and indicative of ontogenetic growth. These results are complementary and strongly support the hypothesis that the sample represents a growth series of a single taxon. Cranial dome growth is positively allometric, proceeds from a flat-headed to a domed state, and confirms the synonymy of Ornatotholus browni as a juvenile Stegoceras. This dataset serves as the first detailed model of growth and variation in a pachycephalosaur. Flat-headed juveniles possess three characters (externally open cranial sutures, tuberculate dorsal surface texture, and open supratemporal fenestrae) that are reduced or eliminated during ontogeny. These characters also occur in putative flat-headed taxa, suggesting that they may also represent juveniles of domed taxa. However, open cranial sutures and supratemporal fenestrae are plesiomorphic within Ornithischia, and thus should be expected in the adult stage of a primitive pachycephalosaur. Additional lines of evidence will be needed to resolve the taxonomic validity of flat-headed pachycephalosaur taxa