The osteology of the Late Triassic reptile Scleromochlus taylori from µCT data.

Scleromochlus taylori is one of the most enigmatic members of the herpetofauna from the Lossiemouth Sandstone Formation (Upper Triassic) of Elgin (Moray, Scotland). For many years it was thought to be closely related to pterosaurs and dinosaurs, but the anatomy of this animal is difficult to interpret because of the notoriously poor preservation of the six available specimens, which comprise void space in the sandstone after the bones were destroyed by diagenesis. Historically, these fossils have been studied using physical molds, which provide only incomplete, and potentially distorted, information. Due to these uncertainties, interpretations of the anatomy, phylogenetic relationships, and paleobiology of Scleromochlus taylori have remained contentious. Here, we use microcomputed tomographic (µCT) techniques to redescribe and illustrate the osteology of Scleromochlus in detail, building upon a short redescription of keystone features of the anatomy that we recently published. We digitally visualize, describe, and figure previously inaccessible-and thus unaltered-portions of its skeleton, as well as providing new observations on the exposed parts of each specimen. This work reveals many novel features of the skull, mandible, trunk, tail, girdles, forelimb, and hindlimb (particularly of the manus, femur, and pes), demonstrating that historic molding techniques failed, in some cases, to accurately capture the anatomy of Scleromochlus. Our review sheds light on some of the most controversial aspects of Scleromochlus morphology showing that this taxon retains plesiomorphic features of Avemetatarsalia in the postcranial skeleton, alongside a suite of synapomorphies diagnostic of pterosauromorphs (the broad clade of pterosaurs and taxa more closely related to them than dinosaurs), particularly one subgroup, the lagerpetids. Consistent with recent work, our updated phylogenetic analyses (Maximum Parsimony and Bayesian Inference) demonstrate that Scleromochlus taylori is an avemetatarsalian archosaur that is recovered firmly in an early diverging position within Pterosauromorpha, as a member of Lagerpetidae, thus shedding important information on the origin of pterosaurs, the first group of vertebrates to evolve powered flight.

Reappraisal of the thalattosuchian crocodylomorph record from the Middle-Upper Jurassic Rosso Ammonitico Veronese of northeastern Italy: Age calibration, new specimens and taphonomic biases.

Despite their extremely rare and fragmentary record, aquatic crocodylomorphs from the Middle to Upper Jurassic (Bajocian-Tithonian) Rosso Ammonitico Veronese (RAV) of northeastern Italy have sparked interest since the late 18th century. Among marine reptiles, Thalattosuchia is by far one of the best represented groups from the RAV units, especially in the Middle Jurassic. Although some specimens have been the subject of multiple studies in recent times, most of them still lack precise stratigraphic assignment and taphonomic assessment, while others remain undescribed. Here we provide a comprehensive revision of the thalattosuchian record from the RAV, alongside the most up-to-date age determination, by means of calcareous nannofossils, when available. Three new metriorhynchoid specimens are described for the first time from the Middle Jurassic of Asiago Plateau (Vicenza province). While the taphonomy of the newly described specimens hampers any taxonomic attribution below superfamily/family level, all three were confidently assigned to a precise interval between the upper Bajocian and the upper Bathonian. This revised record has major paleobiogeographical implications: the new specimens confirm an early origin and distribution of Metriorhynchoidea in the Tethys area and suggest a fast colonization of the open-ocean environment since the upper Bajocian.

The ecological diversification and evolution of Teleosauroidea (Crocodylomorpha, Thalattosuchia), with insights into their mandibular biomechanics.

Throughout the Jurassic, a plethora of marine reptiles dominated ocean waters, including ichthyosaurs, plesiosaurs and thalattosuchian crocodylomorphs. These Jurassic ecosystems were characterized by high niche partitioning and spatial variation in dietary ecology. However, while the ecological diversity of many marine reptile lineages is well known, the overall ecological diversification of Teleosauroidea (one of the two major groups within thalattosuchian crocodylomorphs) has never been explored. Teleosauroids were previously deemed to have a morphologically conservative body plan; however, they were in actuality morphofunctionally more diverse than previously thought. Here we investigate the ecology and feeding specializations of teleosauroids, using morphological and functional cranio-dental characteristics. We assembled the most comprehensive dataset to date of teleosauroid taxa (approximately 20 species) and ran a series of principal component analyses (PC) to categorize them into various feeding ecomorphotypes based on 17 dental characteristics (38 specimens) and 16 functionally significant mandibular characters (18 specimens). The results were examined in conjunction with a comprehensive thalattosuchian phylogeny (153 taxa and 502 characters) to evaluate macroevolutionary patterns and significant ecological shifts. Machimosaurids display a well-developed ecological shift from: (1) slender, pointed tooth apices and an elongate gracile mandible; to (2) more robust, pointed teeth with a slightly deeper mandible; and finally, (3) rounded teeth and a deep-set, shortened mandible with enlarged musculature. Overall, there is limited mandibular functional variability in teleosaurids and machimosaurids, despite differing cranial morphologies and habitat preferences in certain taxa. This suggests a narrow feeding ecological divide between teleosaurids and machimosaurids. Resource partitioning was primarily related to snout and skull length as well as habitat; only twice did teleosauroids manage to make a major evolutionary leap to feed distinctly differently, with only the derived machimosaurines successfully radiating into new feeding ecologies.

Scleromochlus and the early evolution of Pterosauromorpha.

Pterosaurs, the first vertebrates to evolve powered flight, were key components of Mesozoic terrestrial ecosystems from their sudden appearance in the Late Triassic until their demise at the end of the Cretaceous1-6. However, the origin and early evolution of pterosaurs are poorly understood owing to a substantial stratigraphic and morphological gap between these reptiles and their closest relatives6, Lagerpetidae7. Scleromochlus taylori, a tiny reptile from the early Late Triassic of Scotland discovered over a century ago, was hypothesized to be a key taxon closely related to pterosaurs8, but its poor preservation has limited previous studies and resulted in controversy over its phylogenetic position, with some even doubting its identification as an archosaur9. Here we use microcomputed tomographic scans to provide the first accurate whole-skeletal reconstruction and a revised diagnosis of Scleromochlus, revealing new anatomical details that conclusively identify it as a close pterosaur relative1 within Pterosauromorpha (the lagerpetid + pterosaur clade). Scleromochlus is anatomically more similar to lagerpetids than to pterosaurs and retains numerous features that were probably present in very early diverging members of Avemetatarsalia (bird-line archosaurs). These results support the hypothesis that the first flying reptiles evolved from tiny, probably facultatively bipedal, cursorial ancestors1.

Ecological signal in the size and shape of marine amniote teeth.

Amniotes have been a major component of marine trophic chains from the beginning of the Triassic to present day, with hundreds of species. However, inferences of their (palaeo)ecology have mostly been qualitative, making it difficult to track how dietary niches have changed through time and across clades. Here, we tackle this issue by applying a novel geometric morphometric protocol to three-dimensional models of tooth crowns across a wide range of raptorial marine amniotes. Our results highlight the phenomenon of dental simplification and widespread convergence in marine amniotes, limiting the range of tooth crown morphologies. Importantly, we quantitatively demonstrate that tooth crown shape and size are strongly associated with diet, whereas crown surface complexity is not. The maximal range of tooth shapes in both mammals and reptiles is seen in medium-sized taxa; large crowns are simple and restricted to a fraction of the morphospace. We recognize four principal raptorial guilds within toothed marine amniotes (durophages, generalists, flesh cutters and flesh piercers). Moreover, even though all these feeding guilds have been convergently colonized over the last 200 Myr, a series of dental morphologies are unique to the Mesozoic period, probably reflecting a distinct ecosystem structure.

Novel track morphotypes from new tracksites indicate increased Middle Jurassic dinosaur diversity on the Isle of Skye, Scotland.

Dinosaur fossils from the Middle Jurassic are rare globally, but the Isle of Skye (Scotland, UK) preserves a varied dinosaur record of abundant trace fossils and rare body fossils from this time. Here we describe two new tracksites from Rubha nam Brathairean (Brothers' Point) near where the first dinosaur footprint in Scotland was found in the 1980s. These sites were formed in subaerially exposed mudstones of the Lealt Shale Formation of the Great Estuarine Group and record a dynamic, subtropical, coastal margin. These tracksites preserve a wide variety of dinosaur track types, including a novel morphotype for Skye: Deltapodus which has a probable stegosaur trackmaker. Additionally, a wide variety of tridactyl tracks shows evidence of multiple theropods of different sizes and possibly hints at the presence of large-bodied ornithopods. Overall, the new tracksites show the dinosaur fauna of Skye is more diverse than previously recognized and give insight into the early evolution of major dinosaur groups whose Middle Jurassic body fossil records are currently sparse.

Revision of the Late Jurassic deep-water teleosauroid crocodylomorph Teleosaurus megarhinus Hulke, 1871 and evidence of pelagic adaptations in Teleosauroidea.

Teleosauroids were a successful group of semi-aquatic crocodylomorphs that were an integral part of coastal marine/lagoonal faunas during the Jurassic. Their fossil record suggests that the group declined in diversity and abundance in deep water deposits during the Late Jurassic. One of the few known teleosauroid species from the deeper water horizons of the well-known Kimmeridge Clay Formation is 'Teleosaurus' megarhinus Hulke, 1871, a poorly studied, gracile longirostrine form. The holotype is an incomplete snout from the Aulacostephanus autissiodorensis Sub-Boreal ammonite Zone of Kimmeridge, England. The only other referred specimen is an almost complete skull from the slightly older A. eudoxus Sub-Boreal ammonite Zone of Quercy, France. Recently, the validity of this species has been called into question. Here we re-describe the holotype as well as the referred French specimen and another incomplete teleosauroid, DORCM G.05067i-v (an anterior rostrum with three osteoderms and an isolated tooth crown), from the same horizon and locality as the holotype. We demonstrate that all specimens are referable to 'Teleosaurus' megarhinus and that the species is indeed a valid taxon, which we assign to a new monotypic genus, Bathysuchus. In our phylogenetic analysis, the latest iteration of the ongoing Crocodylomorph SuperMatrix Project, Bathysuchus megarhinus is found as sister taxon to Aeolodon priscus within a subclade containing Mycterosuchus nasutus and Teleosaurus cadomensis. Notably Bathysuchus has an extreme reduction in dermatocranial ornamentation and osteoderm size, thickness and ornamentation. These features are mirrored in Aeolodon priscus, a species with a well-preserved post-cranial skeleton and a similar shallow and inconspicuous dermal ornamentation. Based on these morphological features, and sedimentological evidence, we hypothesise that the Bathysuchus + Aeolodon clade is the first known teleosauroid lineage that evolved a more pelagic lifestyle.

The long-term ecology and evolution of marine reptiles in a Jurassic seaway.

Marine reptiles flourished in the Mesozoic oceans, filling ecological roles today dominated by crocodylians, large fish, sharks and cetaceans. Many groups of these reptiles coexisted for over 50 million years (Myr), through major environmental changes. However, little is known about how the structure of their ecosystems or their ecologies changed over millions of years. We use the most common marine reptile fossils-teeth-to establish a quantitative system that assigns species to dietary guilds and then track the evolution of these guilds over the roughly 18-million-year history of a single seaway, the Jurassic Sub-Boreal Seaway of the United Kingdom. Groups did not significantly overlap in guild space, indicating that dietary niche partitioning enabled many species to live together. Although a highly diverse fauna was present throughout the history of the seaway, fish and squid eaters with piercing teeth declined over time while hard-object and large-prey specialists diversified, in concert with rising sea levels. High niche partitioning and spatial variation in dietary ecology related to sea depth also characterize modern marine tetrapod faunas, indicating a conserved ecological structure of the world's oceans that has persisted for over 150 Myr.

Evidence of macrophagous teleosaurid crocodylomorphs in the Corallian Group (Oxfordian, Late Jurassic) of the UK.

Teleosaurids were a group of semi-aquatic crocodylomorphs with a fossil record that spanned the Jurassic Period. In the UK, abundant specimens are known from the Oxford Clay Formation (OCF, Callovian to lower Oxfordian), but are very rare in the Kimmeridge Clay Formation (KCF, Kimmeridgian to lower Tithonian), despite their abundance in some contemporaneous deposits in continental Europe. Unfortunately, due to the paucity of material from the intermediate 'Corallian Gap' (middle to upper Oxfordian), we lack an understanding of how and why teleosaurid taxic abundance and diversity declined from the OCF to the KCF. The recognition of an incomplete teleosaurid lower jaw from the Corallian of Weymouth (Dorset, UK) begins to rectify this. The vertically oriented dentition, blunt tooth apices, intense enamel ornamentation that shifts to an anastomosed pattern apically, and deep reception pits on the dentary unambiguously demonstrates the affinity of this specimen with an unnamed sub-clade of macrophagous/durophagous teleosaurids ('Steneosaurus' obtusidens + Machimosaurus). The high symphyseal tooth count allows us to exclude the specimen from M. hugii and M. mosae, but in absence of more diagnostic material we cannot unambiguously assign DORCM G.3939 to a more specific level. Nevertheless, this specimen represents the first mandibular material referable to Teleosauridae from the poorly sampled middle-upper Oxfordian time-span in the UK.

Dental ontogeny and replacement in Pliosauridae.

Dental morphology and patterns of tooth replacement in representatives of the clade Pliosauridae (Reptilia, Sauropterygia) are evaluated in detail. The jaws of one basal (Thalassiodracon hawkinsii) and two derived species (Pliosaurus carpenteri, Pliosaurus kevani) were visualized by µCT scans, and the ontogenetic patterns, or 'movement paths', of replacement teeth could be mapped. Other specimens (Peloneustes philarchus and Pliosaurus westbuyensis) with well-preserved jaws containing functional and replacement teeth in situ were also examined directly, and waves of tooth replacement could be inferred from the degree of in situ tooth development and the fusion between functional and replacement alveoli. The analysis revealed symmetrical tooth eruption over the medial axis throughout the length of the jaw in the basal pliosaurid Thalassiodracon. By contrast, symmetrical tooth eruption patterns occur only along the anterior sections of the jaws of derived pliosaurids. In Pliosaurus, replacement schedules differ in the anterior and posterior portions of the jaws and appear to correlate with differences in tooth morphology and symmetrical replacement. The anterior teeth exhibit longer replacement cycle periods and symmetrical replacement, while shorter cycle periods and asymmetry are seen posteriorly. A longer period suggests slower replacement and is characteristic of large, specialized caniniform teeth in the longer snouted Late Jurassic taxa. Smaller posterior teeth have a shorter period and therefore a faster replacement cycle. The transition from long to short replacement period over the length of the jaw is thought to account for the loss of symmetry. This differentiation could relate to differential tooth function and a type of heterodonty. We therefore propose a new model of pliosaurid tooth replacement patterns and present it in a phylogenetic context.

The cranial osteology of Tyrannoneustes lythrodectikos (Crocodylomorpha: Metriorhynchidae) from the Middle Jurassic of Europe.

Tyrannoneustes lythrodectikos is one of numerous metriorhynchid crocodylomorph species known from the Oxford Clay Formation of England (Callovian-Oxfordian; Middle-Late Jurassic). This taxon is of evolutionary importance, as it is the oldest and most basal known macrophagous metriorhynchid. It has a mosaic of plesiomorphic and derived feeding related characteristics, including: teeth with microscopic, poorly formed and non-contiguous denticles; increased tooth apicobasal length; ventrally displaced dentary tooth row (increased gape); reduced dentary tooth count; and a proportionally long mandibular symphysis. However the type specimen, and current referred specimens, all lack a preserved cranium. As such, the craniofacial morphology of this taxon, and its potential feeding ecology, remains poorly understood. Here we describe two skulls and two lower jaws which we refer to T. lythrodectikos. Previously these specimens were referred to 'Metriorhynchus' brachyrhynchus. They share with the T. lythrodectikos holotype: the in-line reception pits on the dentary, dorsal margin of the surangular is strongly concave in lateral view, and the most of the angular ventral margin is strongly convex. Based on our description of these specimens, the skull of T. lythrodectikos has three autapomorphies: very long posterior processes of the premaxilla terminating in line with the 4th or 5th maxillary alveoli, deep lateral notches on the lateral surface of the maxillary with reception pits for dentary teeth, and the premaxilla forms the anterior margin of the first maxillary alveoli. Our description of the cranial anatomy of Tyrannoneustes lythrodectikos confirms that some macrophagous characteristics evolved during the Middle Jurassic, and were not exclusive to the clade Geosaurini. Moreover, the skulls further highlight the mosaic nature of Tyrannoneustes lythrodectikos and wide-gape macrophagous evolution in Geosaurinae.

Functional anatomy and feeding biomechanics of a giant Upper Jurassic pliosaur (Reptilia: Sauropterygia) from Weymouth Bay, Dorset, UK.

Pliosaurs were among the largest predators in Mesozoic seas, and yet their functional anatomy and feeding biomechanics are poorly understood. A new, well-preserved pliosaur from the Kimmeridgian of Weymouth Bay (UK) revealed cranial adaptations related to feeding. Digital modelling of computed tomography scans allowed reconstruction of missing, distorted regions of the skull and of the adductor musculature, which indicated high bite forces. Size-corrected beam theory modelling showed that the snout was poorly optimised against bending and torsional stresses compared with other aquatic and terrestrial predators, suggesting that pliosaurs did not twist or shake their prey during feeding and that seizing was better performed with post-symphyseal bites. Finite element analysis identified biting-induced stress patterns in both the rostrum and lower jaws, highlighting weak areas in the rostral maxillary-premaxillary contact and the caudal mandibular symphysis. A comparatively weak skull coupled with musculature that was able to produce high forces, is explained as a trade-off between agility, hydrodynamics and strength. In the Kimmeridgian ecosystem, we conclude that Late Jurassic pliosaurs were generalist predators at the top of the food chain, able to prey on reptiles and fishes up to half their own length.

Complex rostral neurovascular system in a giant pliosaur.

Pliosaurs were a long-lived, ubiquitous group of Mesozoic marine predators attaining large body sizes (up to 12 m). Despite much being known about their ecology and behaviour, the mechanisms they adopted for prey detection have been poorly investigated and represent a mystery to date. Complex neurovascular systems in many vertebrate rostra have evolved for prey detection. However, information on the occurrence of such systems in fossil taxa is extremely limited because of poor preservation potential. The neurovascular complex from the snout of an exceptionally well-preserved pliosaur from the Kimmeridgian (Late Jurassic, c. 170 Myr ago) of Weymouth Bay (Dorset, UK) is described here for the first time. Using computed tomography (CT) scans, the extensive bifurcating neurovascular channels could be traced through the rostrum to both the teeth and the foramina on the dorsal and lateral surface of the snout. The structures on the surface of the skull and the high concentrations of peripheral rami suggest that this could be a sensory system, perhaps similar to crocodile pressure receptors or shark electroreceptors.

Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia).

Machimosaurus was a large-bodied genus of teleosaurid crocodylomorph, considered to have been durophagous/chelonivorous, and which frequented coastal marine/estuarine ecosystems during the Late Jurassic. Here, we revise the genus based on previously described specimens and revise the species within this genus. We conclude that there were three European Machimosaurus species and another taxon in Ethiopia. This conclusion is based on numerous lines of evidence: craniomandibular, dental and postcranial morphologies; differences in estimated total body length; geological age; geographical distribution; and hypothetical lifestyle. We re-diagnose the type species Machimosaurus hugii and limit referred specimens to only those from Upper Kimmeridgian-Lower Tithonian of Switzerland, Portugal and Spain. We also re-diagnose Machimosaurus mosae, demonstrate that it is an available name and restrict the species to the uppermost Kimmeridgian-lowermost Tithonian of northeastern France. We re-diagnose and validate the species Machimosaurus nowackianus from Harrar, Ethiopia. Finally, we establish a new species, Machimosaurus buffetauti, for the Lower Kimmeridgian specimens of France and Germany (and possibly England and Poland). We hypothesize that Machimosaurus may have been analogous to the Pliocene-Holocene genus Crocodylus in having one large-bodied taxon suited to traversing marine barriers and additional, geographically limited taxa across its range.

Tooth serration morphologies in the genus Machimosaurus (Crocodylomorpha, Thalattosuchia) from the Late Jurassic of Europe.

Machimosaurus was a large-bodied durophagous/chelonivorous genus of teleosaurid crocodylomorph that lived in shallow marine and brackish ecosystems during the Late Jurassic. Among teleosaurids, Machimosaurus and its sister taxon 'Steneosaurus' obtusidens are characterized by having foreshortened rostra, proportionally enlarged supratemporal fenestrae and blunt teeth with numerous apicobasal ridges and a shorter anastomosed ridged pattern in the apical region. A recent study on 'S.' obtusidens dentition found both true denticles and false serrations (enamel ridges which contact the carinae). Here, we comprehensively describe and figure the dentition of Machimosaurus, and find that Machimosaurus buffetauti and Machimosaurus hugii have four types of serration or serration-like structures, including both denticles and false denticles on the carinae. The denticles are irregularly shaped and are not always discrete units, whereas the false denticles caused by the interaction between the superficial enamel ridges and the carinae are restricted to the apical region. Peculiarly, the most 'denticle-like' structures are discrete, bulbous units on the apicobasal and apical anastomosed ridges of M. hugii. These 'pseudo-denticles' have never, to our knowledge, previously been reported among crocodylomorphs, and their precise function is unclear. They may have increased the surface area of the apical region and/or strengthened the enamel, both of which would have been advantageous for a durophagous taxon feeding on hard objects such as turtles.