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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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Anisodontosaurus greeri is an enigmatic small-bodied tetrapod with a heterodont dentition from the Holbrook Member of the Moenkopi Formation (Middle Triassic) of Arizona (U.S.A.). The evolutionary relationships of this taxon have long been debated and remain uncertain. Using micro-computed tomography (µCT) scans we redescribe the holotype of Anisodontosaurus greeri (UCMP 37804), as well as an additional specimen (MNA.V.1478) that had been informally referred to this taxon. Our new data reveal hidden details of the dentition (i.e., ankylothecodonty and absence of replacement teeth) that, in combination with the tricuspid and mediolaterally expanded crowns, support a referral of both specimens to the archosauromorph clade Trilophosauridae. The referral of MNA.V.1478 to Anisodontosaurus greeri is supported by the unique anatomy of the highly differentiated dentition (i.e., 'figure of 8'-shaped premolariforms in occlusal views; mesiodistally short tooth row; presence of a distal 'molariform' crown). Comparison of Anisodontosaurus greeri with other trilophosaurids highlights marked similarities with Variodens inopinatus from the Upper Triassic deposits of the U.K. Our cladistic analyses confirm these observations, and recover, for the first time, two distinct lineages within Trilophosauridae: one geologically long-lived and comprising Anisodontosaurus and Variodens; and one comprising Tricuspisaurus, Trilophosaurus spp., and Spinosuchus. These results imply that: (i) Anisodontosaurus is one of the oldest known trilophosaurids worldwide and the oldest in North America; (ii) trilophosaurids achieved a broad distribution at low latitudes within western Pangea by the Middle Triassic; (iii) small body sizes were more common than previously thought among trilophosaurids. [ABSTRACT FROM AUTHOR]

The metriorhynchid crocodylomorph fauna of the Late Jurassic Kimmeridge Clay Formation (KCF) of the UK was highly diverse. One genus from this Formation, Torvoneustes , was unique in evolving dentition similar to known chelonivorous crocodylomorphs and enlarged attachment sites for the pterygoideus adductor musculature. Here we report the largest known Torvoneustes specimen, the occipital region of a large cranium that was discovered on the shore of Brandy Bay, Dorset, UK (KCF). We also report three tooth crowns discovered during the 19th century from Oxfordshire that can also be referred to Torvoneustes. The partial braincase is unique in having: verticalized basioccipital tuberosities that have thickened ventral margins, the notch between the basioccipital tuberosities is a narrow inverted 'U'-shape and a subrectangular-shaped carotid canal foramina. The presence of 'occipital fossae' (deep concavities with the hypoglossal foramina in their dorsomedial corners) and carotid foramina with raised rims relative to the basioccipital posterior surface, allow us to refer it to Torvoneustes. Although incomplete, the exceptional size of the specimen demonstrates that Torvoneustes attained larger body lengths than previously supposed (3.7–4.7 m). Comparing the dimensions of this specimen to other metriorhynchids suggests that at least some Torvoneustes specimens rivalled Plesiosuchus manselii in body length. [ABSTRACT FROM AUTHOR]

Metriorhynchids are an extinct group of Jurassic-Cretaceous crocodylomorphs secondarily adapted to a marine lifestyle. A new metriorhynchid crocodylomorph from the Oxford Clay Formation (Callovian, Middle Jurassic) of England is described. The specimen is a large, fragmentary skull and associated single ramus of a lower jaw uniquely preserved in a septarian concretion. The description of the specimen reveals a series of autapomorphies (apicobasal flutings on the middle labial surface of the tooth crowns, greatly enlarged basoccipital tuberosities) and a unique combination of characters that warrant the creation of a new genus and species: Ieldraan melkshamensis gen. et sp. nov. This taxon shares numerous characters with the Late Jurassic-Early Cretaceous genus Geosaurus: tooth crowns that have three apicobasal facets on their labial surface, subtly ornamented skull and lower jaws elements, and reception pits along the lateral margin of the dentary (maxillary overbite). Phylogenetic analysis places this new species as the sister taxon to Geosaurus. The new taxon adds valuable information on the time of origin of the macrophagous subclade Geosaurini, which was initially thought to have evolved and radiated during the Late Jurassic. The presence of Ieldraan melkshamensis, the phylogenetic re-evaluation of Suchodus durobrivensis as a Plesiosuchus sister taxon and recently identified Callovian Dakosaurus-like specimens in the Oxford Clay Formation, indicate that all major Geosaurini lineages originated earlier than previously supposed. This has major implications for the evolution of macropredation in the group. Specifically, we can now demonstrate that the four different forms of true ziphodonty observed in derived geosaurins independently evolved from a single non-functional microziphodont common ancestor. http://zoobank.org/urn:lsid:zoobank.org:pub:7A400670-A339-463F-8CD4-8704745CB312 [ABSTRACT FROM AUTHOR]

Two of the best known Mesozoic marine reptile assemblages can be found in units deposited in the Jurassic Sub-Boreal Seaway of the UK: the late Middle Jurassic Oxford Clay Formation (OCF) and Late Jurassic Kimmeridge Clay Formation (KCF). They record two very differently structured faunas, but understanding the turnover between them is hampered by a gap in the fossil record that spans much of the Oxfordian, the so-called "Corallian gap". We provide a comprehensive review of specimens from the Corallian Group (CG) of the UK, which includes the first descriptions of several fossils, particularly teeth. We demonstrate that there is a severe reduction in observed marine reptile diversity during the Oxfordian, with several Callovian taxa well known from the OCF not persisting into the Corallian strata, including small-to-mid-sized pliosaurids and longirostrine teleosaurids. We do, however, find evidence that at least one member of each key OCF lineage (plesiosauroids, pliosaurids, ichthyosaurs, and thalattosuchians) survived into the Corallian interval, and that one keystone KCF lineage (the Torvoneustes line of metriorhynchid thalattosuchians) was present during this time, indicating an earlier radiation of this group than previously thought. We suggest that faunal turnover between the OCF and KCF may have been driven by environmental perturbations during the Oxfordian, which selectively removed small bodied pliosaurids and longirostrine teleosaurids from the Jurassic Sub-Boreal Seaway, but less affected metriorhynchids, plesiosauroids, and ophthalmosaurid ichthyosaurs. The preferential removal of taxa from the sub-Boreal realm may have helped facilitate the radiation of lineages that became dominant during the Late Jurassic. [ABSTRACT FROM AUTHOR]

Two of the best known Mesozoic marine reptile assemblages can be found in units deposited in the Jurassic Sub- Boreal Seaway of the UK: the late Middle Jurassic Oxford Clay Formation (OCF) and Late Jurassic Kimmeridge Clay Formation (KCF). They record two very differently structured faunas, but understanding the turnover between them is hampered by a gap in the fossil record that spans much of the Oxfordian, the so-called "Corallian gap". We provide a comprehensive review of specimens from the Corallian Group (CG) of the UK, which includes the first descriptions of several fossils, particularly teeth. We demonstrate that there is a severe reduction in observed marine reptile diversity during the Oxfordian, with several Callovian taxa well known from the OCF not persisting into the Corallian strata, including small-to-mid-sized pliosaurids and longirostrine teleosaurids. We do, however, find evidence that at least one member of each key OCF lineage (plesiosauroids, pliosaurids, ichthyosaurs, and thalattosuchians) survived into the Corallian interval, and that one keystone KCF lineage (the Torvoneustes line of metriorhynchid thalattosuchians) was present during this time, indicating an earlier radiation of this group than previously thought. We suggest that faunal turnover between the OCF and KCF may have been driven by environmental perturbations during the Oxfordian, which selectively removed small bodied pliosaurids and longirostrine teleosaurids from the Jurassic Sub-Boreal Seaway, but less affected metriorhynchids, plesiosauroids, and ophthalmosaurid ichthyosaurs. The preferential removal of taxa from the sub-Boreal realm may have helped facilitate the radiation of lineages that became dominant during the Late Jurassic. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

The use of more than one nomenclatural code is becoming increasingly common in some biological sub-disciplines. To minimize nomenclatural instability, we have decided to establish a higher level systematization for Thalattosuchia under both the International Code of Phylogenetic Nomenclature ('PhyloCode') and the International Code of Zoological Nomenclature ('Zoological Code'). We undertook a series of phylogenetic analyses with an expanded dataset to examine the origins of Thalattosuchia within Crocodylomorpha, and determined the clade's diagnostic characters. Based on these analyses, we provide updated diagnoses for Thalattosuchia and its subclades under both the PhyloCode and Zoological Code. We also introduce two new nomina that are regulated under the PhyloCode (Neothalattosuchia and Euthalattosuchia), and the nomen Dakosaurina, which is registered under both nomenclatural codes. Moreover, we introduce PhyloCode-compliant phylogenetic definitions for Thalattosuchia and its subclades. As we cannot reliably discriminate between the positional hypotheses for Thalattosuchia within Crocodylomorpha, the clades' origins are as much of a mystery today as they were over a century ago. However, we hope that using the same diagnostic characters to define the same clades, with the same nomina, under both nomenclatural codes will be an example to others to follow. [ABSTRACT FROM AUTHOR]

As ecosystems recovered from the end-Permian extinction, many new animal groups proliferated in the ensuing Triassic. Among these were the sauropterygians, reptiles that evolved from terrestrial ancestors and transitioned to a marine environment. The first sauropterygians were small, marine-adapted taxa such as pachypleurosaurs, which are known from Middle-Late Triassic deposits, particularly in the Tethyan realm of Europe, and more recently from Lagerstätten in southwestern China. Here we report two pachypleurosaurs from Myanmar, the first Triassic vertebrate fossils from the country. These specimens demonstrate that their entombing rocks in northern Shan State, which have received less study than terrestrial sediments in southern Shan State and whose ages have long been uncertain, are Triassic. The specimens may be among the oldest pachypleurosaurs globally, potentially corroborating biogeographic scenarios that posit an eastern Tethyan origin for pachypleurosaurs, and raise the potential for future discoveries of well-preserved Triassic reptiles in Myanmar. [ABSTRACT FROM AUTHOR]

As ecosystems recovered from the end-Permian extinction, many new animal groups proliferated in the ensuing Triassic. Among these were the sauropterygians, reptiles that evolved from terrestrial ancestors and transitioned to a marine environment. The first sauropterygians were small, marine-adapted taxa such as pachypleurosaurs, which are known from Middle-Late Triassic deposits, particularly in the Tethyan realm of Europe, and more recently from Lagerstätten in southwestern China. Here we report two pachypleurosaurs from Myanmar, the first Triassic vertebrate fossils from the country. These specimens demonstrate that their entombing rocks in northern Shan State, which have received less study than terrestrial sediments in southern Shan State and whose ages have long been uncertain, are Triassic. The specimens may be among the oldest pachypleurosaurs globally, potentially corroborating biogeographic scenarios that posit an eastern Tethyan origin for pachypleurosaurs, and raise the potential for future discoveries of well-preserved Triassic reptiles in Myanmar. [ABSTRACT FROM AUTHOR]

A marine tethysuchian crocodyliform from the Isle of Wight, most likely from the Upper Greensand Formation (upper Albian, Lower Cretaceous), is described. However, we cannot preclude it being from the Ferruginous Sands Formation (upper Aptian), or more remotely, the Sandrock Formation (upper Aptian-upper Albian). The specimen consists of the anterior region of the right dentary, from the tip of the dentary to the incomplete fourth alveolus. This specimen increases the known geological range of marine tethysuchians back into the late Lower Cretaceous. Although we refer it to Tethysuchia incertae sedis, there are seven anterior dentary characteristics that suggest a possible relationship with the Maastrichtian- Eocene clade Dyrosauridae. We also review 'middle' Cretaceous marine tethysuchians, including putative Cenomanian dyrosaurids. We conclude that there is insufficient evidence to be certain that any known Cenomanian specimen can be safely referred to Dyrosauridae, as there are some cranial similarities between basal dyrosaurids and Cenomanian- Turonian marine 'pholidosaurids'. Future study of middle Cretaceous tethysuchians could help unlock the origins of Dyrosauridae and improve our understanding of tethysuchian macroevolutionary trends. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 854-871. [ABSTRACT FROM AUTHOR]

Metriorhynchidae was a clade of extinct crocodylomorphs that adapted to a pelagic lifestyle, becoming a key component of Mesozoic lagoonal and coastal marine ecosystems. The type genus Metriorhynchus is one of the best-known genera of Mesozoic crocodylomorphs, and since the mid-19th century, the 'concept' of Metriorhynchus has become associated with the referred species Me. superciliosus. Historically Metriorhynchus has been the most species-rich genus in Metriorhynchidae, with most Middle Jurassic species and many Late Jurassic species referred to the genus at some point in their history. However, the type species Me. geoffroyii has largely been omitted in the literature. Its type series is a chimera of multiple metriorhynchid species, and a type specimen has never been designated. Moreover, phylogenetic analyses have repeatedly shown that the 19th–20th century concept of Metriorhynchus is not monophyletic – to the point where only referring every metriorhynchid species, and some basal metriorhynchoids, to the genus would render it monophyletic. Herein we designate a lectotype for Me. geoffroyii , re-describe it and restrict the genus Metriorhynchus to the type species. We also establish the new genus Thalattosuchus for Me. superciliosus , thereby cutting the 'Gordian knot' of Metriorhynchus with Th. superciliosus. [ABSTRACT FROM AUTHOR]

Amongst Mesozoic marine reptiles, metriorhynchid crocodylomorphs were unique in evolving into pelagically adapted forms with little-to-no posterodorsal retraction of the external nares. Narial retraction is a common adaptation seen in sustained swimmers, notably occurring during cetacean evolution. Mesosaurids and the basalmost known members of ichthyosauriforms, thalattosaurians, saurosphargids, sauropterygians, pleurosaurids and mosasauroids had the external nares divided by an ossified bar, bound by multiple cranial bones and were positioned back from the tip of the rostrum. However, metriorhynchids evolved from taxa with a single external naris bound solely by the premaxilla, and positioned near the tip of an elongate rostrum. We posit that metriorhynchids were uniquely disadvantaged in evolving into sustained swimmers. Herein we describe three Late Jurassic metriorhynchid cranial rostra that display differing degrees of narial retraction. In our new phylogenetic analyses, the backwards migration of the narial fossa posterior margin occurred independently at least four times in Metriorhynchidae, whereas the backwards migration of the anterior margin only occurred twice. Although Rhacheosaurini share the backwards migration of the anterior and posterior narial margins, posterodorsal retraction occurred differently along three lineages. This culminated in the Early Cretaceous, where a rhacheosaurin evolved nares bound by the premaxilla and maxilla, and significantly posterodorsally retracted. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Teleosaurids were a clade of crocodylomorphs that attained near-global distribution during the Jurassic Period. Within Teleosauridae, one particular sub-clade of durophagous/macrophagous taxa achieved large body sizes and were apex predators in shallow marine environments during the Late Jurassic and Early Cretaceous in Europe and around the coast of the Tethys Seaway. Unfortunately, the origins of this clade are still poorly understood. 'Steneosaurus' obtusidens is a little-studied macrophagous species from the Oxford Clay Formation (Callovian, Middle Jurassic) of the UK and near Migné-les-Lourdines (Middle Callovian) in France. Despite being considered a sister taxon of the Late Jurassic taxon Machimosaurus, the taxonomy of 'S.' obtusidens remains unclear. Although three different synonymies have been proposed (variously a subjective synonym of other taxa), these taxonomic hypotheses have not been based on detailed anatomical comparisons and thus have not been tested. Here, we re-describe the holotype of 'S.' obtusidens, demonstrate that it is indeed a valid taxon, restrict the referred specimens to a fragmentary skeleton, nearly complete skull, and partial rostrum, and establish a new monotypic genus, Lemmysuchus. Our re-description reveals five autapomorphies for Lemmysuchus obtusidens and nine apomorphic characters that support the tribe Machimosaurini (Lemmysuchus + Machimosaurus). [ABSTRACT FROM AUTHOR]

Abstract: Thalattosuchia was a diverse clade of marine crocodylomorphs known from the Early Jurassic to the Early Cretaceous. Recent studies have hypothesized that their extinction was two-phased: (1) habitat loss near/at the Jurassic-Cretaceous boundary heavily reduced their morphofunctional diversity, particularly in Europe, while (2) climate change and a shift in marine fauna during the Early Cretaceous (either at the Valanginian-Hauterivian boundary or during the early Hauterivian) finished off the already stressed clade. Unfortunately, the Cretaceous fossil record of thalattosuchians is poor, with only one putative “teleosaurid” specimen and approximately ten metriorhynchid specimens. Here we re-describe the youngest known teleosaurid from the Cretaceous (Valanginian of south-eastern France). Originally considered to be a teleosaurid (possibly Steneosaurus), we demonstrate that it belongs to Metriorhynchidae, and a newly discovered subclade, Plesiosuchina. It differs from Plesiosuchus in the pattern of tooth enamel ornamentation and the variation in dentary alveoli size. Referring this specimen to Metriorhynchidae means there are no definitive Cretaceous teleosaurid specimens. Furthermore, it suggests that both durophagous and piscivorous teleosaurids became extinct at the end of the Jurassic. Interestingly, this is the fourth metriorhynchid lineage known to cross the Jurassic-Cretaceous boundary. As such, it would appear that the two thalattosuchian families responded very differently to the lowering sea levels at the end of the Jurassic: teleosaurids possibly became extinct, while metriorhynchids were seemingly unaffected. [Copyright &y& Elsevier]