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Here we describe a new genus and species of metriorhynchid crocodylomorph, Enalioetes schroederi gen. et sp. nov., from the lower Valanginian Stadthagen Formation (Lower Cretaceous) of north-western Germany. Enalioetes schroederi is the most complete and well-preserved Cretaceous metriorhynchid skull known to date, preserving most of the cranium and mandible, the atlas-axis complex and the first postaxial cervical vertebra. The specimen was previously attributed to the coeval enigmatic metriorhynchid Enaliosuchus (a nomen dubium) and, more recently, to Cricosaurus. Although the specific epithet schroederi has been used frequently in the literature, it has never been formally established. Herein, we demonstrate that the new taxon is distinct from all known metriorhynchids by a unique combination of characters including several autapomorphies such as: the lack of bulbous dorsolateral expansion in the posterior nasal cavity; mediolateral distance between the orbital canals being approximately 1.5 times the diameter of the orbital canals; lacrimal with dorsoventrally deep anterior process ventral to the preorbital fossa being equal to or greater than the depth of the jugal anterior process and the anterodorsal process of the lacrimal; and ascending processes at atlas intercentrum extending far dorsally to level of neural canal. Enalioetes schroederi contributes to the sparse global record of Cretaceous metriorhynchids and represents one of the stratigraphically youngest occurrences of the group. It can thus help to enhance our understanding of the metriorhynchid diversity during the Cretaceous Period. [ABSTRACT FROM AUTHOR]

Thalattosuchian crocodylomorphs underwent a major evolutionary transition, evolving from semiaquatic forms reminiscent of extant crocodylians, into pelagic marine forms with flippers, a tail fin and smooth scaleless skin. These fully aquatic forms – the Metriorhynchidae – evolved a novel suite of endocranial anatomies hypothesised to be related to living in saltwater. However, much remains to be discovered about the evolution of these internal braincase structures. Herein, we describe the endocranial anatomy of an early diverging metriorhynchid, Thalattosuchus superciliosus, using microfocus computed tomography (μCT) data and three-dimensional modelling. We compared it against geosaurine and metriorhynchine metriorhynchids, as well as the early diverging metriorhynchoid Pelagosaurus. We found that non-geosaurine metriorhynchids differ from geosaurines in having less laterally expanded cerebral hemispheres, shallower curvatures of the brain’s dorsal margin, and lacking the ventral deflection of the pneumatic diverticulum ventral to the pituitary fossa chamber. However, early-diverging metriorhynchids have well-defined otoccipital diverticula and lacked the ‘extreme pelagic’ endosseous labyrinth morphology. We hypothesise that early metriorhynchids were not adapted to a sustained pursuit lifestyle. Moreover, we posit that within both metriorhynchid subfamilies there was parallel evolution towards becoming pursuit predators. [ABSTRACT FROM AUTHOR]

The first paleontological works on Mesozoic vertebrates from Portugal, carried out from the end of the 19th and the first half of the 20th century, provided the discovery of significant collections of vertebrate fossils. These collections are particularly relevant because they include several specimens collected from different regions of the Lusitanian Basin (some of the sites are currently inaccessible), whose fossil record is poorly known. Theropod remains are relatively scarce and generally consist of fragmentary material, mostly assigned to the megalosaurid Megalosaurus from the Middle Jurassic of England, the first dinosaur to be named and a “wastebasket” taxon used by many scientists to identify theropod material. The studied fossils mostly consist of isolated teeth and vertebrae collected from Upper Jurassic levels of the coastal region, with also some material from Lower and Upper Cretaceous strata from the central and northern sectors of the Lusitanian Basin. Here specimens attributed to Megalosaurus from different Portuguese institutions are reviewed and their taxonomic affinity and stratigraphic context are updated. Most specimens actually belong to different theropod groups, including several isolated teeth from different Upper Jurassic localities here assigned to Ceratosaurus, Torvosaurus, and Allosaurus, as well as an isolated tooth from the Lower Cretaceous that is attributed to an indeterminate allosauroid. Other theropod remains consist mostly of vertebral fragments of indeterminate avetheropods and allosauroids. Elements of other dinosaur groups are also represented, including a few vertebrae here referred to stegosaurians and iguanodontians, as well as a vertebra and some appendicular remains attributed to sauropods. Two vertebrae assigned to thalattosuchians were also identified. The study of this collection allows to better characterize the diversity of Late Jurassic dinosaur faunas from different areas of the Lusitanian Basin and provides some data on the poorly known Cretaceous fossil record of theropods from Portugal.

Thalattosuchian crocodylomorphs were a ubiquitous component of shallow marine ecosystems during the Jurassic and Early Cretaceous. Alas, their origins remain a mystery. Here we describe three specimens from the Sinemurian (and possibly Early Pliensbachian) of the UK: a partial cranial rostrum, a series of cervical vertebrae, and two dorsal vertebrae adhered with matrix. These specimens are amongst the oldest known thalattosuchian fossils, with the partial cranial rostrum being the oldest known non-neothalattosuchian thalattosuchian. This partial cranial rostrum has a unique combination of rostral characters never seen before in any crocodylomorph, and helps to elucidate early thalattosuchian internal rostrum evolution, suggesting that the reduction in thalattosuchian paranasal sinuses was not related to either the reorganization of rostral neurovasculature seen in later diverging taxa or the increased cancellous bone microstructure. Based on our CT sample, a shift in cranial bone microstructure occurred in the Eoneustes  + Metriorhynchidae subclade, one that coincided with the enlargement of the salt glands and decoupling of the external antorbital fenestra from the paranasal sinuses. Without extensive histological sampling we cannot determine whether the shift to an obligate aquatic lifestyle occurred prior to the evolution of Metriorhynchidae. [ABSTRACT FROM AUTHOR]

Our knowledge of marine reptiles of the Callovian age (Middle Jurassic) is majorly based on the collections from the Oxford Clay Formation of England, which yielded a diverse marine reptile fauna of plesiosaurians, ichthyosaurians, and thalattosuchians. However, outside of Western Europe, marine reptile remains of this age are poorly known. Here, we survey marine reptiles from the Callovian stage of European Russia. The fossils collected over more than a century from 28 localities are largely represented by isolated bones and teeth, although partial skeletons are also known. In addition to the previously described rhomaleosaurid and metriorhynchids, we identify pliosaurids of the genera Liopleurodon and Simolestes; cryptoclidid plesiosaurians, including Cryptoclidus eurymerus, Muraenosaurus sp., and cf. Tricleidus, and ophthalmosaurid ichthyosaurians, including the iconic Ophthalmosaurus icenicus. These findings expand the ranges of several Callovian marine reptile taxa far to the Eastern Europe, and support the exchange of marine reptile faunas between Western and Eastern European seas in the middle to late Callovian. However, some specimens from the lower Callovian of European Russia show differences from typical representatives of the middle Callovian Oxford Clay fauna, possibly representing the earlier stages of evolution of some of these marine reptiles not yet recorded in Western Europe or elsewhere. [ABSTRACT FROM AUTHOR]

The first paleontological works on Mesozoic vertebrates from Portugal, carried out from the end of the 19th and the first half of the 20th century, provided the discovery of significant collections of vertebrate fossils. These collections are particularly relevant because they include several specimens collected from different regions of the Lusitanian Basin (some of the sites are currently inaccessible), whose fossil record is poorly known. Theropod remains are relatively scarce and generally consist of fragmentary material, mostly assigned to the megalosaurid Megalosaurus from the Middle Jurassic of England, the first dinosaur to be named and a "wastebasket" taxon used by many scientists to identify theropod material. The studied fossils mostly consist of isolated teeth and vertebrae collected from Upper Jurassic levels of the coastal region, with also some material from Lower and Upper Cretaceous strata from the central and northern sectors of the Lusitanian Basin. Here specimens attributed to Megalosaurus from different Portuguese institutions are reviewed and their taxonomic affinity and stratigraphic context are updated. Most specimens actually belong to different theropod groups, including several isolated teeth from different Upper Jurassic localities here assigned to Ceratosaurus, Torvosaurus, and Allosaurus, as well as an isolated tooth from the Lower Cretaceous that is attributed to an indeterminate allosauroid. Other theropod remains consist mostly of vertebral fragments of indeterminate avetheropods and allosauroids. Elements of other dinosaur groups are also represented, including a few vertebrae here referred to stegosaurians and iguanodontians, as well as a vertebra and some appendicular remains attributed to sauropods. Two vertebrae assigned to thalattosuchians were also identified. The study of this collection allows to better characterize the diversity of Late Jurassic dinosaur faunas from different areas of the Lusitanian Basin and provides some data on the poorly known Cretaceous fossil record of theropods from Portugal. [ABSTRACT FROM AUTHOR]

Thalattosuchia (Early Jurassic to Early Cretaceous) and Dyrosauridea (Late Cretaceous to Early Eocene) are crocodylomorph archosaurs which diversified in fluvial and marine environments and endured extinction events (i.e. Jurassic–Cretaceous boundary for Thalattosuchia; Cretaceous–Palaeogene for Dyrosauridea). Their postcrania remain globally undervalued in anatomical descriptions and diagnoses, shrouding the locomotive adaptations that possibly underpinned their radiations and longevity. We thoroughly surveyed the postcranial morphology of Dyrosauridea and Thalattosuchia, recreated their girdles in three‐dimensions using tens of high‐precisions 3D scans, and analysed their shape using geometric morphometrics. Dyrosauridea and Thalattosuchia have clearly distinct postcrania, even when found within similar environments, suggesting the existence of clade‐specific features limiting the strength of evolutionary convergence. Moreover, the range of postcranial morphologies evolved by dyrosaurids and thalattosuchians is large compared to extant crocodylians, making the latter unsatisfactory functional analogues for every group of extinct crocodylomorphs. Our work reveals the previously unsuspected potential of postcranial anatomy as an abundant source of phylogenetic and taxonomic characters to assess the relationships within Crocodylomorpha. Incorporation of postcranial anatomy therefore appears crucial to fully assess the ecology, disparity, and relationships of crocodylomorphs. [ABSTRACT FROM AUTHOR]

From the Middle Jurassic to the Early Cretaceous, metriorhynchid crocodylomorphs inhabited marine ecosystems across the European archipelago. Unfortunately, European metriorhynchids are only well known from Germany, France, and the UK, with the Eastern European fossil record being especially poor. This hinders our understanding of metriorhynchid biodiversity across these continuous seaways, and our ability to investigate provincialism. Here we describe eleven isolated tooth crowns and six vertebrae referable to Metriorhynchidae from the Callovian, Oxfordian, Volgian (Tithonian), and Ryazanian (Berriasian) or Valanginian of European Russia. We also describe an indeterminate thalattosuchian tooth from the lower Bajocian of the Volgograd Oblast, the first discovery of a marine reptile from the Bajocian strata of European Russia. These rare fossils, along with previous reports of Russian thalattosuchians, indicate that thalattosuchians have been common in the Middle Russian Sea since it was formed. Palaeolatitude calculations for worldwide metriorhynchid-bearing localities demonstrate that the occurrences in European Russia are the most northern, located mainly between 44-50 degrees north. However, metriorhynchids appear to be rare at these palaeolatitudes, and are absent from palaeolatitudes higher than 50?. These observations support the hypothesis that metriorhynchids evolved an elevated metabolism but were not endo-homeothermic, especially as endo-homeothermic marine reptiles (ichthyosaurs and plesiosaurs) remained abundant at much higher palaeolatitudes. [ABSTRACT FROM AUTHOR]

Metriorhynchids are marine crocodylomorphs found across Jurassic and Lower Cretaceous deposits of Europe and Central and South America. Despite being one of the oldest fossil families named in paleontology, the phylogenetic relationships within Metriorhynchidae have been subject to many revisions over the past 15 years. Herein, we describe a new metriorhynchid from the Kimmeridgian of Porrentruy, Switzerland. The material consists of a relatively complete, disarticulated skeleton preserving pieces of the skull, including the frontal, prefrontals, right postorbital, nasals, maxillae, right premaxillae and nearly the entire mandible, and many remains of the axial and appendicular skeleton such as cervical, dorsal, and caudal vertebrae, ribs, the left ischium, the right femur, and the right fibula. This new specimen is referred to the new species Torvoneustes jurensis sp. nov. as part of the large-bodied macrophagous tribe Geosaurini. Torvoneustes jurensis presents a unique combination of cranial and dental characters including a smooth cranium, a unique frontal shape, acute ziphodont teeth, an enamel ornamentation made of numerous apicobasal ridges shifting to small ridges forming an anastomosed pattern toward the apex of the crown and an enamel ornamentation touching the carina. The description of this new species allows to take a new look at the currently proposed evolutionary trends within the genus Torvoneustes and provides new information on the evolution of this clade. [ABSTRACT FROM AUTHOR]

Thalattosuchian crocodylomorphs were a diverse clade that lived from the Early Jurassic to the Early Cretaceous. The subclade Metriorhynchoidea underwent a remarkable transition, evolving from semi-aquatic ambush predators into fully aquatic forms living in the open oceans. Thalattosuchians share a peculiar palatal morphology with semi-aquatic and aquatic fossil cetaceans: paired anteroposteriorly aligned grooves along the palatal surface of the bony secondary palate. In extant cetaceans, these grooves are continuous with the greater palatine artery foramina, arteries that supply their oral thermoregulatory structures. Herein, we investigate the origins of thalattosuchian palatal grooves by examining CT scans of six thalattosuchian species (one teleosauroid, two early-diverging metriorhynchoids and three metriorhynchids), and CT scans of eleven extant crocodylian species. All thalattosuchians had paired osseous canals, enclosed by the palatines, that connect the nasal cavity to the oral cavity. These osseous canals open into the oral cavity via foramina at the posterior terminus of the palatal grooves. Extant crocodylians lack both the external grooves and the internal canals. We posit that in thalattosuchians these novel palatal canals transmitted hypertrophied medial nasal vessels (artery and vein), creating a novel heat exchange pathway connecting the palatal vascular plexus to the endocranial region. Given the general hypertrophy of thalattosuchian cephalic vasculature, and their increased blood flow and volume, thalattosuchians would have required a more extensive suite of thermoregulatory pathways to maintain stable temperatures for their neurosensory tissues. [ABSTRACT FROM AUTHOR]

From the Middle Jurassic to the Early Cretaceous, metriorhynchid crocodylomorphs inhabited marine ecosystems across the European archipelago. Unfortunately, European metriorhynchids are only well known from Germany, France, and the UK, with the Eastern European fossil record being especially poor. This hinders our understanding of metriorhynchid biodiversity across these continuous seaways, and our ability to investigate provincialism. Here we describe eleven isolated tooth crowns and six vertebrae referable to Metriorhynchidae from the Callovian, Oxfordian, Volgian (Tithonian), and Ryazanian (Berriasian) or Valanginian of European Russia. We also describe an indeterminate thalattosuchian tooth from the lower Bajocian of the Volgograd Oblast, the first discovery of a marine reptile from the Bajocian strata of European Russia. These rare fossils, along with previous reports of Russian thalattosuchians, indicate that thalattosuchians have been common in the Middle Russian Sea since it was formed. Palaeolatitude calculations for worldwide metriorhynchid-bearing localities demonstrate that the occurrences in European Russia are the most northern, located mainly between 44–50 degrees north. However, metriorhynchids appear to be rare at these palaeolatitudes, and are absent from palaeolatitudes higher than 50°. These observations support the hypothesis that metriorhynchids evolved an elevated metabolism but were not endo-homeothermic, especially as endo-homeothermic marine reptiles (ichthyosaurs and plesiosaurs) remained abundant at much higher palaeolatitudes.

Metriorhynchids are marine crocodylomorphs found across Jurassic and Lower Cretaceous deposits of Europe and Central and South America. Despite being one of the oldest fossil families named in paleontology, the phylogenetic relationships within Metriorhynchidae have been subject to many revisions over the past 15 years. Herein, we describe a new metriorhynchid from the Kimmeridgian of Porrentruy, Switzerland. The material consists of a relatively complete, disarticulated skeleton preserving pieces of the skull, including the frontal, prefrontals, right postorbital, nasals, maxillae, right premaxillae and nearly the entire mandible, and many remains of the axial and appendicular skeleton such as cervical, dorsal, and caudal vertebrae, ribs, the left ischium, the right femur, and the right fibula. This new specimen is referred to the new species Torvoneustes jurensis sp. nov. as part of the large-bodied macrophagous tribe Geosaurini. Torvoneustes jurensis presents a unique combination of cranial and dental characters including a smooth cranium, a unique frontal shape, acute ziphodont teeth, an enamel ornamentation made of numerous apicobasal ridges shifting to small ridges forming an anastomosed pattern toward the apex of the crown and an enamel ornamentation touching the carina. The description of this new species allows to take a new look at the currently proposed evolutionary trends within the genus Torvoneustes and provides new information on the evolution of this clade.

Thalattosuchian crocodylomorphs were a diverse clade that lived from the Early Jurassic to the Early Cretaceous. The subclade Metriorhynchoidea underwent a remarkable transition, evolving from semi-aquatic ambush predators into fully aquatic forms living in the open oceans. Thalattosuchians share a peculiar palatal morphology with semi-aquatic and aquatic fossil cetaceans: paired anteroposteriorly aligned grooves along the palatal surface of the bony secondary palate. In extant cetaceans, these grooves are continuous with the greater palatine artery foramina, arteries that supply their oral thermoregulatory structures. Herein, we investigate the origins of thalattosuchian palatal grooves by examining CT scans of six thalattosuchian species (one teleosauroid, two early-diverging metriorhynchoids and three metriorhynchids), and CT scans of eleven extant crocodylian species. All thalattosuchians had paired osseous canals, enclosed by the palatines, that connect the nasal cavity to the oral cavity. These osseous canals open into the oral cavity via foramina at the posterior terminus of the palatal grooves. Extant crocodylians lack both the external grooves and the internal canals. We posit that in thalattosuchians these novel palatal canals transmitted hypertrophied medial nasal vessels (artery and vein), creating a novel heat exchange pathway connecting the palatal vascular plexus to the endocranial region. Given the general hypertrophy of thalattosuchian cephalic vasculature, and their increased blood flow and volume, thalattosuchians would have required a more extensive suite of thermoregulatory pathways to maintain stable temperatures for their neurosensory tissues.

Secondarily marine tetrapod lineages have independently evolved osmoregulatory adaptations for life in salt water but inferring physiological changes in extinct marine tetrapods is difficult. The Mesozoic crocodylomorph clade Thalattosuchia is unique in having both direct evidence from natural endocasts and several proposed osteological correlates for salt exocrine glands. Here, we investigate salt gland evolution in thalattosuchians by creating endocranial reconstructions from CT scans of eight taxa (one basal thalattosuchian, one teleosauroid, two basal metriorhynchoids and four metriorhynchids) and four outgroups (three extant crocodylians and the basal crocodyliform Protosuchus) to identify salt gland osteological correlates. All metriorhynchoids show dorsolateral nasal cavity expansions corresponding to the location of nasal salt glands in natural casts, but smaller expansions in teleosauroids correspond more with the cartilaginous nasal capsule. The different sizes of these expansions suggest the following evolutionary sequence: (1) plesiomorphically small glands present in semi-aquatic teleosauroids draining through the nasal vestibule; (2) moderately sized glands in the basalmost metriorhynchoid Pelagosaurus ; and (3) hypertrophied glands in the clade comprising Eoneustes and metriorhynchids, with a pre-orbital fenestra providing a novel exit for salt drainage. The large gland size inferred from basal metriorhynchoids indicates advanced osmoregulation occurred while metriorhynchoids were semi-aquatic. This pattern does not precisely fit into current models of physiological evolution in marine tetrapods and suggests a unique sequence of changes as thalattosuchians transitioned from land to sea. [ABSTRACT FROM AUTHOR]

Environmental changes governed the diversity of marine ecosystems and the evolution of marine reptiles during the Jurassic. Abrupt climatic changes, mainly cooling, produced crises in marine ecosystems including marine reptiles, but global warming events at the Pliensbachian/Toarcian boundary and the early Toarcian Jenkyns Event led to a second order mass extinction. The Jenkyns Event coincides with exceptional preservation of marine reptiles in black shales, so widespread extinctions are masked to some extent by increases in specimens and diversity in the Lagerstätten. Marine reptile diversity responded to this biotic crisis; in the early Toarcian, stenopterygid ichthyosaurs and marine crocodylomorphs (thalattosuchians) diversified whereas some groups that survived the Jenkyns Event disappeared during the hyperthermal conditions of the middle and late Toarcian, including ichthyosaurs (leptonectids, temnodontosaurids, baso-parvipelvians), and sauropterygians (plesiosaurids, microcleidids). After the crisis, Aalenian climates were cold and the sea-level low with a poor record of marine reptiles. The diversity of marine ecosystems increased from the early Bajocian with a turnover in ichthyosaurs (Early Jurassic ichthyosaurs were replaced by ophthalmosaurids), sauropterygians diversified and increased in size (mainly pliosaurids and cryptocleidids), and pelagosaurid thalattosuchians disappeared at the same time that machimosaurids diversified and the Metriorhynchidae appeared. Marine reptiles reached a new maximum of diversity during the Callovian, but the Callovian/Oxfordian transition was a time of climatic cooling and sea-level fall that negatively impacted on marine ecosystems, including the extinction of rhomaleosaurids. From the middle Oxfordian, the development of large epeiric platforms and increase of temperature favoured the diversification of marine ecosystems, included marine reptiles. During the Late Jurassic, ophthalmosaurids, pliosaurids, cryptocleidids, and metriorhynchids dominated, and marine turtles diversified in coastal environments (eurysternids) and open carbonate platforms (plesiochelyids and thalassemydids). [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]

Crocodylomorphs have colonized various environments from fully terrestrial to fully aquatic, making it an important clade among archosaurs. A remarkable example of the rich past diversity of Crocodylomorpha Hay, 1930 is the marine colonization undergone by several crocodylomorph lineages, particularly Thalattosuchia Fraas, 1901 during the Early Jurassic-Early Cretaceous, and Dyrosauridae de Stefano, 1903 during the Late Cretaceous-Early Eocene. Thalattosuchia represents the most impressive and singular marine radiation among Crocodylomorpha, occupying various ecological niches, before enigmatically disappearing in the Cretaceous. Dyrosauridae, on the other hand, is known for surviving the end-Cretaceous mass extinction in abundance but subsequently vanished. The evolutionary path undertaken by crocodylomorphs into the aquatic environments and the reasons for their disappearance outside marine extinction events during the Mesozoic remains a mystery. Despite a well-preserved fossil record, attention has primarily centered on craniodental adaptations, overlooking the swimming-related adaptations recorded in the postcranial skeleton. This research primarily involves a comprehensive examination of the pectoral girdle of the most representative members of Thalattosuchia and Dyrosauridae, highlighting their evolutionary trajectories over time. Additionally, this work aims to test the phylogenetic signal residing in the postcranial anatomy of Crocodylomorpha. As such, the most recent and complete Crocodylomorpha phylogenetic dataset has been repurposed: 42 new postcranial characters have been added and several others have been revised to address our phylogenetic question. We stress that postcranial anatomy constitutes an important tool supply to better understand the relations of extinct crocodyliforms, but also offers insights on their development, ecology, and biomechanics., (© 2024 American Association for Anatomy.)

Abstract 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.

The genus Mystriosaurus, established by Kaup in 1834, was one of the first thalattosuchian genera to be named. The holotype, an incomplete skull from the lower Toarcian Posidonienschiefer Formation of Altdorf (Bavaria, southern Germany), is poorly known with a convoluted taxonomic history. For the past 60 years, Mystriosaurus has been considered a subjective junior synonym of Steneosaurus. However, our reassessment of the Mystriosaurus laurillardi holotype demonstrates that it is a distinct and valid taxon. Moreover, we find the holotype of “Steneosaurus” brevior, an almost complete skull from the lower Toarcian Whitby Mudstone Formation of Whitby (Yorkshire, UK), to be a subjective junior synonym of M. laurillardi. Mystriosaurus is diagnosed in having: a heavily and extensively ornamented skull; large and numerous neurovascular foramina on the premaxillae, maxillae and dentaries; anteriorly oriented external nares; and four teeth per premaxilla. Our phylogenetic analyses reveal M. laurillardi to be distantly related to Steneosaurus bollensis, supporting our contention that they are different taxa. Interestingly, our analyses hint that Mystriosaurus may be more closely related to the Chinese teleosauroid (previously known as Peipehsuchus) than any European form.

Dyrosauridae is a clade of neosuchian crocodyliforms that diversified in terrestrial and aquatic environments across the Cretaceous-Paleogene transition. The postcranial anatomy of dyrosaurids has long been overlooked, obscuring both their disparity and their locomotive adaptations. Here we thoroughly describe of the postcranial remains of an unusually small dyrosaurid, Cerrejonisuchus improcerus, from the middle-late Paleocene Cerrejón Formation of Colombia, and we provide a wealth of new data concerning the postcranial anatomy of the key dyrosaurids: Congosaurus bequaerti and Hyposaurus rogersii. We identify a series of postcranial autapomorphies in Cerrejonisuchus improcerus (an elliptic-shaped odontoid laterally wide, a ulna possessing a double concavity, a fibula bearing a widely flattened proximal end, a pubis showing a large non-triangular distal surface) as well as functionally-important traits such as a relatively long ulna (85% of the humerus’ length), short forelimb (83% of hindlimb’s length), or thoracic vertebra bearing comparatively large lateral process (with widened parapophysis and diapophysis) along with strongly arched thoracic ribs allowing a more sturdy and cylindrical rib cage. These indicate a more terrestrial lifestyle for Cerrejonisuchus compared to the derived members of the clade. We also built a dataset of 187 traits on 27 taxa, that extensively samples the cranial and postcranial architectures of exemplar crocodyliforms. We analyze these data in via Principal Coordinate Analysis (PCoA) to visualize the postcranial morphospace occupation of Dyrosauridae, Thalattosuchia, and Crocodylia. Our data reveal the existence of a distinctive postcranial anatomy for Dyrosauridae that is markedly distinct from that of crocodylians. As a result, modern crocodylians are probably not good functional analog for extinct crocodyliformes. Postcranial data should also be more widely used in phylogenetic and disparity analyses of Crocodyliformes.

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]

Dyrosauridae is a clade of neosuchian crocodyliforms that diversified in terrestrial and aquatic environments across the Cretaceous-Paleogene transition. The postcranial anatomy of dyrosaurids has long been overlooked, obscuring both their disparity and their locomotive adaptations. Here we thoroughly describe of the postcranial remains of an unusually small dyrosaurid, Cerrejonisuchus improcerus, from the middle-late Paleocene Cerrejón Formation of Colombia, and we provide a wealth of new data concerning the postcranial anatomy of the key dyrosaurids: Congosaurus bequaerti and Hyposaurus rogersii. We identify a series of postcranial autapomorphies in Cerrejonisuchus improcerus (an elliptic-shaped odontoid laterally wide, a ulna possessing a double concavity, a fibula bearing a widely flattened proximal end, a pubis showing a large non-triangular distal surface) as well as functionally-important traits such as a relatively long ulna (85% of the humerus' length), short forelimb (83% of hindlimb's length), or thoracic vertebra bearing comparatively large lateral process (with widened parapophysis and diapophysis) along with strongly arched thoracic ribs allowing a more sturdy and cylindrical rib cage. These indicate a more terrestrial lifestyle for Cerrejonisuchus compared to the derived members of the clade. We also built a dataset of 187 traits on 27 taxa, that extensively samples the cranial and postcranial architectures of exemplar crocodyliforms. We analyze these data in via Principal Coordinate Analysis (PCoA) to visualize the postcranial morphospace occupation of Dyrosauridae, Thalattosuchia, and Crocodylia. Our data reveal the existence of a distinctive postcranial anatomy for Dyrosauridae that is markedly distinct from that of crocodylians. As a result, modern crocodylians are probably not good functional analog for extinct crocodyliformes. Postcranial data should also be more widely used in phylogenetic and disparity analyses of Crocodyliformes. [ABSTRACT FROM AUTHOR]

Metriorhynchid crocodylomorphs were an important component in shallow marine ecosystems during the Middle Jurassic to Early Cretaceous in the European archipelago. While metriorhynchids are well known from western European countries, their central and eastern European record is poor and usually limited to isolated or fragmentary specimens which often hinders a precise taxonomic assignment. However, isolated elements such as tooth crowns, have been found to provide informative taxonomic identifications. Here we describe two isolated metriorhynchid tooth crowns from the upper Valanginian (Lower Cretaceous) of the Štramberk area, Czech Republic. Our assessment of the specimens, including multivariate analysis of dental measurements and surface enamel structures, indicates that the crowns belong to two distinct geosaurin taxa (Plesiosuchina? indet. and Torvoneustes? sp.) with different feeding adaptations. The specimens represent the first evidence of Metriorhynchidae from the Czech Republic and some of the youngest metriorhynchid specimens worldwide. [ABSTRACT FROM AUTHOR]

Teleosauroidea was a clade of ancient crocodylomorphs that were a key element of coastal marine environments during the Jurassic. Despite a 300-year research history and a recent renaissance in the study of their morphology and taxonomy, macroevolutionary studies of teleosauroids are currently limited by our poor understanding of their phylogenetic interrelationships. One major problem is the genus Steneosaurus, a wastebasket taxon recovered as paraphyletic or polyphyletic in phylogenetic analyses. We constructed a newly updated phylogenetic data matrix containing 153 taxa (27 teleosauroids, eight of which were newly added) and 502 characters, which we analysed under maximum parsimony using TNT 1.5 (weighted and unweighted analyses) and Bayesian inference using MrBayes v3.2.6 (standard, gamma and variation). The resulting topologies were then analysed to generate comprehensive higher-level phylogenetic hypotheses of teleosauroids and shed light on species-level interrelationships within the clade. The results from our parsimony and Bayesian analyses are largely consistent. Two large subclades within Teleosauroidea are recovered, and they are morphologically, ecologically and biogeographically distinct from one another. Based on comparative anatomical and phylogenetic results, we propose the following major taxonomic revisions to Teleosauroidea: (1) redefining Teleosauridae; (2) introducing one new family and three new subfamilies; (3) the resurrection of three historical genera; and (4) erecting seven new generic names and one new species name. The phylogeny infers that the Laurasian subclade was more phenotypically plastic overall than the Sub-Boreal-Gondwanan subclade. The proposed phylogeny shows that teleosauroids were more diverse than previously thought, in terms of morphology, ecology, dispersal and abundance, and that they represented some of the most successful crocodylomorphs during the Jurassic.

Teleosauroidea is a clade of ancient crocodylomorphs that were integral components of coastal marine environments throughout the Jurassic. For nearly two centuries, one of the most familiar genera of teleosauroids has been Steneosaurus , encompassing nearly every teleosauroid species at some point. However, no type species has been designated for Steneosaurus under ICZN Code rules; the type specimen of the presumed type species S. rostromajor Geoffroy Saint-Hilaire, 1825 (MNHN.RJN 134c-d) is a chimera that has been largely neglected in the literature. Moreover, there is confusion as to which teleosauroid species it pertains to, and the genus Steneosaurus is often recovered as paraphyletic or polyphyletic in phylogenetic analyses. As such, the validity of Steneosaurus is uncertain. Here we formally designate S. rostromajor as the type species of Steneosaurus , designate a lectotype and re-evaluate MNHN.RJN 134c-d. We compare it with several well-known teleosauroids, including Lemmysuchus and ' S.' edwardsi. Due to lack of autapomorphic characters, poor preservation and a tortured taxonomic history, we find MNHN.RJN 134c-d to be an undiagnostic and unreliable specimen. Thus, we consider S. rostromajor as a nomen dubium and propose that the genus Steneosaurus is undiagnostic. This has profound implications for teleosauroid phylogenetics, which we will clarify in an upcoming paper. [ABSTRACT FROM AUTHOR]

Teleosauroidea was a clade of successful, morphologically diverse, ancient crocodylomorphs that were integral in coastal marine/lagoonal environments during the Jurassic. Within Teleosauroidea, the macrophagous/durophagous tribe Machimosaurini evolved specialized feeding strategies (e.g. hypertrophied jaw musculature and blunt, heavily ornamented dentition) and large body sizes (> 7 m), becoming an important component of Middle and Late Jurassic ecosystems. These ocean-dwelling giants are well known from the Callovian (Lemmysuchus) of Europe and the UK, and from the Kimmeridgian–Tithonian (Machimosaurus) of Europe and northern Africa. There are reports of fragmentary machimosaurin material from the Bathonian of Africa, but the overall Bathonian teleosauroid material is poorly understood. While multiple specimens were described during the 19th and 20th centuries, little research has been done since. Here we re-describe two historically important Bathonian species from near Oxford, UK. We demonstrate that both ' Steneosaurus ' larteti and ' Steneosaurus ' boutilieri are valid taxa and we establish neotypes for both species and two new genera, Deslongchampsina and Yvridiosuchus. Our cladistic analysis finds Yvridiosuchus boutilieri as a basal member of Machimosaurini and Deslongchampsina larteti to be closely related to Steneosaurus heberti. Interestingly, four distinct teleosauroid ecomorphotypes are present in the Bathonian of Europe and teleosauroid ecomorphological diversity continued throughout the Callovian and Kimmeridgian/Tithonian in Europe and England. [ABSTRACT FROM AUTHOR]

Major evolutionary transitions, in which animals develop new body plans and adapt to dramatically new habitats and lifestyles, have punctuated the history of life. The origin of cetaceans from land-living mammals is among the most famous of these events. Much earlier, during the Mesozoic Era, many reptile groups also moved from land to water, but these transitions are more poorly understood. We use computed tomography to study changes in the inner ear vestibular system, involved in sensing balance and equilibrium, as one of these groups, extinct crocodile relatives called thalattosuchians, transitioned from terrestrial ancestors into pelagic (open ocean) swimmers. We find that the morphology of the vestibular system corresponds to habitat, with pelagic thalattosuchians exhibiting a more compact labyrinth with wider semicircular canal diameters and an enlarged vestibule, reminiscent of modified and miniaturized labyrinths of other marine reptiles and cetaceans. Pelagic thalattosuchians with modified inner ears were the culmination of an evolutionary trend with a long semiaquatic phase, and their pelagic vestibular systems appeared after the first changes to the postcranial skeleton that enhanced their ability to swim. This is strikingly different from cetaceans, which miniaturized their labyrinths soon after entering the water, without a prolonged semiaquatic stage. Thus, thalattosuchians and cetaceans became secondarily aquatic in different ways and at different paces, showing that there are different routes for the same type of transition. [ABSTRACT FROM AUTHOR]

Thalattosuchia, a clade of Mesozoic aquatic crocodyliforms, were the only archosaurs that ever became fully adapted to marine ecosystems. They are represented by two clades, the semiaquatic teleosauroids and the metriorhynchoids, which include fully pelagic forms. So far, little is known on the early evolutionary history of Metriorhynchoidea and data are sparse, especially from the early Middle Jurassic. Opisuchus meieri gen. et sp. nov. a metriorhynchoid crocodyliform from the early Aalenian (early Middle Jurassic) of southern Germany, is described here. It is one of the most complete specimens of a non-metriorhynchid metriorhynchoid, and the best-preserved thalattosuchian described from the Aalenian. The new taxon is represented by a nearly complete skull, which has a unique combination of characters distinguishing it from other species of Metriorhynchoidea. It displays a mosaic of plesiomorphic and apomorphic morphological features that sheds new light on early metriorhynchoid evolution. This taxon is an important puzzle piece, which will help to better track the mosaic character distribution in Thalattosuchia. [ABSTRACT FROM AUTHOR]

During the Mesozoic, Crocodylomorpha had a much higher taxonomic and morphological diversity than today. Members of one particularly successful clade, Thalattosuchia, are well‐known for being longirostrine: having long, slender snouts. It has generally been assumed that Thalattosuchia owed their success in part to the evolution of longirostry, leading to a feeding ecology similar to that of the living Indian gharial, Gavialis. Here, we compare form and function of the skulls of the thalattosuchian Pelagosaurus and Gavialis using digital reconstructions of the skull musculoskeletal anatomy and finite element models to show that they had different jaw muscle arrangements and biomechanical behaviour. Additionally, the relevance of feeding‐related mandibular traits linked to longirostry in the radiation of crocodylomorph clades was investigated by conducting an evolutionary rates analysis under the variable rates model. We find that, even though Pelagosaurus and Gavialis share similar patterns of stress distribution in their skulls, the former had lower mechanical resistance. This suggests that compared to Gavialis, Pelagosaurus was unable to process large, mechanically less tractable prey, instead operating as a specialized piscivore that fed on softer and smaller prey. Secondly, innovation of feeding strategies was achieved by rate acceleration of functional characters of the mandible, a key mechanism for the diversification of certain clades like thalattosuchians and eusuchians. Different rates of functional evolution suggest divergent diversification dynamics between teleosaurids and metriorhynchids in the Jurassic. [ABSTRACT FROM AUTHOR]

The geographic origins and distributional patterns of Gondwanan teleosauroids during the Jurassic have been fiercely debated over many years. Unlike the rich thalattosuchian fossil record from Laurasia, teleosauroids described from Gondwanan ecosystems are relatively scarce. Most of the known occurrences consist of isolated and fragmentary bones collected in Madagascar, Morocco, Tunisia, India, and Ethiopia. Nevertheless, these specimens, although fragmentary, have provided substantial information for assessing the evolutionary scenarios of multiple teleosauroid lineages and have shown that certain teleosauroid taxa were widespread rather than endemic to Western Europe. Here, a partial skeleton of a teleosauroid crocodylomorph is described. It was found in the late Middle Jurassic (Callovian) deposits of southeastern Tunisia by a team of French and Tunisian paleontologists; however, it has not been thoroughly studied at both macro- and microscopic scale until now. The new specimen is composed of an incomplete symphyseal portion of a lower jaw in addition to isolated teeth, osteoderms (both dorsal and ventral), thoracic and caudal vertebrae, and several thoracic ribs. The specimen has several morphological characters that are reminiscent of longirostrine teleosauroids. Due to the total absence of other cranial bones, as well as the pectoral and pelvic girdles, the specimen is not diagnostic to the generic level. However, these new remains represent the youngest ascertained occurrence of a definitive non-machimosaurin teleosauroid in Africa, provide additional insights into the geographic distribution of Thalattosuchia, and raise once again the question whether the origins of this clade were Gondwanan or Laurasian. [ABSTRACT FROM AUTHOR]

The genus Mystriosaurus, established by Kaup in 1834, was one of the first thalattosuchian genera to be named. The holotype, an incomplete skull from the lower Toarcian Posidonienschiefer Formation of Altdorf (Bavaria, southern Germany), is poorly known with a convoluted taxonomic history. For the past 60 years, Mystriosaurus has been considered a subjective junior synonym of Steneosaurus. However, our reassessment of the Mystriosaurus laurillardi holotype demonstrates that it is a distinct and valid taxon. Moreover, we find the holotype of "Steneosaurus" brevior, an almost complete skull from the lower Toarcian Whitby Mudstone Formation of Whitby (Yorkshire, UK), to be a subjective junior synonym of M. laurillardi. Mystriosaurus is diagnosed in having: a heavily and extensively ornamented skull; large and numerous neurovascular foramina on the premaxillae, maxillae and dentaries; anteriorly oriented external nares; and four teeth per premaxilla. Our phylogenetic analyses reveal M. laurillardi to be distantly related to Steneosaurus bollensis, supporting our contention that they are different taxa. Interestingly, our analyses hint that Mystriosaurus may be more closely related to the Chinese teleosauroid (previously known as Peipehsuchus) than any European form. [ABSTRACT FROM AUTHOR]

Neptunidraco ammoniticus is a thalattosuchian crocodylomorph from the Rosso Ammonitico Veronese Formation (RAVF, Middle Jurassic) of northern Italy. Erected from one partial specimen, Neptunidraco is pivotal in reconstructing thalattosuchian evolution, being it the oldest known member of Metriorhynchidae. Two additional RAVF thalattosuchians have been referred to Neptunidraco. A revised diagnosis of N. ammoniticus is provided here. Using a well-sampled phylogenetic data set of Crocodylomorpha, the affinities of all three RAVF thalattosuchian specimens are investigated simultaneously for the first time using parsimony tree-search strategies and Bayesian inference using the Fossilized Birth-Death with Sampled Ancestor (FBDSA) model. The results of the alternative analyses are not consistent in the placement of the RAVF specimens. The holotype of N. ammoniticus is consequently referred to Metriorhynchidae incertae sedis. The first referred specimen is recovered in various alternative placements among Metriorhynchoidea. The third and most fragmentary specimen is recovered as a crocodylomorph of uncertain affinities in the parsimony analysis and in the undated Bayesian analysis, and a metriorhynchoid sister taxon of the second RAVF specimen in the tip-dated Bayesian analysis. Only a subset of the results in the parsimony-based analyses supports the referral of the latter two specimens to Neptunidraco. The unusually high rate of morphological divergence for the Neptunidraco branch, inferred in previous iterations of the Bayesian inference analyses but not recovered in the novel analysis, was likely an artifact of the a priori constraint of all RAVF thalattosuchians into a single taxonomic unit, and of the arbitrarily fixed tip-age priors for the terminal taxa. These results confirm the utility of specimen-level morphological analysis and of combined tree-search strategies for inferring the affinities and the inclusiveness of fragmentary but significant fossil taxa, and reinforce the importance of incorporating stratigraphic uncertainty as prior in tip-dated Bayesian inference analyses. [ABSTRACT FROM AUTHOR]

Thalattosuchians are a group of Mesozoic crocodylomorphs known from aquatic deposits of the Early Jurassic–Early Cretaceous that comprises two main lineages of almost exclusively marine forms, Teleosauridae and Metriorhynchoidea. Teleosaurids were found in shallow marine, brackish and freshwater deposits, and have been characterized as semiaquatic near-shore forms, whereas metriorhynchids are a lineage of fully pelagic forms, supported by a large set of morphological characters of the skull and postcranial anatomy. Recent contributions on Thalattosuchia have been focused on the study of the endocranial anatomy. This newly available information provides novel evidence to suggest adaptations on the neuroanatomy, senses organs, vasculature, and behavioral evolution of these crocodylomorphs. However, is still not clear if the major morphological differences between teleosaurids and metriorhynchids were also mirrored by changes in the braincase and endocranial anatomy. Based on X-ray CT scanning and digital endocast reconstructions we describe the braincase and endocranial anatomy of two well-preserved specimens of Thalattosuchia, the semiaquatic teleosaurid Steneosaurus bollensis and the pelagic metriorhynchid Cricosaurus araucanensis. We propose that some morphological traits, such as: an enlarged foramen for the internal carotid artery, a carotid foramen ventral to the occipital condyle, a single CN XII foramen, absence of brain flexures, well-developed cephalic vascular system, lack of subtympanic foramina and the reduction of the paratympanic sinus system, are distinctive features of Thalattosuchia. It has been previously suggested that the enlarged foramen for the internal carotid artery, the absence of brain flexures, and the hypertrophied cephalic vascular system were synapomorphies of Metriorhynchidae; however, new information revealed that all of these features were already established at the base of Thalattosuchia and might have been exapted later on their evolutionary history. Also, we recognized some differences within Thalattosuchia that previously have not been received attention or even were overlooked (e.g., circular/bilobate trigeminal foramen, single/double CN XII foramen, separation of the cranioquadrate canal from the external otic aperture, absence/presence of lateral pharyngeal foramen). The functional significances of these traits are still unclear. Extending the sampling to other Thalattosuchia will help to test the timing of acquisition and distribution of these morphological modifications among the whole lineage. Also comparison with extant marine tetrapods (including physiological information) will be crucial to understand if some (and/or which) of the morphological peculiarities of thalattosuchian braincases are products of directional natural selection resulting in a fully adaptation to a nektonic life style.

En 1808 puis en 1824, Georges Cuvier signala dans la région du Mans la présence de restes de crocodiles fossiles à partir desquels fut créée l’espèce Crocodilus maunyi GRAY, 1831. Il avait été informé de ces découvertes par le naturaliste manceau Louis Jean Charles Maulny (1758-1815) et par Jean Antoine Daudin (1749-1832), premier conservateur du Muséum d’histoire naturelle du Mans. Faute de figures publiées, ce matériel est resté énigmatique durant deux siècles. Des aquarelles inédites conservées dans les archives de Georges Cuvier permettent de dévoiler les spécimens qu’il avait décrits sous le nom de « crocodiles fossiles des environs du Mans ». Ils comprennent des dents, des vertèbres et des portions de mâchoires de crocodylomorphes marins et de plésiosauriens découvertes dans le Mésozoïque de la Sarthe (Bathonien, Callovien, Cénomanien ?). Un de ces dessins présente une dent de pliosaure du Bathonien de Ber- nay-en-Champagne qui serait la plus ancienne découverte d’un représentant de ce groupe. Deux autres aquarelles montrent une portion de mandibule d’un crocodylomorphe marin (Thalattosuchia, Metriorhynchidae indéterminé) et une vertèbre de plésiosaurien (Plesiosauria indéterminé) du Callovien moyen de Chaufour-Notre-Dame, toujours conservées au Musée Vert, le muséum d’histoire naturelle du Mans. Ces deux spécimens furent découverts en 1816 par un amateur local, le docteur Eusèbe Marie Tendron (1778-1854). Crocodilus maunyi GRAY, 1831 reste néanmoins considéré comme un nomen dubium au regard du manque de caractères diagnostiques de l’ensemble de ses syntypes.

Thalattosuchians are a group of Mesozoic crocodylomorphs known from aquatic deposits of the Early Jurassic-Early Cretaceous that comprises two main lineages of almost exclusively marine forms, Teleosauridae and Metriorhynchoidea. Teleosaurids were found in shallowmarine, brackish and freshwater deposits, and have been characterized as semiaquatic near-shore forms, whereas metriorhynchids are a lineage of fully pelagic forms, supported by a large set of morphological characters of the skull and postcranial anatomy. Recent contributions on Thalattosuchia have been focused on the study of the endocranial anatomy. This newly available information provides novel evidence to suggest adaptations on the neuroanatomy, senses organs, vasculature, and behavioral evolution of these crocodylomorphs. However, is still not clear if the major morphological differences between teleosaurids and metriorhynchids were also mirrored by changes in the braincase and endocranial anatomy. Based on X-ray CT scanning and digital endocast reconstructions we describe the braincase and endocranial anatomy of two well-preserved specimens of Thalattosuchia, the semiaquatic teleosaurid Steneosaurus bollensis and the pelagic metriorhynchid Cricosaurus araucanensis. We propose that somemorphological traits, such as: an enlarged foramen for the internal carotid artery, a carotid foramen ventral to the occipital condyle, a single CN XII foramen, absence of brain flexures, welldeveloped cephalic vascular system, lack of subtympanic foramina and the reduction of the paratympanic sinus system, are distinctive features of Thalattosuchia. It has been previously suggested that the enlarged foramen for the internal carotid artery, the absence of brain flexures, and the hypertrophied cephalic vascular system were synapomorphies of Metriorhynchidae; however, new information revealed that all of these features were already established at the base of Thalattosuchia and might have been exapted later on their evolutionary history. Also, we recognized some differences within Thalattosuchia that previously have not been received attention or even were overlooked (e.g., circular/bilobate trigeminal foramen, single/double CN XII foramen, separation of the cranioquadrate canal from the external otic aperture, absence/presence of lateral pharyngeal foramen). The functional significances of these traits are still unclear. Extending the sampling to other Thalattosuchia will help to test the timing of acquisition and distribution of these morphological modifications among the whole lineage. Also comparison with extant marine tetrapods (including physiological information) will be crucial to understand if some (and/or which) of the morphological peculiarities of thalattosuchian braincases are products of directional natural selection resulting in a fully adaptation to a nektonic life style. [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]

Thalattosuchians were highly specialised aquatic archosaurs of the Jurassic and Early Cretaceous, and represent a peak of aquatic adaptation among crocodylomorphs. Relatively little is known of their endocranial anatomy or its relevance for the evolution of sensory systems, physiology, and other aspects of biology. Nevertheless, such data have significance for two reasons: (1) thalattosuchians represent an important data point regarding adaptation to marine life in tetrapods; and (2) as early-diverging members of the crocodylian stem-lineage, thalattosuchians provide information on the evolutionary assembly of the brain and other endocranial structures in crocodylomorphs. Here we use µCT data to virtually reconstruct the endocranial anatomy of Pelagosaurus typus, an early thalattosuchian with plesiomorphic traits of relevance to the split between the two major subgroups: Teleosauroidea and Metriorhynchoidea. Interpretation of these data in a broad comparative context indicate that several key endocranial features may be unique to thalattosuchians, including: a pyramidal morphology of the semicircular canals, the presence of an elongate endosseous cochlear duct that may indicate enhanced hearing ability, the presence of large, paired canals extending anteriorly from an enlarged pituitary fossa, a relatively straight brain (possibly due to the presence of large, laterally placed orbits), and an enlarged venous sinus projecting dorsally from the endocast that is confluent with the paratympanic sinus system. Notably, we document a large expansion of the nasal cavity anterior to the orbits in Pelagosaurus as an osteological correlate of an enlarged salt gland previously only documented in Late Jurassic metriorhynchoids. This is the first anatomical evidence of this structure in early thalattosuchians. Pelagosaurus also shares the presence of paired olfactory bulbs with metriorhynchoids, and shows an enlarged cerebrum, which may also be present in teleosauroids. Taken together, our findings indicate that physiological and sensory adaptations to marine life occurred early in thalattosuchian evolution, predating the origins of flippers, tail flukes, and hydrodynamic body forms seen later in metriorhynchoids.

Morphological and physiological features indicate Metriorhynchidae as the only group of crocodylomorphs with a pelagic lifestyle. Some of these features have evolved convergently in several clades of tetrapods secondarily adapted to aquatic life. One striking feature of metrio-rhynchids as compared to other crocodylomorphs is the morphology of the pelvic region (i.e., jWiflW ventrally deflected sacral ribs and reduced pelvic girdle), which increases significantly the depth of this region. This morphology, as a whole, resembles that of other viviparous Mesozoic marine reptiles not phylogenetically related to metriorhynchids. We tested two alternative hypotheses of reproductive strategies in this clade: oviparity ra. viviparity. Given the lack of direct evidence supporting one or the other, we explored the use of evidence that may disconfirm either of these hypotheses. Using this counter-inductive approach, we found no cases contradicting viviparity in metriorhynchids, except for their phylogenetic position as archosaurs. A survey of reproductive modes amongst amniotes depicts the evolutionär}' plasticity of the transition to viviparity, and a widespread occurrence among tetrapods secondarily adapted to a marine life. Assuming oviparity for metriorhynchids implies egg-laying out of the water. However, their postcranial morphology (i.e., features of fore and hind limbs, pelvic girdle, and tail) contradicts this possibility. In this context, we rejected oviparity for metriorhynchids. [ABSTRACT FROM AUTHOR]

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.

Marine crocodylomorphs were particularly abundant in Europe during the Middle Jurassic, but were very scarce in Africa. New finds of thalattosuchian cranial remains in Morocco suggest that this scarcity is probably related to poor sampling rather than original diversity. These remains pertain to the coastal thalattosuchians, the teleosauroids, and particularly to the clade grouping the blunt-toothed ' Steneosaurus' obtusidens and the genus Machimosaurus. A new tribe is erected grouping these two taxa: Machimosaurini. Until now the machimosaurins have been known from the middle Callovian. The new material extends the presence of this group further back to the lower Bathonian, nearly 5 myr earlier. The machimosaurins are the only teleosauroid group that has been recently reviewed, and the difference between the revised diversity provided herein and that previously reported is large. A review of other teleosauroids and clear establishment of their phylogenetic relationships are also likely to have considerable impact on their observed diversity. So, until a complete review of the teleosauroids is carried out, the results of crocodylomorph diversity analyses should be treated with caution. [ABSTRACT FROM AUTHOR]

A new teleosaurid from the Lower Cretaceous of Tataouine (Tunisia), Machimosaurus rex sp. nov., definitively falsifies that these crocodylomorphs faced extinction at the end of the Jurassic. Phylogenetic analysis supports its placement closer to M. hugii and M. mosae than M. buffetauti . With the skull length up to 160 cm and an estimated body length of 10 m, M. rex results the largest known thalattosuchian, and the largest known crocodylomorph at its time. This giant thallatosuchian probably was an ambush predator in the lagoonal environments that characterized the Tethyan margin of Africa during the earliest Cretaceous. Whether the Jurassic-Cretaceous mass extinction was real or artefact is debated. The discovery of M . rex supports that the end-Jurassic crisis affected primarily Laurasian biota and its purported magnitude is most likely biased by the incomplete Gondwanan fossil record. The faunal turnovers during the J-K transition are likely interpreted as local extinction events, triggered by regional ecological factors, and survival of widely-distributed and eurytypic forms by means of habitat tracking. [ABSTRACT FROM AUTHOR]

Résumé C’est à partir de la collection d’un certain « abbé Bachelet [sic] , naturaliste à Rouen » que Georges Cuvier donna en 1808 les premières descriptions scientifiques de dinosaures et de crocodiliens marins (Thalattosuchia). Selon Cuvier, « Bachelet » n’avait rien publié sur les circonstances de la découverte de ces fossiles et un doute subsistait jusqu’à présent sur leur origine stratigraphique. Cet article dévoile pour la première fois l’identité de l’abbé Charles Bacheley (1716–1795) et présente une biographie de ce pionnier de la paléontologie normande. Il est l’auteur d’une notice publiée en 1778 sur les pétrifications des côtes du pays d’Auge entre les Vaches Noires et Trouville-sur-Mer. Ce travail passé totalement inaperçu révèle que les spécimens de la collection Bacheley étudiés par Cuvier proviennent des Marnes de Dives (Callovien supérieur) ou de Villers (Oxfordien inférieur) visibles au pied des falaises des Vaches Noires. Bacheley attribue ces restes de dinosaures et de crocodiliens marins à des « poissons », terme qui servait encore à cette époque à désigner, selon l’usage populaire, les cétacés. Bacheley n’excluait pas la possibilité que ces ossements pétrifiés pussent appartenir à des formes animales sans analogues vivants. Georges Cuvier published in 1808 the first scientific descriptions of dinosaur and marine crocodilian (Thalattosuchia) remains from the collection of a certain “ Abbé Bachelet [sic]”, a naturalist in Rouen (Normandy, France). According to Cuvier, “Bachelet” never published any papers about the circumstances of the discovery of these fossils. Since then, there has been some doubt about their precise geographic and stratigraphic origin. This article reveals for the first time the identity of Charles Bacheley (1716–1795) and presents the biography of this pioneer in Norman palaeontology. He is the author of a note published in 1778 on the petrifactions found on the coast of Pays d’Auge (Calvados) between the Vaches Noires and Trouville-sur-Mer. This study has gone completely unnoticed and reveals that the specimens from the Bacheley collection studied by Cuvier come from the “Marnes de Dives” (Upper Callovian) or the “Marnes de Villers” (Lower Oxfordian) of the Vaches Noires cliffs. Bacheley identified these dinosaurs and marine crocodilian remains as belonging to “fishes,” which at that time also included cetaceans according to popular usage. Interestingly, Bacheley did not exclude the possibility that these petrified bones could belong to animals distinct from living forms. [ABSTRACT FROM AUTHOR]