Search

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]

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]

The purpose of this application, under Article 55.3 of the ICZN Code, is to remove the homonymy between the unused Cricosaurinae Koken, 1888 (Archosauria, Crocodylomorpha, Metriorhynchidae) (type genus Cricosaurus Wagner, 1858) and the in-use senior homonym Cricosaurinae Cope, 1875 (Lepidosauria, Squamata, Xantusiidae) (type genus Cricosaura Gundlach & Peters in Peters, 1863). We propose to emend the stem of the name of the type genus of the unused Cricosaurinae Koken, 1888 from Cricosaur- to Cricosaurus-, which would result in Cricosaurusinae Koken, 1888. Reversal of precedence (Article 23.9 of the Code) is invoked for the subjective synonymy between Cricosaurini Koken, 1888, herewith treated as a nomen oblitum, and Rhacheosaurini Young, Bell & Brusatte, 2011, herewith treated as a nomen protectum. [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]

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]

Marine reptiles from the Upper Jurassic of Central Europe are rare and often fragmentary, which hinders their precise taxonomic identification and their placement in a palaeobiogeographic context. Recent fieldwork in the Kimmeridgian of Krzyżanowice, Poland, a locality known from turtle remains originally discovered in the 1960s, has reportedly provided additional fossils thought to indicate the presence of a more diverse marine reptile assemblage, including giant pliosaurids, plesiosauroids, and thalattosuchians. Based on its taxonomic composition, the marine tetrapod fauna from Krzyżanowice was argued to represent part of the “Matyja-Wierzbowski Line”—a newly proposed palaeobiogeographic belt comprising faunal components transitional between those of the Boreal and Mediterranean marine provinces. Here, we provide a detailed re-description of the marine reptile material from Krzyżanowice and reassess its taxonomy. The turtle remains are proposed to represent a “plesiochelyid” thalassochelydian (Craspedochelys? sp.) and the plesiosauroid vertebral centrum likely belongs to a cryptoclidid. However, qualitative assessment and quantitative analysis of the jaws originally referred to the colossal pliosaurid Pliosaurus clearly demonstrate a metriorhynchid thalattosuchian affinity. Furthermore, these metriorhynchid jaws were likely found at a different, currently indeterminate, locality. A tooth crown previously identified as belonging to the thalattosuchian Machimosaurus is here considered to represent an indeterminate vertebrate. The revised taxonomy of the marine reptiles from Krzyżanowice, as well as the uncertain provenance of the metriorhynchid specimen reported from the locality, cast doubt on the palaeobiogeographic significance of the assemblage.

Here we describe a new species of the metriorhynchid thalattosuchian Cricosaurus, C. bambergensis sp. nov., from the Upper Jurassic Torleite Formation of Wattendorf near Bamberg, Bavaria (southern Germany). The holotype and only known specimen is a nearly complete skeleton that shows a number of diagnostic traits including a bicarinate dentition formed by labiolingually compressed tooth crowns that lack a conspicuous enamel ornamentation and the presence of a distinct midline ridge with paired depressions on the palatines. Our phylogenetic analysis recovers a grouping of Cricosaurus bambergensis sp. nov. with C. elegans and C. suevicus. The implications of the new Cricosaurus species to the species complex from the late Kimmeridgian–early Tithonian of southern Germany is discussed. Our description of C. bambergensis demonstrates that the specific, and morphological, diversity of Cricosaurus in southern Germany was higher than previously thought. This coincides with the recent trend of re-evaluating the species-complexes of extant taxa, and the identification of new “cryptic species”. As such, the crocodylomorph fossil record will need to be re- examined to ensure there is not an underestimation of their biodiversity.

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]

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]

Cricosaurus represents one of the most important radiations within Metriorhynchidae, a specialized marine clade of Crocodylomorpha that flourished in marine environments from the Middle Jurassic until the Early Cretaceous. Here, we describe a new species of Cricosaurus, Cricosaurus rauhuti sp. nov., from the Mörnsheim Formation (lower Tithonian) of southern Germany. The specimen is the first three-dimensionally preserved Cricosaurus from this realm, and it is represented by a partially preserved cranium and lower jaw that has autapomorphic characters that distinguish it from any other species of Cricosaurus. The phylogenetic analysis shows that the new species is more closely related to some non-European forms from the Late Jurassic and one taxon from the Valanginian of Germany (all 3D-preserved) than to other Late Jurassic Cricosaurus from southern Germany (all 2D-preserved). Our exploratory analyses confirm that this is not a preservation bias. For the first time, this shows that there are two clades of Cricosaurus present during the Late Jurassic in Germany. Furthermore, it provides additional evidence of a close faunal relationship between Europe and South America in the Late Jurassic, as also evidenced by the fossil record of other marine reptiles. Finally, the distribution of metriorhynchids in localities of the Solnhofen Archipelago suggests a change in the taxic diversity of the region during the early Tithonian, which might be the result of environmental changes in the Solnhofen Archipelago. [ABSTRACT FROM AUTHOR]

Marine reptiles from the Upper Jurassic of Central Europe are rare and often fragmentary, which hinders their precise taxonomic identification and their placement in a palaeobiogeographic context. Recent fieldwork in the Kimmeridgian of Krzyżanowice, Poland, a locality known from turtle remains originally discovered in the 1960s, has reportedly provided additional fossils thought to indicate the presence of a more diverse marine reptile assemblage, including giant pliosaurids, plesiosauroids, and thalattosuchians. Based on its taxonomic composition, the marine tetrapod fauna from Krzyżanowice was argued to represent part of the "Matyja-Wierzbowski Line"--a newly proposed palaeobiogeographic belt comprising faunal components transitional between those of the Boreal and Mediterranean marine provinces. Here, we provide a detailed re-description of the marine reptile material from Krzyżanowice and reassess its taxonomy. The turtle remains are proposed to represent a "plesiochelyid" thalassochelydian (Craspedochelys? sp.) and the plesiosauroid vertebral centrum likely belongs to a cryptoclidid. However, qualitative assessment and quantitative analysis of the jaws originally referred to the colossal pliosaurid Pliosaurus clearly demonstrate a metriorhynchid thalattosuchian affinity. Furthermore, these metriorhynchid jaws were likely found at a different, currently indeterminate, locality. A tooth crown previously identified as belonging to the thalattosuchian Machimosaurus is here considered to represent an indeterminate vertebrate. The revised taxonomy of the marine reptiles from Krzyżanowice, as well as the uncertain provenance of the metriorhynchid specimen reported from the locality, cast doubt on the palaeobiogeographic significance of the assemblage. [ABSTRACT FROM AUTHOR]

Teleosauridae and Metriorhynchidae were thalattosuchian crocodylomorph clades that secondarily adapted to marine life and coexisted during the Middle to Late Jurassic. While teleosaurid diversity collapsed at the end of the Jurassic, most likely as a result of a global cooling of the oceans and associated marine regressions, metriorhynchid diversity was largely unaffected, although the fossil record of Thalattosuchia is poor in the Cretaceous. In order to investigate the possible differences in thermophysiologies between these two thalattosuchian lineages, we analysed stable oxygen isotope compositions (expressed as δ18O values) of tooth apatite from metriorhynchid and teleosaurid specimens. We then compared them with the δ18O values of coexisting endo-homeothermic ichthyosaurs and plesiosaurs, as well as ecto-poikilothermic chondrichthyans and osteichthyans. The distribution of δ18O values suggests that both teleosaurids and metriorhynchids had body temperatures intermediate between those of typical ecto-poikilothermic vertebrates and warm-blooded ichthyosaurs and plesiosaurs, metriorhynchids being slightly warmer than teleosaurids. We propose that metriorhynchids were able to raise their body temperature above that of the ambient environment by metabolic heat production, as endotherms do, but could not maintain a constant body temperature compared with fully homeothermic ichthyosaurs and plesiosaurs. Teleosaurids, on the other hand, may have raised their body temperature by mouth-gape basking, as modern crocodylians do, and benefited from the thermal inertia of their large body mass to maintain their body temperature above the ambient one. Endothermy in metriorhynchids might have been a by-product of their ecological adaptations to active pelagic hunting, and it probably allowed them to survive the global cooling of the Late Jurassic, thus explaining the selective extinction affecting Thalattosuchia at the Jurassic-Cretaceous boundary. [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.

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]

Here we describe a new species of the metriorhynchid thalattosuchian Cricosaurus, C. bambergensis sp. nov., from the Upper Jurassic Torleite Formation of Wattendorf near Bamberg, Bavaria (southern Germany). The holotype and only known specimen is a nearly complete skeleton that shows a number of diagnostic traits including a bicarinate dentition formed by labiolingually compressed tooth crowns that lack a conspicuous enamel ornamentation and the presence of a distinct midline ridge with paired depressions on the palatines. Our phylogenetic analysis recovers a grouping of Cricosaurus bambergensis sp. nov. with C. elegans and C. suevicus. The implications of the new Cricosaurus species to the species complex from the late Kimmeridgian-early Tithonian of southern Germany is discussed. Our description of C. bambergensis demonstrates that the specific, and morphological, diversity of Cricosaurus in southern Germany was higher than previously thought. This coincides with the recent trend of re-evaluating the species-complexes of extant taxa, and the identification of new "cryptic species". As such, the crocodylomorph fossil record will need to be re-examined to ensure there is not an underestimation of their biodiversity. [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]

The fossil record of metriorhynchids and plesiosaurians from the Rosso Ammonitico Veronese Formation (RAVFm, Middle–Upper Jurassic, Italy) is represented by elements collected between the eighteenth and twentieth centuries. All the metriorhynchid material is referred to the genusNeptunidraco. The first RAVFm plesiosaurian material was collected in the nineteenth century and referred toPlesiosaurus: elements are here described and interpreted as a chimerical association of crocodylomorph and plesiosaurian bones, providing the first co-occurrence of these clades in the RAVFm. The second plesiosaurian is the associated skeleton that we refer toAnguanax zignoigen. et sp. nov. Bayesian phylogenetic analysis confirms the basal geosaurine affinities ofNeptunidracoresulted by parsimony analysis. Using both methods,Anguanaxwas recovered as a basal pliosaurid, sister group of the clade includingMarmornectesand Thalassophonea. Bayesian inference methods indicate that both Italian lineages diverged from other known lineages between 176 and 171 Mya, also showing divergence rates significantly higher than any other representative of their respective clades. We suggest a phase of rapid evolutionary adaptation to deeper marine environments in the ancestors of the Rosso Ammonitico Veronese reptiles as a response to the latest Liassic regressive regime in Northern Tethys. [ABSTRACT FROM PUBLISHER]

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]

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

We report the largest known British specimen of the metriorhynchid crocodylomorph genusDakosaurusdiscovered offshore from Chesil Beach, Dorset, England (Kimmeridge Clay Formation). This specimen is large toothed, which has extreme enamel spalling on the labial surface, enlarged carinae (‘carinal flanges’), carinal wear, macroziphodont denticles, and the crown retains much of its labiolingual width along most of its apicobasal length. This suite of morphologies is unique toDakosaurus. All known Kimmeridge Clay FormationDakosaurusspecimens are isolated tooth crowns. A skull previously referred toDakosauruslacks all the cranial apomorphies ofD. maximusandD. andiniensis, and cannot be referred to this genus. Furthermore, the vast majority of putativeDakosaurustooth crowns from the ‘Potton Sands’ ( = Woburn Sands Formation) do indeed representDakosaurus(as well as twoPlesiosuchusspecimens), and they most likely originate from the Kimmeridge Clay Formation. [ABSTRACT FROM AUTHOR]

We report the oldest known specimen of the metriorhynchid crocodylomorph genusGeosaurus, and the first specimen discovered from the Lower Kimmeridge Clay Formation of England (Kimmeridgian stage,Rasenia cymodoceSub-Boreal ammonite Zone). This specimen, an isolated tooth, has the tri-facetted labial surface morphology that is unique to Late Jurassic species ofGeosaurus. Its dimensions suggest that it is one of the posterior teeth. Macrowear on the labial surface (enamel spalling) shows evidence of tooth–food abrasion. The presence ofGeosaurusnear the base of the lower Kimmeridgian, along with the recently re-described middle Oxfordian specimen attributable toTorvoneustes, shows that the subclade Geosaurini evolved and diversified prior to the Kimmeridgian. Members Geosaurini evolved a diverse array of sophisticated feeding mechanisms, including durophagous forms (Torvoneustes), barracuda-like forms with ‘scissor-like’ occlusion (Geosaurus), species reminiscent of false killer whales/Type 1 North Atlantic killer whales (Dakosaurus) and a large genus similar to extant Type 2 North Atlantic killer whales (Plesiosuchus). With the description of the isolatedGeosaurustooth, all four Geosaurini genera are now known from the Lower Kimmeridgian Clay Formation. The craniodental plasticity of Geosaurini, and the astonishing range of feeding mechanisms they evolved, is unparalleled amongst thalattosuchians.http://www.zoobank.org/urn:lsid:zoobank.org:act:36B1FB4C-C20E-41C2-BC19-BAAB4E9D1612. [ABSTRACT FROM PUBLISHER]

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]

Re-description of a long-known, but little studied, metriorhynchid crocodylomorph from the Oxfordian of England begins to help fill the ‘Corallian Gap’. This ‘Gap’ represents a paucity of vertebrate remains from the Oxfordian of England. Filling the ‘Corallian Gap’ has important implications for our understanding of metriorhynchids from England, especially as there was a shift in this ecosystem from being metriorhynchine dominated in the older Oxford Clay Formation, to a geosaurine-dominated fauna in the younger Kimmeridge Clay Formation. The re-description of the specimen reveals that it is a member of the derived subclade Geosaurini, which most likely pertains to the genusTorvoneustes. [ABSTRACT FROM PUBLISHER]

The Oxford Clay Formation of England has yielded numerous sympatric species of metriorhynchid crocodylomorphs, although disagreement has persisted regarding the number of valid species. For over 140 years teeth reminiscent of the genusDakosaurushave been known from the Oxford Clay Formation but these have never been properly described and their taxonomy and systematic affinity remain contentious. Furthermore, an enigmatic mandible and associated postcranial skeleton discovered by Alfred Leeds in the Fletton brick pits near Peterborough also remains undescribed. We show that this specimen, and several isolated teeth, represents the oldest known remains of a large-bodied predatory metriorhynchid. This material is described herein and referred toTyrannoneustes lythrodectikosgen. et sp. nov. This species has a unique occlusal pattern: the dentition was arranged so that the posterior maxillodentary teeth interlock in the same plane and occlude mesiodistally. It is the first described crocodylomorph with microscopic denticles that are not contiguous along the carinae (forming short series of up to 10 denticles) and do not noticeably alter the height of the keel. Additionally, the dorsally expanded and curved posterior region of the mandible ventrally displaced the dentary tooth row relative to the jaw joint facilitating the enlargement of the dentition and increasing optimum gape. Therefore,Tyrannoneusteswould have been a large-bodied marine predator that was well-suited to feed on larger prey than other contemporaneous metriorhynchids. A new phylogenetic analysis findsTyrannoneustesto be the sister taxon to the subclade Geosaurini. An isolated tooth, humerus, and well-preserved mandible suggest a second species of metriorhynchid super-predator may also have lived in the Oxford Clay sea. Finally, we revise the diagnoses and descriptions of the other Oxford Clay metriorhynchid species, providing a guide for differentiating the many contemporaneous taxa from this exceptional fossil assemblage. http://zoobank.org/urn:lsid:zoobank.org:pub:C6222858-EC36-430D-8E28-1F636A6697CA [ABSTRACT FROM PUBLISHER]

Metriorhynchids are the only crocodyliforms adapted to pelagic marine life. Snout natural endocasts of the Tithonian ( Late Jurassic) metriorhynchid Cricosaurus araucanensis indicated that skeletal changes defining the peculiar metriorhynchid body plan were coupled with changes of the soft cephalic anatomy such as the enlarged salt glands and restructuring of the paranasal sinus system. Seven new natural endocasts of the snout and a 3-D reconstruction of C. araucanensis are described. Data from these casts and the reconstruction are congruent, and they are combined into an accurate reconstruction that improves our knowledge of the pre-orbital anatomy. The olfactory tract, bulbs, olfactory nasal region and the anterior extension of the antorbital sinus within the maxilla are recognized. Osteological correlates of the salt gland body are also proposed. Palaeobiological inferences are erected based on the integration of natural endocasts and 3-D reconstruction data. It is proposed that C. araucanensis nasal salt glands were highly vascularized with a blood supply comparable with those of extant marine birds. Reduced olfactory bulbs and olfactory nasal region indicate that the aerial olfaction, differing from extant crocodilians, was not well developed. [ABSTRACT FROM AUTHOR]

Metriorhynchoid thalattosuchians were a marine clade of Mesozoic crocodylomorphs that evolved from semi-aquatic, "gharial"-like species into the obligately pelagic subclade Metriorhynchidae. To explore whether the sensory and physiological demands of underwater life necessitates a shift in rostral anatomy, both in neurology and vasculature, we investigate the trigeminal innervation and potential somatosensory abilities of metriorhynchoids by digitally segmenting the rostral neurovascular canals in CT scans of 10 extant and extinct crocodyliforms. The dataset includes the terrestrial, basal crocodyliform Protosuchus haughtoni, two semi-aquatic basal metriorhynchoids, four pelagic metriorhynchids and three extant, semi-aquatic crocodylians. In the crocodylian and basal metriorhynchoid taxa, we find three main neurovascular channels running parallel to one another posteroanteriorly down the length of the snout, whereas in metriorhynchids there are two, and in P. haughtoni only one. Crocodylians appear to be unique in their extensive trigeminal innervation, which is used to supply the integumentary sensory organs (ISOs) involved with their facial somatosensory abilities. Crocodylians have a far higher number of foramina on the maxillary bones than either metriorhynchoids or P. haughtoni, suggesting that the fossil taxa lacked the somatosensory abilities seen in extant species. We posit that the lack of ISO osteological correlates in metriorhynchoids is due to their basal position in Crocodyliformes, rather than a pelagic adaptation. This is reinforced by the hypothesis that extant crocodyliforms, and possibly some neosuchian clades, underwent a long "nocturnal bottleneck"-hinting that their complex network of ISOs evolved in Neosuchia, as a sensory trade-off to compensate for poorer eyesight., (© 2021 American Association for Anatomy.)

Metriorhynchid crocodylomorphs were the only group of archosaurs to fully adapt to a pelagic lifestyle. During the Jurassic and Early Cretaceous, this group diversified into a variety of ecological and morphological types, from large super-predators with a broad short snout and serrated teeth to specialized piscivores/teuthophages with an elongate tubular snout and uncarinated teeth. Here, we use an integrated repertoire of geometric morphometric (form), biomechanical finite-element analysis (FEA; function) and phylogenetic data to examine the nature of craniofacial evolution in this clade. FEA stress values significantly correlate with morphometric values representing skull length and breadth, indicating that form and function are associated. Maximum-likelihood methods, which assess which of several models of evolution best explain the distribution of form and function data on a phylogenetic tree, show that the two major metriorhynchid subclades underwent different evolutionary modes. In geosaurines, both form and function are best explained as evolving under 'random' Brownian motion, whereas in metriorhynchines, the form metrics are best explained as evolving under stasis and the function metric as undergoing a directional change (towards most efficient low-stress piscivory). This suggests that the two subclades were under different selection pressures, and that metriorhynchines with similar skull shape were driven to become functionally divergent. [ABSTRACT FROM AUTHOR]

Abstract: Metriorhynchidae is a clade of marine-adapted crocodilians known from several Middle Jurassic–Early Cretaceous specimens collected predominantly in South America and Europe, but poorly known in the northern margin of Gondwana. The “Portomaggiore crocodile” is the most complete specimen of an Italian metriorhynchid to date: it consists of a partial skeleton that has been provisionally referred to an unnamed species of Late Jurassic Metriorhynchus or Geosaurus. The specimen is preserved in the reddish, nodular limestone of the Rosso Ammonitico Veronese Formation (Bajocian–Tithonian); new data on microfossil associations constrain the age of the metriorhynchid to the late Bajocian–earliest Bathonian. On the basis of cranial synapomorphies, the “Portomaggiore crocodile” falls as the closest sister-taxon of the Late Jurassic–Early Cretaceous geosaurines, and is referred to Neptunidraco ammoniticus gen. et sp. nov. It is unique among Middle Jurassic metriorhynchids in showing an incipient streamlining of the skull, shared with Late Jurassic and Cretaceous taxa. Since Neptunidraco is the oldest known member of Metriorhynchidae, its phylogenetic position supports the hypothesis that the timing of the initial metriorhynchid and geosaurine diversifications should start in the Bajocian. [ABSTRACT FROM AUTHOR]