A globally distributed durophagous marine reptile clade supports the rapid recovery of pelagic ecosystems after the Permo-Triassic mass extinction.

Marine ecosystem recovery after the Permo-Triassic mass extinction (PTME) has been extensively studied in the shallow sea, but little is known about the nature of this process in pelagic ecosystems. Omphalosauridae, an enigmatic clade of open-water durophagous marine reptiles, potentially played an important role in the recovery, but their fragmentary fossils and uncertain phylogenetic position have hindered our understanding of their role in the process. Here we report the large basal ichthyosauriform Sclerocormus from the Early Triassic of China that clearly demonstrates an omphalosaurid affinity, allowing for the synonymy of the recently erected Nasorostra with Omphalosauridae. The skull also reveals the anatomy of the unique feeding apparatus of omphalosaurids, likely an adaptation for feeding on hard-shelled pelagic invertebrates, especially ammonoids. Morphofunctional analysis of jaws shows that omphalosaurids occupy the morphospace of marine turtles. Our discovery adds another piece of evidence for an explosive radiation of marine reptiles into the ocean in the Early Triassic and the rapid recovery of pelagic ecosystems after the PTME.

Bone histology and microanatomy of Edaphosaurus and Dimetrodon (Amniota, Synapsida) vertebrae from the Lower Permian of Texas.

Here we describe the histology and microanatomy of vertebral centra of the iconic pelycosaur-grade synapsids Edaphosaurus boanerges and Dimetrodon spp. Vertebrae from different axial positions and, in the case of Dimetrodon, from different ontogenetic stages were selected. For the histological description, we produced histological petrographic thin sections ground to a thickness of 50-80 µm of the vertebrae in the sagittal and transversal cutting planes. After the preparation process, the thin sections were examined under transmitted and cross-polarized light in a polarized microscope. The analyzed vertebrae reveal similar bone tissues, where both taxa have cortical parallel-fibered bone (PFB). PFB and lamellar bone (LB) forms in the cancellous part. However, in juvenile Dimetrodon, woven-fibered bone (WFB) is also deposited and shows a high degree of vascularity. This suggests that Dimetrodon had slightly faster bone growth than Edaphosaurus, which is mainly made of PFB and LB and shows poorly developed vascular canals. In addition, one specimen of Dimetrodon displays the preservation of an ossified notochord, which can be assumed to be indicative of how the intervertebral tissues were developed. Historically, evidence of how the joint between Dimetrodon vertebral centra was built was lacking until this specimen appeared. If the notochord ran persistently through the vertebrae, it would have possibly increased the stiffness of the vertebral column and would have affected the limbs and locomotion. Furthermore, the organization of trabeculae and relative thickness of the vertebral cortex gives insights into how the animals were adapted to their habitat.

Early giant reveals faster evolution of large body size in ichthyosaurs than in cetaceans.

Body sizes of marine amniotes span six orders of magnitude, yet the factors that governed the evolution of this diversity are largely unknown. High primary production of modern oceans is considered a prerequisite for the emergence of cetacean giants, but that condition cannot explain gigantism in Triassic ichthyosaurs. We describe the new giant ichthyosaur Cymbospondylus youngorum sp. nov. with a 2-meter-long skull from the Middle Triassic Fossil Hill Fauna of Nevada, USA, underscoring rapid size evolution despite the absence of many modern primary producers. Surprisingly, the Fossil Hill Fauna rivaled the composition of modern marine mammal faunas in terms of size range, and energy-flux models suggest that Middle Triassic marine food webs were able to support several large-bodied ichthyosaurs at high trophic levels, shortly after ichthyosaur origins.

Palaeontological evidence reveals convergent evolution of intervertebral joint types in amniotes.

The intervertebral disc (IVD) has long been considered unique to mammals. Palaeohistological sampling of 17 mostly extinct clades across the amniote tree revealed preservation of different intervertebral soft tissue types (cartilage, probable notochord) seen in extant reptiles. The distribution of the fossilised tissues allowed us to infer the soft part anatomy of the joint. Surprisingly, we also found evidence for an IVD in fossil reptiles, including non-avian dinosaurs, ichthyosaurs, plesiosaurs, and marine crocodiles. Based on the fossil dataset, we traced the evolution of the amniote intervertebral joint through ancestral character state reconstruction. The IVD evolved at least twice, in mammals and in extinct diapsid reptiles. From this reptilian IVD, extant reptile groups and some non-avian dinosaurs independently evolved a synovial ball-and-socket joint. The unique birds dorsal intervertebral joint evolved from this dinosaur joint. The tuatara and some geckos reverted to the ancestral persisting notochord.

Conflicting evidence for the use of caudal autotomy in mesosaurs.

The early Permian mesosaurs were the first amniotes to re-invade aquatic environments. One of their most controversial and puzzling features is their distinctive caudal anatomy, which has been suggested as a mechanism to facilitate caudal autotomy. Several researchers have described putative fracture planes in mesosaur caudal vertebrae - unossified regions in the middle of caudal vertebral centra - that in many extant squamates allow the tail to separate and the animal to escape predation. However, the reports of fracture planes in mesosaurs have never been closely investigated beyond preliminary descriptions, which has prompted scepticism. Here, using numerous vertebral series, histology, and X-ray computed tomography, we provide a detailed account of fracture planes in all three species of mesosaurs. Given the importance of the tail for propulsion in many other aquatic reptiles, the identification of fracture planes in mesosaurs has important implications for their aquatic locomotion. Despite mesosaurs apparently having the ability to autotomize their tail, it is unlikely that they actually made use of this behaviour due to a lack of predation pressure and no record of autotomized tails in articulated specimens. We suggest that the presence of fracture planes in mesosaurs is an evolutionary relic and could represent a synapomorphy for an as-yet undetermined terrestrial clade of Palaeozoic amniotes that includes the earliest radiation of secondarily aquatic tetrapods.

Hematological convergence between Mesozoic marine reptiles (Sauropterygia) and extant aquatic amniotes elucidates diving adaptations in plesiosaurs.

Plesiosaurs are a prominent group of Mesozoic marine reptiles, belonging to the more inclusive clades Pistosauroidea and Sauropterygia. In the Middle Triassic, the early pistosauroid ancestors of plesiosaurs left their ancestral coastal habitats and increasingly adapted to a life in the open ocean. This ecological shift was accompanied by profound changes in locomotion, sensory ecology and metabolism. However, investigations of physiological adaptations on the cellular level related to the pelagic lifestyle are lacking so far. Using vascular canal diameter, derived from osteohistological thin-sections, we show that inferred red blood cell size significantly increases in pistosauroids compared to more basal sauropterygians. This change appears to have occurred in conjunction with the dispersal to open marine environments, with cell size remaining consistently large in plesiosaurs. Enlarged red blood cells likely represent an adaptation of plesiosaurs repeated deep dives in the pelagic habitat and mirror conditions found in extant marine mammals and birds. Our results emphasize physiological aspects of adaptive convergence among fossil and extant marine amniotes and add to our current understanding of plesiosaur evolution.

Neck mobility in the Jurassic plesiosaur Cryptoclidus eurymerus: finite element analysis as a new approach to understanding the cervical skeleton in fossil vertebrates.

The sauropterygian clade Plesiosauria arose in the Late Triassic and survived to the very end of the Cretaceous. Plesiosauria evolved the greatest species diversity of any marine reptile clade, attaining a global distribution. Plesiosauria consist of two clades, Rhomaleosauridae and Neoplesiosauria. Basal Neoplesiosauria have long necks with at least 30 cervicals, but show qualitative osteological evidence for a stiff neck. Here we quantify neck mobility in lateral, ventral, and dorsal directions based on finite element modeling of neck vertebrae from the Middle Jurassic plesiosaur Cryptoclidus eurymerus. We model the mobility in a single motion segment, consisting of two adjacent cervical vertebrae and the joints connecting them. Based on the model with a maximum intervertebral spacing of 3 mm, we find that in Cryptoclidus, the maximum angle of lateral deflection in the motion segment was 2°. The maximum angle of ventral deflection was 5° and of dorsal deflection was 5°. When these values are multiplied by the number of cervical vertebrae, it becomes apparent that neck mobility was limited in all directions. The maximum angle of total lateral deflection in the neck was 67°. The maximum angle of total ventral deflection was 148° and of total dorsal deflection was 157°. This raises the question of the function of such a long, multi-segment but immobile neck. We posit that the long neck served in hydrodynamic and visual camouflage, hiding the bulk of the body from the small but abundant prey, such as schooling fish and squid. Neck immobility may have been advantageous in withstanding strong hydrodynamic forces acting on the neck during predatory strikes.

Quantitative histological models suggest endothermy in plesiosaurs.

BACKGROUND: Plesiosaurs are marine reptiles that arose in the Late Triassic and survived to the Late Cretaceous. They have a unique and uniform bauplan and are known for their very long neck and hydrofoil-like flippers. Plesiosaurs are among the most successful vertebrate clades in Earth's history. Based on bone mass decrease and cosmopolitan distribution, both of which affect lifestyle, indications of parental care, and oxygen isotope analyses, evidence for endothermy in plesiosaurs has accumulated. Recent bone histological investigations also provide evidence of fast growth and elevated metabolic rates. However, quantitative estimations of metabolic rates and bone growth rates in plesiosaurs have not been attempted before. METHODS: Phylogenetic eigenvector maps is a method for estimating trait values from a predictor variable while taking into account phylogenetic relationships. As predictor variable, this study employs vascular density, measured in bone histological sections of fossil eosauropterygians and extant comparative taxa. We quantified vascular density as primary osteon density, thus, the proportion of vascular area (including lamellar infillings of primary osteons) to total bone area. Our response variables are bone growth rate (expressed as local bone apposition rate) and resting metabolic rate (RMR). RESULTS: Our models reveal bone growth rates and RMRs for plesiosaurs that are in the range of birds, suggesting that plesiosaurs were endotherm. Even for basal eosauropterygians we estimate values in the range of mammals or higher. DISCUSSION: Our models are influenced by the availability of comparative data, which are lacking for large marine amniotes, potentially skewing our results. However, our statistically robust inference of fast growth and fast metabolism is in accordance with other evidence for plesiosaurian endothermy. Endothermy may explain the success of plesiosaurs consisting in their survival of the end-Triassic extinction event and their global radiation and dispersal.

A Triassic plesiosaurian skeleton and bone histology inform on evolution of a unique body plan.

Secondary marine adaptation is a major pattern in amniote evolution, accompanied by specific bone histological adaptations. In the aftermath of the end-Permian extinction, diverse marine reptiles evolved early in the Triassic. Plesiosauria is the most diverse and one of the longest-lived clades of marine reptiles, but its bone histology is least known among the major marine amniote clades. Plesiosaurians had a unique and puzzling body plan, sporting four evenly shaped pointed flippers and (in most clades) a small head on a long, stiffened neck. The flippers were used as hydrofoils in underwater flight. A wide temporal, morphological, and morphometric gap separates plesiosaurians from their closest relatives (basal pistosaurs, Bobosaurus). For nearly two centuries, plesiosaurians were thought to appear suddenly in the earliest Jurassic after the end-Triassic extinctions. We describe the first Triassic plesiosaurian, from the Rhaetian of Germany, and compare its long bone histology to that of later plesiosaurians sampled for this study. The new taxon is recovered as a basal member of the Pliosauridae, revealing that diversification of plesiosaurians was a Triassic event and that several lineages must have crossed into the Jurassic. Plesiosaurian histology is strikingly uniform and different from stem sauropterygians. Histology suggests the concurrent evolution of fast growth and an elevated metabolic rate as an adaptation to cruising and efficient foraging in the open sea. The new specimen corroborates the hypothesis that open ocean life of plesiosaurians facilitated their survival of the end-Triassic extinctions.