Cranial anatomy,taxonomic implications and palaeopathology of an Upper Jurassic Pliosaur (Reptilia: Sauropterygia) from Westbury,Wiltshire, UK

Abstract: Complete skulls of giant marine reptiles of the Late Jurassic are rare, and so the discovery of the 1.8‐m‐long skull of a pliosaur from the Kimmeridge Clay Formation (Kimmeridgian) of Westbury, Wiltshire, UK, is an important find. The specimen shows most of the cranial and mandibular anatomy, as well as a series of pathological conditions. It was previously referred to Pliosaurus brachyspondylus, but it can be referred reliably only to the genus Pliosaurus, because species within the genus are currently in need of review. The new specimen, together with another from the same locality, also referred to P. brachyspondylus, will be crucial in that systematic revision, and it is likely that the genus Pliosaurus will be found to include several genera. The two Westbury Pliosaurus specimens share many features, including the form of the teeth, but marked differences in the snout and parietal crest suggest sexual dimorphism; the present specimen is probably female. The large size of the animal, the extent of sutural fusion and the pathologies suggest this is an ageing individual. An erosive arthrotic condition of the articular glenoids led to prolonged jaw misalignment, generating a suite of associated bone and dental pathologies. Further damage to the jaw joint may have been the cause of death.     

MARINE REPTILES FROM THE LOWER CRETACEOUS OF SOUTH AUSTRALIA: ELEMENTS OF A HIGH-LATITUDE COLD-WATER ASSEMBLAGE

Abstract: The Lower Cretaceous rocks of South Australia have yielded a diverse marine reptile assemblage of up to five families of plesiosaur (including a new cryptoclidid or cimoliasaurid, indeterminate elasmosaurids, a possible polycotylid, rhomaleosaurids, and pliosaurid) and one family of ichthyosaur (ophthalmosaurid). Other common associated vertebrates include chimaerids and osteichthyans. Sharks, dipnoans and dinosaurs are uncommon and marine turtles are notably absent. The main fossil‐producing strata belong to the Lower Aptian–Lower Albian Bulldog Shale although the Upper Albian Oodnadatta Formation has produced isolated elements. Both these units comprise finely laminated shaly mudstones and claystones deposited in a transgressive shallow coastal, epicontinental marine environment. Estimates of palaeolatitude place South Australia between 60° and 70°S, in the late Early Cretaceous. Sedimentary structures (including lonestone boulders and glendonites), fossils, isotope data and climatic modelling also indicate that seasonally cool–cold conditions (possibly with winter freezing) prevailed during deposition of the Bulldog Shale. This contrasts markedly with climate regimes typically tolerated by modern aquatic reptiles but suggests that some of the South Australian Mesozoic taxa may have possessed adaptations (including elevated metabolic levels and/or annual migration) to cope with low temperatures. A high proportion of juvenile plesiosaur remains in the Bulldog Shale might also indicate that nutrient‐rich cold‐water coastal habitats functioned as both ‘safe calving grounds’ and refuges for young animals prior to their entering the open sea as adults. The occurrence of plesiosaurs and ichthyosaurs in the high‐latitude Lower Cretaceous of southern Australia, along with plesiosaurs and mosasaurs in the Upper Cretaceous of South America, Antarctica, New Zealand and the Chatham Islands, demonstrates that Mesozoic marine reptiles utilized southern high‐latitude environments over a considerable period of time, and that these records do not represent casual occupation by isolated taxa.     

The true world's first sculptures of antediluvian animals, which never were…

In 1852, the French state commissioned the artists Frémiet and Jacquemart to execute bronzes of a plesiosaur and a pterodactyl for the Jardin des Plantes in Paris. The orders were cancelled before the sculptures could be realized, largely because of petty jealousies among the professors of the Muséum national d’Histoire naturelle, who maintained that the long-extinct animals were too poorly understood for accurate reconstructions. In this way an important opportunity to educate and inspire the French public about the life of the past was lost.     

Charles de Gerville et les premiers restes de Plesiosauria signalés dans le Jurassique inférieur de France (Hettangien,région de Valognes,Manche)

The historian, archaeologist, and naturalist Charles de Gerville (1769–1853) reported as early as January 1816 the presence of fossil bones in the quarries of the region of Valognes, in the department of Manche. These quarries exploited the “Calcaire de Valognes” and the underlying “Argiles et Calcaires d’Huberville”, both formations dated as Lower Hettangian (Lower Jurassic). De Gerville communicated to Jacques-Louis-Marin Defrance a watercolour drawing of one of these fossil bones, discovered in July 1820, in which it is possible to recognize a plesiosaur femur. This femur, as well as a fragment of rib and three vertebrae of Plesiosauria from de Gerville's collection, was then figured in a publication by Arcisse de Caumont, in 1825. The plesiosaur femur was later acquired by Pierre Tesson of Caen, who owned one of the most important collections of fossils in Normandy. The Tesson collection was later purchased in 1857 by the British Museum (now The Natural History Museum, London), where the femur has been rediscovered. This specimen is the first known plesiosaur specimen ever discovered in the Lower Jurassic of France, before the genus Plesiosaurus was established upon material from the Lias of southwestern England. One of the vertebrae figured by de Caumont has been rediscovered in the “Muséum Emmanuel-Liais” (Cherbourg-en-Cotentin), which had acquired part of de Gerville's collection. The interpretation of the “fossil bones of Valognes” by Georges Cuvier, Henri Marie Ducrotay de Blainville, Arcisse de Caumont, Jacques-Amand Eudes-Deslongchamps and Constant Duméril is revealed through unpublished documents.     

New information on Bobosaurus forojuliensis (Reptilia: Sauropterygia): implications for plesiosaurian evolution

Preparation of the holotype specimen of Bobosaurus forojuliensis, a large sauropterygian from the lower Carnian of northeastern Italy, revealed new morphological data relevant in establishing its phylogenetic affinities among pistosauroid taxa and its relationships with plesiosaurians. Inclusion of B. forojuliensis in two phylogenetic analyses focusing, respectively, on sauropterygians and pistosauroids agreed in placing the Italian taxon as closer to plesiosaurians than to other pistosauroids. The phylogenetic interpretation of Bobosaurus was not biased by assumptions on character weighting, is consistent with its relatively younger age compared to most pistosauroids, extends the fossil record of the plesiosaurian basal lineage back to the Carnian and supports the earliest diversification of the clade during the Late Triassic in agreement with the record of several distinct lineages of rhomaleosaurids, plesiosauroids and pliosauroids in the lowermost Jurassic. Bobosaurus shows that the evolution of the plesiosaurian body plan from the ancestral pistosauroid grade was a step-wise process, and that some of the vertebral and appendicular specialisations of Jurassic and Cretaceous plesiosaurians had already developed in the earliest Late Triassic.     

Reassessment of the Lower Cretaceous (Barremian) pliosauroid Leptocleidus superstes Andrews, 1922 and other plesiosaur remains from the nonmarine Wealden succession of southern England

The Lower Cretaceous (upper Berriasian to lowermost Aptian) nonmarine Wealden succession of southern England has been a prolific source of vertebrate fossils for over 180 years. The sequence is most famous for terrestrial reptiles, particularly dinosaurs; however, significant aquatic tetrapod discoveries including rare nonmarine plesiosaurs have also been reported. The record of Wealden plesiosaurs currently incorporates a single valid taxon, Leptocleidus superstes Andrews, 1922a, based on a partial skeleton and skull from the Barremian Upper Weald Clay Formation of Berwick, Sussex. Traditional classifications place this plesiomorphic pliosauroid with basal Jurassic rhomaleosaurids; however, the genus Leptocleidus has since become a ‘waste basket’ for various Cretaceous rhomaleosaurid‐like plesiosaurs from around the globe. In an attempt to clarify this situation, the type specimen of L. superstes was reexamined and redescribed. Previously unrecorded anatomical features were identified including an elongate, gracile paraoccipital process on the exoccipital‐opisthotic, and tooth ornament comprising widely spaced, coarse striations that are restricted to the lingual surface of the crown (mesiodistal ‘carinae’ are absent). Other indeterminate pliosauroid remains (recovered along with coeval elasmosaurids) from the upper Berriasian–Valanginian Hastings Beds Group also exhibit potentially diagnostic traits: an atlas centrum with no anterolateral exposure and with ventral margin formed by the intercentrum; a single‐headed rib articulation on the atlas centrum extending onto the axis centrum; and epipodials that are longer than broad. The placement of L. superstes is controversial in recent phylogenies. To test the competing hypotheses, L. superstes together with all closely related species were rescored into the most comprehensive published phylogenetic data sets of Plesiosauria and Pliosauroidea. Separate maximum parsimony and Bayesian analyses of each matrix unanimously supported a relationship between L. superstes and pliosauroids but could not confirm placement within either Rhomaleosauridae sensu stricto, or a discrete ‘leptocleidoid’ clade. Examination of character states advocating affinities amongst Leptocleidus spp. suggests homoplasy rather than clear homology between what are potentially palaeobiogeographically disparate genus‐level taxa. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 663–691.     

The internal cranial anatomy of the Plesiosauria (Reptilia, Sauropterygia): evidence for a functional secondary palate

In the late 19th Century, the choanae (or internal nares) of the Plesiosauria were identified as a pair of palatal openings located rostral to the external nares, implying a rostrally directed respiratory duct and air path inside the rostrum. Despite obvious functional shortcomings, this idea was firmly established in the scientific literature by the first decade of the 20th Century. The functional consequences of this morphology were only re-examined by the end of the 20th Century, leading to the conclusion that the choanae were not involved in respiration but instead in underwater olfaction, the animals supposedly breathing with the mouth agape. Re-evaluation of the palatal and internal cranial anatomy of the Plesiosauria reveals that the traditional identification of the choanae as a pair of fenestrae situated rostral to the external nares appears erroneous. These openings more likely represent the bony apertures of ducts that lead to internal salt glands situated inside the maxillary rostrum. The 'real' functional choanae (or caudal interpterygoid vacuities), are situated at the caudal end of the bony palate between the sub-temporal fossae, as was suggested in the mid-19th Century. The existence of a functional secondary palate in the Plesiosauria is therefore strongly supported, and the anatomical, physiological, and evolutionary implications of such a structure are discussed.     

Global interrelationships of Plesiosauria (Reptilia,Sauropterygia) and the pivotal role of taxon sampling in determining the outcome of phylogenetic analyses

Previous attempts to resolve plesiosaurian phylogeny are reviewed and a new phylogenetic data set of 66 taxa (67% of ingroup taxa examined directly) and 178 characters (eight new) is presented. We recover two key novel results: a monophyletic Plesiosauridae comprising Plesiosaurus dolichodeirus, Hydrorion brachypterygius, Microcleidus homalospondylus, Occitanosaurus tournemirensis and Seeleyosaurus guilelmiimperatoris; and five plesiosaurian taxa recovered outside the split between Plesiosauroidea and Pliosauroidea. These taxa are Attenborosaurus conybeari, ‘Plesiosaurus’macrocephalus and a clade comprising Archaeonectrus rostratus, Macroplata tenuiceps and BMNH 49202. Based on this result, a new name, Neoplesiosauria, is erected for the clade comprising Plesiosauroidea and Pliosauroidea. Taxon subsamples of the new dataset are used to simulate previous investigations of global plesiosaurian relationships. Based on these simulations, most major differences between previous global phylogenetic hypotheses can be attributed to differences in taxon sampling. These include the position of Leptocleididae and Polycotylidae and the monophyly or paraphyly of Rhomaleosauridae. On this basis we favour the results recovered by our, larger analysis. Leptocleididae and Polycotylidae are sister taxa, forming a monophyletic clade within Plesiosauroidea, indicating that the large‐headed, short‐necked ‘pliosauromorph’ body plan evolved twice within Plesiosauria. Rhomaleosauridae forms the monophyletic sister taxon of Pliosauridae within Pliosauroidea. Problems are identified with previous phylogenetic definitions of plesiosaurian clades and new, stem‐based definitions are presented that should maintain their integrity over a range of phylogenetic hypotheses. New, rank‐free clade names Cryptoclidia and Leptocleidia are erected to replace the superfamilies Cryptoclidoidea and Leptocleidoidea. These were problematic as they were nested within the superfamily Plesiosauroidea. The incongruence length difference test indicates no significant difference in levels of homoplasy between cranial and postcranial characters.     

A new specimen of the Triassic pistosauroid Yunguisaurus,with implications for the origin of Plesiosauria (Reptilia,Sauropterygia)

An adult skeleton of the pistosauroid sauropterygian Yunguisaurus liae reveals a number of morphological features not observed in the holotype, such as the complete morphology of the skull roof, stapes, atlas and axis, ventral view of the postcranium, and nearly complete limbs and tail. Size and morphological differences between the two specimens are mostly regarded as ontogenetic variation, and newly added data did not affect the phylogenetic relationships with other pistosauroids significantly. The number of mesopodia (11 carpals and 8 tarsals) exceeds that known in any other Triassic marine reptiles and does not serve as a precursor of the plesiosaurian pattern with fewer mesopodia of different topology; it demonstrates variability of the limb morphology among the Triassic pistosauroids. The pectoral girdles of Corosaurus, Augustasaurus and Yunguisaurus may indicate early stages of the adaptation towards the plesiosaurian style of paraxial limb movements with ventroposterior power stroke.