First record of a pterosaur landing trackway.

The terrestrial progression of pterosaurs, the flying reptiles of the Mesozoic Era, has been debated for over two centuries. The recent discovery of quadrupedal pterodactyloid pterosaur tracks from Late Jurassic sediments near Crayssac, France, shows that the hindlimbs moved parasagittally, as in mammals, birds and other dinosaurs, and the hypertrophied forelimbs could make tracks both close to the body wall and far outside it. Their manus tracks are unique in form, position and kinematics, which would be expected because the forelimbs were used for flight. Here, we report the first record of a pterosaur landing track, which differs substantially from typical walking trackways. The individual landed on both hind feet in parallel fashion, dragged its toes slightly as it left the track, landed again almost immediately and placed the hindfeet parallel again, then placed its forelimbs on the ground, took another short step with both hindlimbs and adjusted its forelimbs, and then began to walk off normally. The trackway shows that pterosaurs stalled to land, a reflection of their highly developed capacity for flight control and manoeuverability.

The comparative osteology of Plesiochelys bigleri n. sp., a new coastal marine turtle from the Late Jurassic of Porrentruy (Switzerland).

BACKGROUND: During the Late Jurassic, several groups of eucryptodiran turtles inhabited the shallow epicontinental seas of Western Europe. Plesiochelyidae are an important part of this first radiation of crown-group turtles into coastal marine ecosystems. Fossils of Plesiochelyidae occur in many European localities, and are especially abundant in the Kimmeridgian layers of the Swiss Jura Mountains (Solothurn and Porrentruy). In the mid-19th century, the quarries of Solothurn (NW Switzerland) already provided a large amount of fossil turtles, most notably Plesiochelys etalloni, the best-known plesiochelyid species. Recent excavations in the Porrentruy area (NW Switzerland) revealed new fossils of Plesiochelys, including numerous well-preserved shells with associated cranial and postcranial material. METHODS/RESULTS: Out of 80 shells referred to Plesiochelys, 41 are assigned to a new species, Plesiochelys bigleri n. sp., including a skull-shell association. We furthermore refer 15 shells to Plesiochelys etalloni, and 24 shells to Plesiochelys sp. Anatomical comparisons show that Plesiochelys bigleri can clearly be differentiated from Plesiochelys etalloni by cranial features. The shell anatomy and the appendicular skeleton of Plesiochelys bigleri and Plesiochelys etalloni are very similar. However, a statistical analysis demonstrates that the thickness of neural bones allows to separate the two species based on incomplete material. This study furthermore illustrates the extent of intraspecific variation in the shell anatomy of Plesiochelys bigleri and Plesiochelys etalloni.

Thalassemys bruntrutana n. sp., a new coastal marine turtle from the Late Jurassic of Porrentruy (Switzerland), and the paleobiogeography of the Thalassemydidae.

Background. The Swiss Jura Mountains are a key region for Late Jurassic eucryptodiran turtles. Already in the mid 19th century, the Solothurn Turtle Limestone (Solothurn, NW Switzerland) yielded a great amount of Kimmeridgian turtles that are traditionally referred to Plesiochelyidae, Thalassemydidae, and Eurysternidae. In the past few years, fossils of these coastal marine turtles were also abundantly discovered in the Kimmeridgian of the Porrentruy region (NW Switzerland). These findings include numerous sub-complete shells, out of which we present two new specimens of Thalassemys (Thalassemydidae) in this study. Methods. We compare the new material from Porrentruy to the type species Th. hugii, which is based on a well preserved specimen from the Solothurn Turtle Limestone (Solothurn, Switzerland). In order to improve our understanding of the paleogeographic distribution of Thalassemys, anatomical comparisons are extended to Thalassemys remains from other European countries, notably Germany and England. Results. While one of the two Thalassemys specimens from Porrentruy can be attributed to Th. hugii, the other specimen represents a new species, Th. bruntrutana n. sp. It differs from Th. hugii by several features: more elongated nuchal that strongly thickens anterolaterally; wider vertebral scales; proportionally longer plastron; broader and less inclined xiphiplastron; wider angle between scapular process and acromion process. Our results show that Th. hugii and Th. bruntrutana also occur simultaneously in the Kimmeridgian of Solothurn as well as in the Kimmeridgian of England (Kimmeridge Clay). This study is an important step towards a better understanding of the paleobiogeographic distribution of Late Jurassic turtles in Europe.

Portlandemys gracilis n. sp., a New Coastal Marine Turtle from the Late Jurassic of Porrentruy (Switzerland) and a Reconsideration of Plesiochelyid Cranial Anatomy.

BACKGROUND: Several groups of stem cryptodires became adapted to coastal marine environments as early as the Late Jurassic, 40 million years before the Pan-Chelonioidea. The Plesiochelyidae are a major component of this first radiation of crown-group turtles into marine habitats. They are abundant in many European localities, but their systematics is still greatly confused. Only three species are represented by cranial material: Plesiochelys etalloni, Plesiochelys planiceps, and Portlandemys mcdowelli. METHODOLOGY/PRINCIPAL FINDINGS: In the present study, we describe a cranium and a mandible from the Kimmeridgian of Porrentruy (Switzerland), which we refer to a new species, Portlandemys gracilis n. sp. This new taxon differs from Portlandemys mcdowelli in several aspects of the cranium and mandible, notably in being generally more gracile, but the two species share a narrow skull, a more acute angle between the labial ridges on the mandible, and a unique configuration of the anterodorsal part of the basicranium. The cranial anatomy of plesiochelyid turtles is discussed in details based primarily on these new specimens and new cranial material of Plesiochelys etalloni from Solothurn, Switzerland. CONCLUSIONS/SIGNIFICANCE: Several characters (e.g., the contribution of the parietal to the foramen nervi trigemini, the configuration of the dorsum sellae and sella turcica, the presence of an infolding ridge on the posterior surface of the quadrate) appear as potential candidates to help elucidate plesiochelyid relationships. Some of these characters are included in a previously published phylogenetic dataset and help to stabilize the relationships of plesiochelyid turtles and closely related taxa. For the first time, our results suggest that plesiochelyids, 'Thalassemys' moseri, and Solnhofia parsonsi (representing the Eurysternidae) form a clade at the base of Eucryptodira.

A taxonomic review of the Late Jurassic eucryptodiran turtles from the Jura Mountains (Switzerland and France).

Background. Eucryptodiran turtles from the Late Jurassic (mainly Kimmeridgian) deposits of the Jura Mountains (Switzerland and France) are among the earliest named species traditionally referred to the Plesiochelyidae, Thalassemydidae, and Eurysternidae. As such, they are a reference for the study of Late Jurassic eucryptodires at the European scale. Fifteen species and four genera have been typified based on material from the Late Jurassic of the Jura Mountains. In the past 50 years, diverging taxonomic reassessments have been proposed for these turtles with little agreement in sight. In addition, there has been a shift of focus from shell to cranial anatomy in the past forty years, although most of these species are only represented by shell material. As a result, the taxonomic status of many of these 15 species remains ambiguous, which prevents comprehensive comparison of Late Jurassic turtle assemblages throughout Europe and hinders description of new discoveries, such as the new assemblage recently unearthed in the vicinity of Porrentruy, Switzerland. Methods. An exhaustive reassessment of the available material provides new insights into the comparative anatomy of these turtles. The taxonomic status of each of the 15 species typified based on material from the Late Jurassic of the Jura Mountains is evaluated. New diagnoses and general descriptions are provided for each valid taxon. Results. Six out of the 15 available species names are recognized as valid: Plesiochelys etalloni, Craspedochelys picteti, Craspedochelys jaccardi, Tropidemys langii, Thalassemys hugii, and 'Thalassemys' moseri. The intraspecific variability of the shell of P. etalloni is discussed based on a sample of about 30 relatively complete specimens from Solothurn, Switzerland. New characters are proposed to differentiate P. etalloni, C. picteti, and C. jaccardi, therefore rejecting the previously proposed synonymy of these forms. Based partly on previously undescribed specimens, the plastral morphology of Th. hugii is redescribed. The presence of lateral plastral fontanelles is notably revealed in this species, which calls into question the traditional definitions of the Thalassemydidae and Eurysternidae. Based on these new data, Eurysternum ignoratum is considered a junior synonym of Th. hugii. The Eurysternidae are therefore only represented by Solnhofia parsonsi in the Late Jurassic of the Jura Mountains. Finally, 'Th.' moseri is recognized as a valid species, although a referral to the genus Thalassemys is refuted.