Description and phylogenetic relationships of a new species of Torvoneustes (Crocodylomorpha, Thalattosuchia) from the Kimmeridgian of Switzerland.

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.

A new species of the large-headed coastal marine turtle Solnhofia (Testudinata, Thalassochelydia) from the Late Jurassic of NW Switzerland.

BACKGROUND: The large-headed turtle Solnhofia parsonsi is known by a handful of specimens from the Late Jurassic of Germany and Switzerland (maybe also France). Solnhofia parsonsi is traditionally regarded as a "eurysternid" Thalassochelydia, a group of small to medium sized, mostly lagoonal or marginal turtles found almost exclusively in the Late Jurassic of Europe. More recently, Solnhofia parsonsi has been proposed to be a close relative of Sandownidae, an enigmatic group of Cretaceous to Paleogene turtles characterized by a derived cranial anatomy and a wider geographical distribution. Sandownids may therefore have evolved from thalassochelydian ancestors such as Solnhofia parsonsi. METHODS: We herein describe new material of Solnhofia from the Kimmeridgian (Late Jurassic) of Porrentruy, NW Switzerland. The bulk of the material consists of an association of a cranium and over 180 shell bones found together in a block of marly limestone. A second cranium and a mandible from slightly younger, but nearby localities are also described. RESULTS: We refer the new material to Solnhofia brachyrhyncha n. sp. The new species shares with Solnhofia parsonsi a relatively large head, an extensive secondary palate formed primarily by the maxillae, a greatly developed processus trochlearis oticum with a contribution from the parietal and quadratojugal, a large jugal-palatine contact in the floor of the fossa orbitalis, and a posteromedial process of the jugal running on the dorsal surface of the maxilla and pterygoid. Some of these characteristics are also present in sandownids, but our morphological study clearly shows that Solnhofia brachyrhyncha is closer to Solnhofia parsonsi than to any sandownids. DISCUSSION: Solnhofia brachyrhyncha differs from Solnhofia parsonsi in many aspects, notably: a shortened and broader cranium, a shorter and posteriorly broader upper triturating surface with a slightly sinusoidal lateral margin and without contribution from the palatine, a processus trochlearis oticum more oblique in dorsal or ventral view and less concave in anterior view, choanae that do not extend posteriorly on the pterygoids, a more developed processus pterygoideus externus, a condylus mandibularis situated anterior to the level of the occipital plane, a greater ventral exposure of the parabasisphenoid, a mandible about as wide as long, a relatively short symphysis, a lower triturating surface widened posterolaterally thanks to the presence of large laterally projecting dentary tubercles, a stouter and shorter coronoid process, a splenial positioned more anteriorly along the mandibular ramus, costo-peripheral fontanelles extending more anteriorly and posteriorly along the costal series, and an escutcheon shaped central plastral fontanelle formed mostly by the hypoplastra. In addition to the morphology of the new species, we also briefly discuss about observable ontogenetic variations and possible taphonomic origin of the assemblage.

Correction: Novel insights into the morphology of Plesiochelys bigleri from the early Kimmeridgian of Northwestern Switzerland.

[This corrects the article DOI: 10.1371/journal.pone.0214629.].

Novel insights into the morphology of Plesiochelys bigleri from the early Kimmeridgian of Northwestern Switzerland.

Plesiochelyidae were relatively large coastal marine turtles, which inhabited the epicontinental seas of Western Europe during the Late Jurassic. Their fossil record can be tracked in Germany, Switzerland, the United Kingdom, France, Spain and Portugal. The Jura Mountains, in northwestern Switzerland, have been the main source for the study of this group, mostly thanks to the rich and famous historical locality of Solothurn. In the last two decades, numerous plesiochelyid remains have been collected from Kimmeridgian deposits (Lower Virgula Marls and Banné Marls) in the area of Porrentruy (Canton of Jura, Switzerland). This material was revealed by construction works of the A16 Transjurane highway between 2000 and 2011, and led to the recent description of the new species Plesiochelys bigleri. In the years 2014 and 2016, new fragmentary turtle material was collected from the Banné Marls (Reuchenette Formation, lower Kimmeridgian) near the village of Glovelier, Canton of Jura, Switzerland. The new material consists of a complete shell, additional shell elements, a few bones from the appendicular and vertebral skeleton, and a fragmentary basicranium. This material can be confidently assigned to the species P. bigleri. It supports the presence of this species in the Banné Marls, slightly extends its spatial distribution and confirms the differences with the closely related species P. etalloni. The new material reveals that the split between the cerebral and palatine branches of the internal carotid artery occurs in a vertical plane in P. bigleri. This condition could not be observed in the type material due to poor preservation. This new character clearly distinguishes P. bigleri from P. etalloni and seems to be unique among thalassochelydians.

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.

A Jurassic stem pleurodire sheds light on the functional origin of neck retraction in turtles.

Modern turtles are composed of two monophyletic groups, notably diagnosed by divergent neck retraction mechanisms. Pleurodires (side-necked turtles) bend their neck sideways and protect their head under the anterior margin of the carapace. Cryptodires (hidden-necked turtles) withdraw their neck and head in the vertical plane between the shoulder girdles. These two mechanisms of neck retraction appeared independently in the two lineages and are usually assumed to have evolved for protective reasons. Here we describe the neck of Platychelys oberndorferi, a Late Jurassic early stem pleurodire, and find remarkable convergent morphological and functional similarities with modern cryptodires. Partial vertical neck retraction in this taxon is interpreted to have enabled fast forward projection of the head during underwater prey capture and offers a likely explanation to the functional origin of neck retraction in modern cryptodires. Complete head withdrawal for protection may therefore have resulted from an exaptation in that group.

First report of Plesiochelys etalloni and Tropidemys langii from the Late Jurassic of the UK and the palaeobiogeography of plesiochelyid turtles.

Plesiochelyidae is a clade of relatively large coastal marine turtles that inhabited the shallow epicontinental seas that covered western Europe during the Late Jurassic. Although the group has been reported from many deposits, the material is rarely identified at the species level. Here, we describe historical plesiochelyid material from the Kimmeridge Clay Formation of England and compare it with contemporaneous localities from the continent. An isolated basicranium is referred to the plesiochelyid Plesiochelys etalloni based notably on the presence of a fully ossified pila prootica. This specimen represents the largest individual known so far for this species and is characterized by remarkably robust features. It is, however, uncertain whether this represents an ontogenetic trend towards robustness in this species, some kind of specific variation (temporal, geographical or sexual), or an abnormal condition of this particular specimen. Four other specimens from the Kimmeridge Clay are referred to the plesiochelyid Tropidemys langii. This contradicts a recent study that failed to identify this species in this formation. This is the first time, to the best of our knowledge, that the presence of Plesiochelys etalloni and Tropidemys langii is confirmed outside the Swiss and French Jura Mountains. Our results indicate that some plesiochelyids had a wide palaeobiogeographic distribution during the Kimmeridgian.

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.

Establishment of a lectotype for the species Plesiochelys langii Rütimeyer, 1873.

Background. In a recent paper, we proposed a lectotype for the species Plesiochelys langii Rütimeyer, 1873. However, we failed to register this publication in ZooBank as required by the International Commission on Zoological Nomenclature (ICZN) for electronic publication. Although our conclusions remain unchanged, this particular nomenclatural act cannot be considered as published under ICZN regulations. Results. The present work fulfills the requirements of the ICZN for a lectotype designation and has been registered in ZooBank.

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.

The rediscovery and redescription of the holotype of the Late Jurassic turtle Plesiochelys etalloni.

Plesiochelyidae are a major component of Late Jurassic shallow marine environments throughout Europe. However, the taxonomy of plesiochelyid turtles is rather confused. Over the years, many taxa have been synonymized with Plesiochelys etalloni, one of the first described species. However, the holotype of P. etalloni (and only specimen known from Lect, the type locality) was lost for more than 150 years. This specimen has been recently rediscovered in the collections of the Musée d'archéologie du Jura in Lons-le-Saunier, France. For the first time since its original description in 1857, the holotype of P. etalloni is redescribed and compared to relevant material. The taxonomic status of this taxon is revised accordingly. Based on the morphology of the newly rediscovered holotype and on a reassessment of specimens from Solothurn (Switzerland), the species P. solodurensis, P. sanctaeverenae and P. langii are synonymized with P. etalloni. Known skull-shell associations for P. etalloni are re-evaluated in light of the new morphological information available since the rediscovery of this holotype specimen. Finally, we confirm that Plesiochelys is represented by a single species in the Late Jurassic of the Jura Mountains.

A new stem turtle from the Middle Jurassic of Scotland: new insights into the evolution and palaeoecology of basal turtles.

The discovery of a new stem turtle from the Middle Jurassic (Bathonian) deposits of the Isle of Skye, Scotland, sheds new light on the early evolutionary history of Testudinata. Eileanchelys waldmani gen. et sp. nov. is known from cranial and postcranial material of several individuals and represents the most complete Middle Jurassic turtle described to date, bridging the morphological gap between basal turtles from the Late Triassic-Early Jurassic and crown-group turtles that diversify during the Late Jurassic. A phylogenetic analysis places the new taxon within the stem group of Testudines (crown-group turtles) and suggests a sister-group relationship between E. waldmani and Heckerochelys romani from the Middle Jurassic of Russia. Moreover, E. waldmani also demonstrates that stem turtles were ecologically diverse, as it may represent the earliest known aquatic turtle.