Updated cranial and mandibular description of the Late Cretaceous (Maastrichtian) baenid turtle Saxochelys gilberti based on micro-computed tomography scans and new information on the holotype-shell association.

Saxochelys gilberti is a baenid turtle from the Late Cretaceous Hell Creek Formation of the United States of America known from cranial, shell, and other postcranial material. Baenid turtles are taxonomically diverse and common fossil elements within Late Cretaceous through Eocene faunas. Detailed anatomical knowledge is critical to understanding the systematics and morphological evolution of the group. This is particularly important as baenids represent an important group of continental vertebrates that survived the mass extinction event associated with the Cretaceous/Paleogene boundary. High-resolution micro-computed tomography scanning of the holotype skull reveals additional anatomical details for the already well-known Saxochelys gilberti. This includes the revision of some anatomical statements from the original description, but also detailed knowledge on internal anatomical features of the braincase and the description of a well-preserved axis (cervical vertebra 2). Our new detailed description and previous work on the shell and postcrania make Saxochelys one of the best-described, nearly complete baenid turtles, which are often only known from either isolated shell or cranial material. A revised phylogenetic analysis confirms the position of Saxochelys gilberti as a derived baenid (Eubaeninae) more closely related to Baena arenosa than to Eubaena cephalica. Supplementary Information: The online version contains supplementary material available at 10.1186/s13358-023-00301-6.

Turtle body size evolution is determined by lineage-specific specializations rather than global trends.

Organisms display a considerable variety of body sizes and shapes, and macroevolutionary investigations help to understand the evolutionary dynamics behind such variations. Turtles (Testudinata) show great body size disparity, especially when their rich fossil record is accounted for. We explored body size evolution in turtles, testing which factors might influence the observed patterns and evaluating the existence of long-term directional trends. We constructed the most comprehensive body size dataset for the group to date, tested for correlation with paleotemperature, estimated ancestral body sizes, and performed macroevolutionary model-fitting analyses. We found no evidence for directional body size evolution, even when using very flexible models, thereby rejecting the occurrence of Cope's rule. We also found no significant effect of paleotemperature on overall through-time body size patterns. In contrast, we found a significant influence of habitat preference on turtle body size. Freshwater turtles display a rather homogeneous body size distribution through time. In contrast, terrestrial and marine turtles show more pronounced variation, with terrestrial forms being restricted to larger body sizes, up to the origin of testudinids in the Cenozoic, and marine turtles undergoing a reduction in body size disparity after the extinctions of many groups in the mid-Cenozoic. Our results, therefore, suggest that long-term, generalized patterns are probably explained by factors specific to certain groups and related at least partly to habitat use.

A well-preserved cranium from the Judith River Formation (Montana, USA) reveals the inner ear and neuroanatomy of a Campanian baenid turtle.

A cranium belonging to a baenid turtle was recently recovered from the lower half of the Judith River Formation, Montana. Badlands Dinosaur Museum (BDM) 004 is a well-preserved partial cranium that includes the posterior cranial vault, cranial base, and otic capsules. Based on diagnostic characters, the skull can be attributed to Plesiobaena antiqua, which has been previously reported from the Judith River Formation. It also shares with palatobaenines projecting posterior processes of the tubercula basioccipitale and a prominent condylus occipitalis with a deep central pit, demonstrating variation within the Pl. antiqua hypodigm. In a phylogenetic analysis, an operational taxonomic unit of BDM 004 was positioned within Baenodda in an unresolved polytomy with Pl. antiqua, Edowa zuniensis, Palatobaeninae, and Eubaeninae. Microcomputed tomographic (µCT) scans revealed morphology of the middle and inner ear and endocast that are largely unknown in baenids. Semicircular canals of BDM 004 are virtually identical to those of Eubaena cephalica and consistent in dimensions to those of other turtle taxa, including anterior and posterior semicircular canals that are robust and taller than the common crus and diverge from each other at an angle of approximately 90°. The digital endocast reveals a moderately flexed brain with rounded cerebral hemispheres and minimal separation between the metencephalon and myelencephalon. Its well-preserved columella auris (stapes) is gracile with a posterodorsally flared basis columella. It arcs across the middle ear and flattens near its terminus. This study adds to the understanding of baenid middle and inner ear and neuroanatomical morphology and expands the morphological understanding of Pl. antiqua.

100 million years of turtle paleoniche dynamics enable the prediction of latitudinal range shifts in a warming world.

Past responses to environmental change provide vital baseline data for estimating the potential resilience of extant taxa to future change. Here, we investigate the latitudinal range contraction that terrestrial and freshwater turtles (Testudinata) experienced from the Late Cretaceous to the Paleogene (100.5-23.03 mya) in response to major climatic changes. We apply ecological niche modeling (ENM) to reconstruct turtle niches, using ancient and modern distribution data, paleogeographic reconstructions, and the HadCM3L climate model to quantify their range shifts in the Cretaceous and late Eocene. We then use the insights provided by these models to infer their probable ecological responses to future climate scenarios at different representative concentration pathways (RCPs 4.5 and 8.5 for 2100), which project globally increased temperatures and spreading arid biomes at lower to mid-latitudes. We show that turtle ranges are predicted to expand poleward in the Northern Hemisphere, with decreased habitat suitability at lower latitudes, inverting a trend of latitudinal range contraction that has been prevalent since the Eocene. Trionychids and freshwater turtles can more easily track their niches than Testudinidae and other terrestrial groups. However, habitat destruction and fragmentation at higher latitudes will probably reduce the capability of turtles and tortoises to cope with future climate changes.

Cranial osteology, taxonomic reassessment, and phylogenetic relationships of the Early Cretaceous (Aptian-Albian) turtle Trinitichelys hiatti (Paracryptodira).

We describe the skull of the Early Cretaceous (Aptian-Albian) baenid turtle Trinitichelys hiatti using micro-computed tomography to provide new insights into the cranial anatomy of basal baenids and into the evolution of paracryptodires. We show that the validity of Trinitichelys hiatti vs Arundelemys dardeni still holds true, that the most basal known baenids for which skull material is known share an intriguing combination of features that are typical of either Pleurosternidae or Baenidae, and that the carotid system of Trinitichelys hiatti is intermediate to that of pleurosternids and more advanced baenids. Our expanded phylogenetic analysis confirms the traditional placement of Arundelemys dardeni, Lakotemys australodakotensis, and Trinitichelys hiatti as basal baenids, retrieves Helochelydridae along the stem of Baenoidea, but recovers Dinochelys whitei, Glyptops ornatus, Dorsetochelys typocardium, and Uluops uluops as basal branching Paracryptodira.

A review of helochelydrid shell material from late Albian to early Cenomanian greensands of Southern England, United Kingdom.

A number of helochelydrid turtle shell remains were recovered over the course of the 19th century from mid-Cretaceous sediments throughout Southern England, including the poorly figured and described types of Trachydermochelys phlyctaenus from the Cambridge Greensand of Cambridgeshire, Plastremys lata from the Upper Greensand of the Isle of Wight, and "Trachydermochelys" rutteri from the Melbury Sandstone of Dorset. A review of stratigraphic provenience suggests that all material originates from late Late Aptian portions of the Upper Greensand or early Early Cenomanian portions of the West Melbury Marly Chalk Formation, a relatively narrow time window geologically speaking. As described, Trachydermochelys phlyctaenus is a problematic taxon, because the most plausible type series is a chimera that includes two helochelydrid morphotypes in addition to protostegid remains. This conundrum is resolved through the designation of a lectotype. A review of all historic material confirms the presence of three English taxa distinct from Helochelydra danubina, a coeval taxon named from Germany. At least four helochelydrid taxa, therefore, occurred in western Europe during the Early to Late Cretaceous transition.

New insights into the cranial osteology of the Early Cretaceous paracryptodiran turtle Lakotemys australodakotensis.

Lakotemys australodakotensis is an Early Cretaceous paracryptodire known from two shells and a skull from the Lakota Formation of South Dakota, USA. Along with the Early Cretaceous Arundelemys dardeni and the poorly known Trinitichelys hiatti, Lakotemys australodakotensis is generally retrieved as an early branching baenid, but more insights into the cranial anatomy of these taxa is needed to obtain a better understanding of paracryptodiran diversity and evolution. Here, we describe the skull of Lakotemys australodakotensis using micro-computed tomography to provide the anatomical basis for future phylogenetic analyses that will be needed to investigate more precisely the intrarelationships of Paracryptodira. Preliminary comparisons reveal that the cranial anatomy of Lakotemys australodakotensis is very similar to that of the Aptian-Albian basal baenid Arundelemys dardeni, that both taxa exhibit a remarkable combination of derived characters found in baenodds and characters found in non-baenid paracryptodires, particularly Pleurosternidae, and that Lakotemys australodakotensis is the only known baenid to date to possess a canal for the palatine artery.

Symmetry-Breaking Stabilities in Carapace Curvature on Testudo (Reptilia, Testudinidae).

The aim of this research was to contribute to the study of the doming geometry of Testudo carapace as an unstable point of equilibrium when animals are overturned. We performed this research using geometric morphometric using a sample of 64 Testudo individuals belonging to different species (T. hermannin = 30, T. graecan = 3, T. marginata n = 13 and T. horsfieldii n = 18), sexes and ages. A set of four sagittal landmarks (discrete homologous points) and 15 pairs of semi-landmarks, on the frontal doming of the carapace, were digitized on individual carapace pictures. Significative fluctuating asymmetry was detected, defined as small, completely random departures from bilateral symmetry, but much less than directional asymmetry, which appeared highly significative. Anti-symmetry did not appear. Carapace asymmetry was dominated by a clear right directionality. A possible biological speculation could be that this asymmetry more that easing the self-righting potential ("kinematic instability", understood as the ability to self-right without effort), makes stable ventral turning difficult ("static stability", understood as the ability to resist passively turning the body produced by destabilizing forces). This asymmetry is present among both sexes but more marked among males. An explanation for this sexually differentiated pattern could be the higher locomotion and the fight for mating in males, making them consequently more prone to losing their balance and falling on their back. These data may be useful in studying adaptative traits in Testudo species as well as establishing a seminal base for future studies. This research is the first attempt to explore a suitable method to assess doming asymmetry which could be useful in future, more extensive investigations, on a larger interspecific sample.

A redescription of the Late Jurassic (Tithonian) turtle Uluops uluops and a new phylogenetic hypothesis of Paracryptodira.

We study the Late Jurassic (Tithonian) turtle Uluops uluops using micro-computed tomography scans to investigate the cranial anatomy of paracryptodires, and provide new insights into the evolution of the internal carotid artery and facial nerve systems, as well as the phylogenetic relationships of this group. We demonstrate the presence of a canalis caroticus lateralis in Uluops uluops, the only pleurosternid for which a palatine artery canal can be confidently identified. Our phylogenetic analysis retrieves Uluops uluops as the earliest branching pleurosternid, Helochelydridae within Pleurosternidae, and Compsemydidae including Kallokibotion bajazidi within Baenidae, which suggests at least two independent losses of the palatine artery within paracryptodires. We expect future studies will provide additional insights into the evolution of the circulation system of paracryptodires, as well as clarifying relationships along the turtle stem. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1186/s13358-021-00234-y.

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.

A reevaluation of the basal turtle Indochelys spatulata from the Early-Middle Jurassic (Toarcian-Aalenian) of India, with descriptions of new material.

BACKGROUND: Indochelys spatulata is an extinct turtle from the Early to Middle Jurassic Kota Formation of the Pranhita-Godavari Gondwana basin, India. The holotype and previously only known specimen is a partially eroded shell that had been collected near Kota village, north of Sironcha, in Maharashtra State. Phylogenetic analyses have consistently suggested placement at the base of the clade Mesochelydia. METHODS: We here figure and describe the holotype of Indochelys spatulata and two new specimens, which were collected from the Kota Formation near Kistapur village, Telengana State, about 60 km NW from the type locality. We furthermore explore the relationships of this fossil turtle by updating its scoring based on all available material in the most recent analysis of basal turtle relationships. RESULTS: The revision of the holotype of Indochelys spatulata provides minor adjustments to the morphology of this specimen, in particular recognition of a transverse break across the carapace, presence of only eight neurals, of which the eight is octagonal, and presence of a pathological element located between neurals VII and VIII. The new material provides new anatomical insights, in particular presence of a broad cervical, a vertebral V that inserts deeply into vertebral IV, narrow pleurals within increasingly short posteromedial contacts with the vertebrals towards the posterior, at least three pairs of musk duct foramina, and numerous insights into the morphology of the girdles and stylopodium. In combination, all material allows affirming the validity of Indochelys spatulata with confidence. The phylogenetic analysis affirms the placement of Indochelys spatulata as a basal mesochelydian, but cannot resolve its relationships relative to the roughly coeval Condorchelys antiqua and Kayentachelys aprix.

The ecomorphology of the shell of extant turtles and its applications for fossil turtles.

Turtles are a successful clade of reptiles that originated in the Late Triassic. The group adapted during its evolution to different types of environments, ranging from dry land to ponds, rivers, and the open ocean, and survived all Mesozoic and Cenozoic extinction events. The body of turtles is characterized by a shell, which has been hypothesized to have several biological roles, like protection, thermal and pH regulation, but also to be adapted in its shape to the ecology of the animal. However, only few studies have investigated the relationships between shell shape and ecology in a global context or clarified if shape can be used to diagnose habitat preferences in fossil representatives. Here, we assembled a three-dimensional dataset of 69 extant turtles and three fossils, in particular, the Late Triassic Proganochelys quenstedtii and Proterochersis robusta and the Late Jurassic Plesiochelys bigleri to test explicitly for a relationship between shell shape and ecology. 3D models were obtained using surface scanning and photogrammetry. The general shape of the shells was captured using geometric morphometrics. The habitat ecology of extant turtles was classified using the webbing of their forelimbs as a proxy. Principal component analysis (PCA) highlights much overlap between habitat groups. Discriminant analyses suggests significant differences between extant terrestrial turtles, extant fully aquatic (i.e., marine and riverine) turtles, and an unspecialized assemblage that includes extant turtles from all habitats, mostly freshwater aquatic forms. The paleoecology of the three fossil species cannot be determined with confidence, as all three fall within the unspecialized category, even if Plesiochelys bigleri plots closer to fully aquatic turtles, while the two Triassic species group closer to extant terrestrial forms. Although the shape of the shell of turtles indeed contains an ecological signal, it is overall too weak to uncover using shell shape in paleoecological studies, at least with the methods we selected.

Rhinochelys amaberti Moret (1935), a protostegid turtle from the Early Cretaceous of France.

Modern marine turtles (chelonioids) are the remnants of an ancient radiation that roots in the Cretaceous. The oldest members of that radiation are first recorded from the Early Cretaceous and a series of species are known from the Albian-Cenomanian interval, many of which have been allocated to the widespread but poorly defined genus Rhinochelys, possibly concealing the diversity and the evolution of early marine turtles. In order to better understand the radiation of chelonioids, we redescribe the holotype and assess the taxonomy of Rhinochelys amaberti Moret (1935) (UJF-ID.11167) from the Late Albian (Stoliczkaia dispar Zone) of the Vallon de la Fauge (Isère, France). We also make preliminary assessments of the phylogenetic relationships of Chelonioidea using two updated datasets that widely sample Cretaceous taxa, especially Rhinochelys. Rhinochelys amaberti is a valid taxon that is supported by eight autapomorphies; an emended diagnosisis proposed. Our phylogenetic analyses suggest that Rhinochelys could be polyphyletic, but constraining it as a monophyletic entity does not produce trees that are significantly less parsimonious. Moreover, support values and stratigraphic congruence indexes are fairly low for the recovered typologies, suggesting that missing data still strongly affect our understanding of the Cretaceous diversification of sea turtles.

Shell variability in the stem turtles Proterochersis spp.

BACKGROUND: Turtle shells tend to exhibit frequent and substantial variability, both in bone and scute layout. Aside from secondary changes, caused by diseases, parasites, and trauma, this variability appears to be inherent and result from stochastic or externally induced flaws of developmental programs. It is, thus, expected to be present in fossil turtle species at least as prominently, as in modern populations. Descriptions of variability and ontogeny are, however, rare for fossil turtles, mainly due to rarity, incompleteness, damage, and post-mortem deformation of their remains. This paper is an attempt at description and interpretation of external shell variability in representatives of the oldest true turtles, Proterochersis robusta and Proterochersis porebensis (Proterochersidae, the sister group to all other known testudinatans) from the Late Triassic (Norian) of Germany and Poland. METHODS: All the available shell remains of Proterochersis robusta (13 specimens) and Proterochersis porebensis (275 specimens) were studied morphologically in order to identify any ontogenetic changes, intraspecific variability, sexual dimorphism, and shell abnormalities. To test the inferred sexual dimorphism, shape analyses were performed for two regions (gular and anal) of the plastron. RESULTS: Proterochersis spp. exhibits large shell variability, and at least some of the observed changes seem to be correlated with ontogeny (growth of gulars, extragulars, caudals, and marginals, disappearance of middorsal keel on the carapace). Several specimens show abnormal layout of scute sulci, several others unusual morphologies of vertebral scute areas, one has an additional pair of plastral scutes, and one extraordinarily pronounced, likely pathological, growth rings on the carapace. Both species are represented in a wide spectrum of sizes, from hatchlings to old, mature individuals. The largest fragmentary specimens of Proterochersis porebensis allow estimation of its maximal carapace length at approximately 80 cm, while Proterochersis robusta appears to have reached lower maximal sizes. DISCUSSION: This is the second contribution describing variability among numerous specimens of Triassic turtles, and the first to show evidence of unambiguous shell abnormalities. Presented data supplement the sparse knowledge of shell scute development in the earliest turtles and suggest that at least some aspects of the developmental programs governing scute development were already similar in the Late Triassic to these of modern forms.

Homeotic shift at the dawn of the turtle evolution.

All derived turtles are characterized by one of the strongest reductions of the dorsal elements among Amniota, and have only 10 dorsal and eight cervical vertebrae. I demonstrate that the Late Triassic turtles, which represent successive stages of the shell evolution, indicate that the shift of the boundary between the cervical and dorsal sections of the vertebral column occurred over the course of several million years after the formation of complete carapace. The more generalized reptilian formula of at most seven cervicals and at least 11 dorsals is thus plesiomorphic for Testudinata. The morphological modifications associated with an anterior homeotic change of the first dorsal vertebra towards the last cervical vertebra in the Triassic turtles are partially recapitulated by the reduction of the first dorsal vertebra in crown-group Testudines, and they resemble the morphologies observed under laboratory conditions resulting from the experimental changes of Hox gene expression patterns. This homeotic shift hypothesis is supported by the, unique to turtles, restriction of Hox-5 expression domains, somitic precursors of scapula, and brachial plexus branches to the cervical region, by the number of the marginal scute-forming placodes, which was larger in the Triassic than in modern turtles, and by phylogenetic analyses.

Archelosaurian Color Vision, Parietal Eye Loss, and the Crocodylian Nocturnal Bottleneck.

Vertebrate color vision has evolved partly through the modification of five ancestral visual opsin proteins via gene duplication, loss, and shifts in spectral sensitivity. While many vertebrates, particularly mammals, birds, and fishes, have had their visual opsin repertoires studied in great detail, testudines (turtles) and crocodylians have largely been neglected. Here I examine the genomic basis for color vision in four species of turtles and four species of crocodylians, and demonstrate that while turtles appear to vary in their number of visual opsins, crocodylians experienced a reduction in their color discrimination capacity after their divergence from Aves. Based on the opsin sequences present in their genomes and previous measurements of crocodylian cones, I provide evidence that crocodylians have co-opted the rod opsin (RH1) for cone function. This suggests that some crocodylians might have reinvented trichromatic color vision in a novel way, analogous to several primate lineages. The loss of visual opsins in crocodylians paralleled the loss of various anatomical features associated with photoreception, attributed to a "nocturnal bottleneck" similar to that hypothesized for Mesozoic mammals. I further queried crocodylian genomes for nonvisual opsins and genes associated with protection from ultraviolet light, and found evidence for gene inactivation or loss for several of these genes. Two genes, encoding parietopsin and parapinopsin, were additionally inactivated in birds and turtles, likely co-occurring with the loss of the parietal eye in these lineages.

Latitudinal diversity gradients in Mesozoic non-marine turtles.

The latitudinal biodiversity gradient (LBG)-the pattern of increasing taxonomic richness with decreasing latitude-is prevalent in the structure of the modern biota. However, some freshwater taxa show peak richness at mid-latitudes; for example, extant Testudines (turtles, terrapins and tortoises) exhibit their greatest diversity at 25° N, a pattern sometimes attributed to recent bursts of climatically mediated species diversification. Here, we test whether this pattern also characterizes the Mesozoic distribution of turtles, to determine whether it was established during either their initial diversification or as a more modern phenomenon. Using global occurrence data for non-marine testudinate genera, we find that subsampled richness peaks at palaeolatitudes of 15-30° N in the Jurassic, 30-45° N through the Cretaceous to the Campanian, and from 30° to 60° N in the Maastrichtian. The absence of a significant diversity peak in southern latitudes is consistent with results from climatic models and turtle niche modelling that demonstrate a dearth of suitable turtle habitat in Gondwana during the Jurassic and Late Cretaceous. Our analyses confirm that the modern testudinate LBG has a deep-time origin and further demonstrate that LBGs are not always expressed as a smooth, equator-to-pole distribution.

A toothed turtle from the Late Jurassic of China and the global biogeographic history of turtles.

BACKGROUND: Turtles (Testudinata) are a successful lineage of vertebrates with about 350 extant species that inhabit all major oceans and landmasses with tropical to temperate climates. The rich fossil record of turtles documents the adaptation of various sub-lineages to a broad range of habitat preferences, but a synthetic biogeographic model is still lacking for the group. RESULTS: We herein describe a new species of fossil turtle from the Late Jurassic of Xinjiang, China, Sichuanchelys palatodentata sp. nov., that is highly unusual by plesiomorphically exhibiting palatal teeth. Phylogenetic analysis places the Late Jurassic Sichuanchelys palatodentata in a clade with the Late Cretaceous Mongolochelys efremovi outside crown group Testudines thereby establishing the prolonged presence of a previously unrecognized clade of turtles in Asia, herein named Sichuanchelyidae. In contrast to previous hypotheses, M. efremovi and Kallokibotion bajazidi are not found within Meiolaniformes, a clade that is here reinterpreted as being restricted to Gondwana. CONCLUSIONS: A revision of the global distribution of fossil and recent turtle reveals that the three primary lineages of derived, aquatic turtles, including the crown, Paracryptodira, Pan-Pleurodira, and Pan-Cryptodira can be traced back to the Middle Jurassic of Euramerica, Gondwana, and Asia, respectively, which resulted from the primary break up of Pangaea at that time. The two primary lineages of Pleurodira, Pan-Pelomedusoides and Pan-Chelidae, can similarly be traced back to the Cretaceous of northern and southern Gondwana, respectively, which were separated from one another by a large desert zone during that time. The primary divergence of crown turtles was therefore driven by vicariance to the primary freshwater aquatic habitat of these lineages. The temporally persistent lineages of basal turtles, Helochelydridae, Meiolaniformes, Sichuanchelyidae, can similarly be traced back to the Late Mesozoic of Euramerica, southern Gondwana, and Asia. Given the ambiguous phylogenetic relationships of these three lineages, it is unclear if their diversification was driven by vicariance as well, or if they display a vicariance-like pattern. The clean, primary signal apparent among early turtles is secondarily obliterated throughout the Late Cretaceous to Recent by extensive dispersal of continental turtles and by multiple invasions of marine habitats.

Evolution of neck vertebral shape and neck retraction at the transition to modern turtles: an integrated geometric morphometric approach.

The unique ability of modern turtles to retract their head and neck into the shell through a side-necked (pleurodiran) or hidden-necked (cryptodiran) motion is thought to have evolved independently in crown turtles. The anatomical changes that led to the vertebral shapes of modern turtles, however, are still poorly understood. Here we present comprehensive geometric morphometric analyses that trace turtle vertebral evolution and reconstruct disparity across phylogeny. Disparity of vertebral shape was high at the dawn of turtle evolution and decreased after the modern groups evolved, reflecting a stabilization of morphotypes that correspond to the two retraction modes. Stem turtles, which had a very simple mode of retraction, the lateral head tuck, show increasing flexibility of the neck through evolution towards a pleurodiran-like morphotype. The latter was the precondition for evolving pleurodiran and cryptodiran vertebrae. There is no correlation between the construction of formed articulations in the cervical centra and neck mobility. An increasing mobility between vertebrae, associated with changes in vertebral shape, resulted in a more advanced ability to retract the neck. In this regard, we hypothesize that the lateral tucking retraction of stem turtles was not only the precondition for pleurodiran but also of cryptodiran retraction. For the former, a kink in the middle third of the neck needed to be acquired, whereas for the latter modification was necessary between the eighth cervical vertebra and first thoracic vertebra. Our paper highlights the utility of 3D shape data, analyzed in a phylogenetic framework, to examine the magnitude and mode of evolutionary modifications to vertebral morphology. By reconstructing and visualizing ancestral anatomical shapes, we provide insight into the anatomical features underlying neck retraction mode, which is a salient component of extant turtle classification.