Skull osteology, neuroanatomy, and jaw-related myology of the pig-nosed turtle Carettochelys insculpta (Cryptodira, Trionychia).

The osteology, neuroanatomy, and musculature are known for most primary clades of turtles (i.e., "families"), but knowledge is still lacking for one particular clade, the Carettochelyidae. Carettochelyids are represented by only one living taxon, the pig-nosed turtle Carettochelys insculpta. Here, we use micro-computed tomography of osteological and contrast-enhanced stained specimens to describe the cranial osteology, neuroanatomy, circulatory system, and jaw musculature of Carettochelys insculpta. The jaw-related myology is described in detail for the first time for this taxon, including m. zygomaticomandibularis, a muscular unit only found in trionychians. We also document a unique arterial pattern for the internal carotid artery and its subordinate branches and provide an extensive list of osteological ontogenetic differences. The present work provides new insights into the craniomandibular anatomy of turtles and will allow a better understanding of the evolutionary history of the circulatory system of trionychians and intraspecific variation among turtles.

A pan-chelydrid, Chelydropsis aubasi sp. nov., from the Middle Eocene (MP 15, early Bartonian) of Chéry-Chartreuve, France.

Snapping turtles (Pan-Chelydridae) play an important role in modern ecosystems throughout North America, but their fossil record is notably poor. We here describe a new species of fossil pan-chelydrid, Chelydropsis aubasi, from the Middle Eocene (MP15, Bartonian) of Chéry-Chartreuve, Department of Aisne, France, based on a series of fragments that document most of the shell. The new species not only bridges the morphological gap between earlier pan-chelydrids from North America and later pan-chelydrids from Europe but also advances the arrival of the group from North America to Europe by at least 10 million years. Although pan-chelydrids are notably absent from Early and Middle Eocene localities throughout Europe, the new fossil gives credence to the idea that pan-chelydrids may have dispersed from North America to Europe in concert with other turtles during the Paleocene-Eocene Thermal Maximum, but remain undetected to date.

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.

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.

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.

New interpretation of the cranial osteology of the Early Cretaceous turtle Arundelemys dardeni (Paracryptodira) based on a CT-based re-evaluation of the holotype.

Arundelemys dardeni is an Early Cretaceous paracryptodire known from a single, incomplete, but generally well-preserved skull. Phylogenetic hypotheses of paracryptodires often find Arundelemys dardeni as an early branching baenid. As such, it has a central role in understanding the early evolution of the successful clade Baenidae, which survived the Cretaceous-Paleogene mass extinction, as well as the diversification of Paracryptodira into its subclades, which recent research suggests to perhaps include helochelydrids, compsemydids, pleurosternids, and baenids. Computer tomography scans of the holotype material that were produced for the initial description of Arundelemeys dardeni reveal several errors in the initial anatomical description of the species, which we correct based on element-by-element segmentation. In addition, we provide entirely novel anatomical information, including descriptions of several previously undescribed cranial bones, the endosseous labyrinth, and the cranial scutes, the latter of which are unknown for most paracryptodires. We provide an interpretation of cranial scutes which homologizes the scutes of Arundelemys dardeni with those of other stem turtles.

A new pelomedusoid turtle, Sahonachelys mailakavava, from the Late Cretaceous of Madagascar provides evidence for convergent evolution of specialized suction feeding among pleurodires.

The Maevarano Formation in northwestern Madagascar has yielded a series of exceptional fossils over the course of the last three decades that provide important insights into the evolution of insular ecosystems during the latest Cretaceous (Maastrichtian). We here describe a new genus and species of pelomedusoid turtle from this formation, Sahonachelys mailakavava, based on a nearly complete skeleton. A phylogenetic analysis suggests close affinities of Sahonachelys mailakavava with the coeval Madagascan Sokatra antitra. These two taxa are the only known representatives of the newly recognized clade Sahonachelyidae, which is sister to the speciose clade formed by Bothremydidae and Podocnemidoidae. A close relationship with coeval Indian turtles of the clade Kurmademydini is notably absent. A functional assessment suggests that Sahonachelys mailakavava was a specialized suction feeder that preyed upon small-bodied invertebrates and vertebrates. This is a unique feeding strategy among crown pelomedusoids that is convergent upon that documented in numerous other clades of turtles and that highlights the distinct evolutionary pathways taken by Madagascan vertebrates.

A review of the carotid artery and facial nerve canal systems in extant turtles.

The cranial circulation and innervation systems of turtles have been studied for more than two centuries and extensively used to understand turtle systematics. Although a significant number of studies related to these structures exists, a broader comprehension of variation across the tree has been hindered by poor sampling and a lack of synthetic studies that addressed both systems together. We here provide new insights regarding the carotid circulation and facial nerve innervation systems in a broad set of extant turtles using CT (computed tomography) scans, which allow us to trace the canals these structures form in bone and understand the interaction between both systems. We document that the palatine artery, including the lateral carotid canal, is absent in all pleurodires and carettochelyids and was likely reduced or lost several times independently within Testudinoidea. We also highlight osteological correlates for the location of the mandibular artery. We finally summarize variation regarding the placement of the mandibular artery, location of the geniculate ganglion, placement of the hyomandibular and vidian nerves, and situations where we recommend caution when assessing canals in fossils. A morphometric study confirms that the relative sizes of the carotid canals are correlated with one another. Our results have the potential for building new phylogenetic characters and investigating the circulation systems of fossil taxa, which are expected to shed light on the evolution of the circulation system of turtles and clarify some unresolved relationships between fossil turtle clades.

Cranial osteology of the Early Cretaceous turtle Pleurosternon bullockii (Paracryptodira: Pleurosternidae).

Pleurosternon bullockii is a turtle from the Early Cretaceous of Europe known from numerous postcranial remains. Only one skull has so far been referred to the species. Pleurosternon bullockii belongs to a group of turtles called pleurosternids, which is thought to include several poorly known taxa from the Late Jurassic and Early Cretaceous of Europe and North America. Pleurosternids and baenids, a group of North American turtles that lived from the Late Cretaceous to the Eocene, define a clade called Paracryptodira. Additionally, Paracryptodira likely includes compsemydids, and, potentially, helochelydrids. Character support for Paracryptodira is relatively weak, and many global phylogenetic studies fail to support paracryptodiran monophyly altogether. Proposed paracryptodiran synapomorphies are largely cranial, despite the poor characterization of pleurosternid cranial material. In addition to their questionable monophyly, the global position of paracryptodires is debated. Early studies suggest crown-turtle affinities, but most phylogenies find them as stem-turtles, irrespective of their monophyly. Here, we document the cranial osteology of Pleurosternon bullockii with the use of three-dimensional models derived from segmenting high-resolution X-ray micro-computed tomography (CT) scans. Pleurosternon bullockii has a primitive basipterygoid region of the skull, but a cryptodire-like acustico-jugular region. A surprising number of similarities with pleurodires exist, particularly in the laterally expanded external process of the pterygoid and in the posterior orbital wall. Our observations constitute an important step toward a phylogenetic re-evaluation of Paracryptodira.