Márton Rabi

A new xinjiangchelyid turtle from the Middle Jurassic of Xinjiang, China and the evolution of the basipterygoid process in Mesozoic turtles

BackgroundMost turtles from the Middle and Late Jurassic of Asia are referred to the newly defined clade Xinjiangchelyidae, a group of mostly shell-based, generalized, small to mid-sized aquatic froms that are widely considered to represent the stem lineage of Cryptodira. Xinjiangchelyids provide us with great insights into the plesiomorphic anatomy of crown-cryptodires, the most diverse group of living turtles, and they are particularly relevant for understanding the origin and early divergence of the primary clades of extant turtles.ResultsExceptionally complete new xinjiangchelyid material from the ?Qigu Formation of the Turpan Basin (Xinjiang Autonomous Province, China) provides new insights into the anatomy of this group and is assigned to Xinjiangchelys wusu n. sp. A phylogenetic analysis places Xinjiangchelys wusu n. sp. in a monophyletic polytomy with other xinjiangchelyids, including Xinjiangchelys junggarensis, X. radiplicatoides, X. levensis and X. latiens. However, the analysis supports the unorthodox, though tentative placement of xinjiangchelyids and sinemydids outside of crown-group Testudines. A particularly interesting new observation is that the skull of this xinjiangchelyid retains such primitive features as a reduced interpterygoid vacuity and basipterygoid processes.ConclusionsThe homology of basipterygoid processes is confidently demonstrated based on a comprehensive review of the basicranial anatomy of Mesozoic turtles and a new nomenclatural system is introduced for the carotid canal system of turtles. The loss of the basipterygoid process and the bony enclosure of the carotid circulation system occurred a number of times independently during turtle evolution suggesting that the reinforcement of the basicranial region was essential for developing a rigid skull, thus paralleling the evolution of other amniote groups with massive skulls.

A new species of the side-necked turtle Foxemys (Pelomedusoides: Bothremydidae) from the Late Cretaceous of Hungary and the historical biogeography of the Bothremydini

The continental deposits of the Upper Cretaceous (Santonian) Csehbanya Formation of the Bakony Mountains in Hungary yielded abundant remains of a bothremydid side-necked turtle, which are attributed to a new species of the genus Foxemys , Foxemys trabanti . F. trabanti shows strong affinities with the European monophyletic group Foxemydina owing to the absence of pits in the upper and lower triturating surfaces, the exclusion of the jugal from the triturating surface, the separation of the Eustachian tube and the stapes by a narrow fissure, the presence of deep and narrow fossa pterygoidei, the partially closed foramen jugulare posterius and the pentagonal shape of the basisphenoid in ventral view. Among the Foxemydina the bothremydid from Iharkut is more closely related to F. mechinorum than to Polysternon provinciale from the Early Campanian of France, mainly because of the position of the occipital condyle relative to the mandibular condyles of the quadrate. The new remains represent the only record of the Foxemydina outside of Western Europe and provide the earliest known occurrence of this endemic, freshwater group in the former Mediterranean Basin. The historical biogeography of the tribe Bothremydini is investigated and a hypothesis of migration from Africa to North America via the high-latitude Thulean route is put forward.

Osteology, relationships, and ecology of Annemys (Testudines, Eucryptodira) from the Late Jurassic of Shar Teg, Mongolia, and phylogenetic definitions for Xinjiangchelyidae, Sinemydidae, and Macrobaenidae

ABSTRACT A complete description of the xinjiangchelyid turtles Annemys levensis and A. latiens is provided, based on all available material from the Upper Jurassic type locality of Shar Teg, Mongolia. Annemys latiens was previously known almost exclusively from shell material, but an undescribed skull from Shar Teg is referable to this species and its distinct morphology confirms the presence of two taxa at this locality. Annemys latiens has an elongated skull that markedly differs in proportions from those of A. levensis and resembles the shape of some recent, piscivorous turtles. The overall similarity of the shells of the two Annemys species combined with significant differences in the skull indicate that these turtles probably partitioned the aquatic niche by exploring different feeding strategies. Among xinjiangchelyids, at least three different skull morphotypes can be differentiated, which implies a moderate level of ecological diversification among Late Jurassic Asian turtles. Phylogenetic analysis weakly supports the inclusion of Annemys spp. into Xinjiangchelyidae and places xinjiangchelyids at the stem of Testudines, but the latter result is considered tentative. Phylogenetic definitions of Xinjiangchelyidae, Sinemydidae, and Macrobaenidae are provided for nomenclatural clarity and precision.

Cranial morphology of Theriosuchus sympiestodon (Mesoeucrocodylia, Atoposauridae) and the widespread occurrence of Theriosuchus in the Late Cretaceous of Europe

Abstract We present a detailed morphological description of the type-locality cranial material of Theriosuchus sympiestodon Martin, Rabi, and Csiki, 2010 from the Maastrichtian Densuş-Ciula Formation of the Haţeg Basin, Romania together with new material of isolated cranial elements and teeth from various sites of the same general area. The recognition of several individuals of distinct sizes allows for an assessment of ontogenetic variation in this taxon. New material, consisting of isolated teeth and an incomplete maxilla with in situ teeth, coming from various late Campanian/early Maastrichtian sites in southern France is referable to ?Theriosuchus sp. and hints to a rare but widespread distribution of Theriosuchus in the Late Cretaceous European archipelago.

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

BackgroundTurtles (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.ResultsWe 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.ConclusionsA 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.

A sinemydid turtle from the Jehol Biota provides insights into the basal divergence of crown turtles.

Morphological phylogenies stand in a major conflict with molecular hypotheses regarding the phylogeny of Cryptodira, the most diverse and widely distributed clade of extant turtles. However, molecular hypotheses are often considered a better estimate of phylogeny given that it is more consistent with the stratigraphic and geographic distribution of extinct taxa. That morphology fails to reproduce the molecular topology partly originates from problematic character polarization due to yet another contradiction around the composition of the cryptodiran stem lineage. Extinct sinemydids are one of these problematic clades: they have been either placed among stem-cryptodires, stem-chelonioid sea turtles, or even stem-turtles. A new sinemydid from the Early Cretaceous Jehol Biota (Yixian Formation, Barremian-Early Aptian) of China, Xiaochelys ningchengensis gen. et sp. nov., allows for a reassessment of the phylogenetic position of Sinemydidae. Our analysis indicates that sinemydids mostly share symplesiomorphies with sea turtles and their purported placement outside the crown-group of turtles is an artefact of previous datasets. The best current phylogenetic estimate is therefore that sinemydids are part of the stem lineage of Cryptodira together with an array of other Jurassic to Cretaceous taxa. Our study further emphasises the importance of using molecular scaffolds in global turtle analyses.

A new specimen of Manchurochelys manchoukuoensis from the Early Cretaceous Jehol Biota of Chifeng, Inner Mongolia, China and the phylogeny of Cretaceous basal eucryptodiran turtles

BackgroundManchurochelys manchoukuoensis is an emblematic turtle from the Cretaceous Yixian Formation of Liaoning, China, a geological rock unit that is famous for yielding perfectly preserved skeletons of fossil vertebrates, including that of feathered dinosaurs. Manchurochelys manchoukuoensis was one of the first vertebrates described from this fauna, also known as the Jehol Biota. The holotype was lost during World War II and only one additional specimen has been described since. Manchurochelys manchoukuoensis is a critical taxon for unraveling the phylogenetic relationships of Cretaceous pancryptodires from Asia, a group that is considered to be of key importance for the origin of crown-group hidden-neck turtles (Cryptodira).ResultsA new specimen of Manchurochelys manchoukuoensis is described here from the Jiufotang Formation of Qilinshan, Chifeng, Inner Mongolia, China. This is the third specimen described and expands the range of this taxon from the Yixian Formation of the Fuxin-Yixian Basin in Liaoning to the Jiufotang Formation of the Chifeng-Yuanbaoshan Basin. A possible temporal extension of the range is less certain. The new finding adds to our understanding of the morphology of this taxon and invites a thorough revision of the phylogeny of Macrobaenidae, Sinemydidae, and closely allied forms.ConclusionsOur comprehensive phylogenetic analyses of Cretaceous Asian pancryptodires yielded two main competing hypotheses: in the first these taxa form a paraphyletic grade, whereas in the second they form a monophyletic clade. The inclusion of problematic tree changing taxa, such as Panpleurodires (stem + crown side-neck turtles) has a major influence on the phylogenetic relationships of Sinemydidae and closely allied forms. Manchurochelys manchoukuoensis nests within Sinemydidae together with Sinemys spp. and Dracochelys bicuspis in the majority of our analyses.

Gross morphology and microstructure of type locality ossicles of Psephophorus polygonus Meyer, 1847 (Testudines, Dermochelyidae)

Psephophorus polygonus Meyer, 1847, the first fossil leatherback turtle to be named, was described on the basis of shell ossicles from the middle Miocene (MN6-7/8?) of Slovakia. The whereabouts of this material is uncertain but a slab on display at the Naturhistorisches Museum Wien is considered the neotype. We rediscovered further type locality ossicles in four European institutions, re-evaluated their gross morphology and described for the first time their microstructure by comparing them with Dermochelys coriacea, the only living dermochelyid turtle. The gross morphology is congruent with that already described for P. polygonus, but with two significant exceptions: the ridged ossicles of P. polygonus may have a distinctly concave ventral surface as well as a tectiform shape in cross-section. They do not develop the external keel typical of many ossicles of D. coriacea. Both ridged and non-ridged ossicles of P. polygonus are characterized by compact diploe structures with an internal cortex consisting of a coarse fibrous meshwork, whereas the proportionately thinner ossicles of D. coriacea tend to lose the internal cortex, and thus their diploe, during ontogeny. The ossicles of both P. polygonus and D. coriacea differ from those of other lineages of amniotes whose carapace is composed of polygonal ossicles or platelets, in having growth centres situated at the plate centres just interior to the external bone surface and not within the cancellous core or closer to the internal compact layer. The new diagnosis of P. polygonus allows us to preliminarily re-evaluate the taxonomy of some of the Psephophorus-like species. Despite some macro-and micromorphological differences, it seems likely that Psephophorus was as cosmopolitan as extant Dermochelys and had a broadly similar ecology, with a possible difference concerning the dive depth.

Hettangian (Early Jurassic) Dinosaur Tracksites from the Mecsek Mountains, Hungary

Isolated theropod dinosaur tracks were first collected in Hungary from Hettangian (Lower Jurassic) beds of the Mecsek Coal Formation in 1966 and described as Komlosaurus carbonis Kordos, 1983. Our study is based on newly collected material from additional track-bearing beds. The description of the two largest preserved surfaces containing a total of 102 tracks that can be referred to as 21 trackways is provided here. This represents the first attempt to measure, map and compare the tracks of these bipedal, functionally tridactyl dinosaurs in several associated trackways. Significant morphological variability can be observed (e.g., depth, presence or absence of a metatarsal impression, digit length, digit divarication angle) that is explained by differences in physical parameters of the substrate. The mean of pes length is 16.3 cm in tracksite PB1 and 19.9 cm in tracksite PB2. Stride length of trackways usually ranges between 120 and 170 cm; pace angulation is 160–175°. The speed of the trackmaker is calculated to range between 6 and 14 km/h. Imprints are diagnosed by a pes length/width ratio lower than 2.0; metatarsal pads and hallux impressions are frequent. Based on the similarity of several morphological characters, the herein described tracks are referred to the ichnotaxon Komlosaurus carbonis, which is clearly distinct from Grallator and Kayentapus.

Upper Oligocene marine fishes from nearshore deposits of the Central Paratethys (Máriahalom, Hungary)

A rich and diverse ichthyofauna is described from the upper Oligocene (Egerian) sands of Máriahalom, Hungary. The site is dominated by brackish molluscs that are preserved together with rare marine and terrestrial vertebrates. Based on the isolated elasmobranch and bony fish remains, eight sharks, four rays and seven teleost taxa were identified from Máriahalom. The ichthyofauna represents a nearshore marine subtropical community dominated by odontaspidid and carcharhinid sharks and by euryhaline durophagous sparids and by sciaenid teleost fishes. Epibenthic feeders were common, whereas piscivorous taxa included barracudas, billfishes and numerous sharks, including the macropredatory Otodus angustidens. Palaeontological, sedimentological and stratigraphical data indicates a tide-influenced and fluvial-influenced nearshore palaeoenvironment with brackish lagoons and normal marine littoral habitats. The remains of terrestrial, brackish and nearshore marine invertebrates and vertebrates accumulated in a tidal channel. The systematics of the ichthyofauna is consistent with the palaeogeography inferred from invertebrates, i.e. a Central Paratethys connected to the Mediterranean area during the Egerian.