On the way from Asia to America: eutriconodontan mammals from the Early Cretaceous of Yakutia, Russia.

Eutriconodonta are an important group of early crown mammals with a wide distribution in the Jurassic-Cretaceous of the Northern Hemisphere and few occurrences in the Southern Hemisphere. Three taxa of eutriconodontans are known from the Early Cretaceous high-latitude Teete vertebrate assemblage in Yakutia, Russia: Sangarotherium aquilonium (Eutriconodonta incertae sedis), Gobiconodon sp. A (large), and Gobiconodon sp. B (small) (Gobiconodontidae). These three taxa are based on four specimens and indicate a remarkable taxonomic diversity of eutriconodontans at this locality. The coexistence of two Gobiconodon species, large and small, is characteristic for several Early Cretaceous vertebrate assemblages in Asia. Gobiconodon sp. A from the Teete locality is the largest species of this genus known from Asia, but is smaller than the North American G. ostromi. The spreading of Gobiconodon from Asia to North America likely occurred during the Aptian-Albian faunal dispersal event. The discovery of Gobiconodon in the Teete locality is further evidence for a dispersal route via Beringia from Asia to North America which previously has been postulated based on the occurrence of Asian dinosaur taxa in western North America at this time. The questionable record of Gobiconodon from Europe and its lack from eastern North America make a dispersal from Asia to North America via Europe less probable.

Middle Jurassic fossils document an early stage in salamander evolution.

Salamanders are an important group of living amphibians and model organisms for understanding locomotion, development, regeneration, feeding, and toxicity in tetrapods. However, their origin and early radiation remain poorly understood, with early fossil stem-salamanders so far represented by larval or incompletely known taxa. This poor record also limits understanding of the origin of Lissamphibia (i.e., frogs, salamanders, and caecilians). We report fossils from the Middle Jurassic of Scotland representing almost the entire skeleton of the enigmatic stem-salamander Marmorerpeton. We use computed tomography to visualize high-resolution three-dimensional anatomy, describing morphologies that were poorly characterized in early salamanders, including the braincase, scapulocoracoid, and lower jaw. We use these data in the context of a phylogenetic analysis intended to resolve the relationships of early and stem-salamanders, including representation of important outgroups alongside data from high-resolution imaging of extant species. Marmorerpeton is united with Karaurus, Kokartus, and others from the Middle Jurassic-Lower Cretaceous of Asia, providing evidence for an early radiation of robustly built neotenous stem-salamanders. These taxa display morphological specializations similar to the extant cryptobranchid "giant" salamanders. Our analysis also demonstrates stem-group affinities for a larger sample of Jurassic species than previously recognized, highlighting an unappreciated diversity of stem-salamanders and cautioning against the use of single species (e.g., Karaurus) as exemplars for stem-salamander anatomy. These phylogenetic findings, combined with knowledge of the near-complete skeletal anatomy of Mamorerpeton, advance our understanding of evolutionary changes on the salamander stem-lineage and provide important data on early salamanders and the origins of Batrachia and Lissamphibia.

Braincase anatomy of extant Crocodylia, with new insights into the development and evolution of the neurocranium in crocodylomorphs.

Present-day crocodylians exhibit a remarkably akinetic skull with a highly modified braincase. We present a comprehensive description of the neurocranial osteology of extant crocodylians, with notes on the development of individual skeletal elements and a discussion of the terminology used for this project. The quadrate is rigidly fixed by multiple contacts with most braincase elements. The parabasisphenoid is sutured to the pterygoids (palate) and the quadrate (suspensorium); as a result, the basipterygoid joint is completely immobilized. The prootic is reduced and externally concealed by the quadrate. It has a verticalized buttress that participates in the canal for the temporal vasculature. The ventrolateral processes of the otoccipitals completely cover the posteroventral region of the braincase, enclose the occipital nerves and blood vessels in narrow bony canals and also provide additional sutural contacts between the braincase elements and further consolidate the posterior portion of the crocodylian skull. The otic capsule of crocodylians has a characteristic cochlear prominence that corresponds to the lateral route of the perilymphatic sac. Complex internal structures of the otoccipital (extracapsular buttress) additionally arrange the neurovascular structures of the periotic space of the cranium. Most of the braincase elements of crocodylians are excavated by the paratympanic pneumatic sinuses. The braincase in various extant crocodylians has an overall similar structure with some consistent variation between taxa. Several newly observed features of the braincase are present in Gavialis gangeticus and extant members of Crocodylidae to the exclusion of alligatorids: the reduced exposure of the prootic buttress on the floor of the temporal canal, the sagittal nuchal crest of the supraoccipital projecting posteriorly beyond the postoccipital processes and the reduced paratympanic pneumaticity. The most distinctive features of the crocodylian braincase (fixed quadrate and basipterygoid joint, consolidated occiput) evolved relatively rapidly at the base of Crocodylomorpha and accompanied the initial diversification of this clade during the Late Triassic and Early Jurassic. We hypothesize that profound rearrangements in the individual development of the braincases of basal crocodylomorphs underlie these rapid evolutionary modifications. These rearrangements are likely reflected in the embryonic development of extant crocodylians and include the involvement of neomorphic dermal anlagen in different portions of the developing chondrocranium, the extensive ossification of the palatoquadrate cartilage as a single expanded quadrate and the anteromedial inclination of the quadrate.

Wear patterns and dental functioning in an Early Cretaceous stegosaur from Yakutia, Eastern Russia.

Isolated stegosaurian teeth from the Early Cretaceous high-latitude (palaeolatitude estimate of N 62°- 66.5°) Teete locality in Yakutia (Eastern Siberia, Russia) are characterized by a labiolingually compressed, slightly asymmetrical and mesiodistally denticulated (9-14 denticles) crown, a pronounced ring-like cingulum, as well as a "complex network of secondary ridges". The 63 teeth (found during on-site excavation in 2012, 2017-2019 and screen-washing in 2017-2019) most likely belong to one species of a derived (stegosaurine) stegosaur. Most of the teeth exhibit a high degree of wear and up to three wear facets has been observed on a single tooth. The prevalence of worn teeth with up to three wear facets and the presence of different types of facets (including steeply inclined and groove-like) indicate the tooth-tooth contact and precise dental occlusion in the Teete stegosaur. The microwear pattern (mesiodistally or slightly obliquely oriented scratches; differently oriented straight and curved scratches on some wear facets) suggest a complex jaw mechanism with palinal jaw motion. Histological analysis revealed that the Teete stegosaur is characterized by relatively short tooth formation time (95 days) and the presence of a "wavy enamel pattern". Discoveries of a "wavy enamel pattern" in the Teete stegosaur, in a Middle Jurassic stegosaur from Western Siberia, and in the basal ceratopsian Psittacosaurus, suggest that this histological feature is common for different ornithischian clades, including ornithopods, marginocephalians, and thyreophorans. A juvenile tooth in the Teete sample indicates that stegosaurs were year-round residents and reproduced in high latitudes. The combination of high degree of tooth wear with formation of multiple wear facets, complex jaw motions, relatively short tooth formation time and possibly high tooth replacement rates is interpreted as a special adaptation for a life in high-latitude conditions or, alternatively, as a common stegosaurian adaptation making stegosaurs a successful group of herbivorous dinosaurs in the Middle Jurassic-Early Cretaceous and enabeling them to live in both low- and high-latitude ecosystems.

A new small-sized stem salamander from the Middle Jurassic of Western Siberia, Russia.

Salamanders (Caudata) are one of the three modern groups of amphibians known from the Middle Jurassic. The early stages of evolution of these amphibians are still poorly known, especially for stem taxa of Jurassic age. A new small-sized stem salamander, Egoria malashichevi gen. et sp. nov., from the Middle Jurassic (Bathonian) Itat Formation of the Berezovsk Quarry locality in Western Siberia, Russia, is described on the basis of isolated vertebrae, including an atlas centrum and a fragmentary trunk vertebra centrum previously referred to an undescribed salamander taxon ("Berezovsk salamander A"). The new taxon is diagnosed by the following unique combination of vertebral characters: atlantal anterior cotyles with elliptical anterior outline, located at an angle of approximately 135-137 degrees to each other; wide posterior portion of the atlantal centrum; ossified portion of the intercotylar tubercle represented by dorsal and ventral lips; absence of a deep depression on the ventral surface of the atlantal centrum; absence of pronounced ventrolateral ridges on the atlas; absence of spinal nerve foramina; presence of a pitted texture on the ventral and lateral surfaces of the centra and lateral surfaces neural arch pedicels; presence of a short atlantal neural arch with its anterior border situated behind the level of the anterior cotyles; short trunk vertebrae; and upper transverse process (= diapophysis) larger than lower transverse process (= parapophysis) on the trunk vertebrae; notochordal canal opens in the upper half of the cotyle (= the lower portion of the centrum is more massive and less compact than the upper portion). The microanatomical organization of the atlas and trunk vertebrae is characterized by the presence of inner cancellous endochondral bone. The small body size (about 180-215 mm) of Egoria malashichevi gen. et sp. nov. indicates that that not all stem salamanders were large neotenic forms (up to 550-600 mm in Urupia and Marmorerpeton) and hints at a broader ecological role for stem salamanders.

A high-latitude fauna of mid-Mesozoic mammals from Yakutia, Russia.

The Early Cretaceous (?Berriasian-Barremian) Teete vertebrate locality in Western Yakutia, East Siberia, Russia, has produced mammal remains that are attributed to three taxa: Eleutherodontidae indet. cf. Sineleutherus sp. (Haramiyida; an upper molariform tooth), Khorotherium yakutensis gen. et sp. nov. (Tegotheriidae, Docodonta; maxillary fragment with three molariform teeth and dentary fragment with one molariform tooth), and Sangarotherium aquilonium gen. et sp. nov. (Eutriconodonta incertae sedis; dentary fragment with one erupted molariform tooth and one tooth in crypt). This is the second occurrence of Mesozoic mammals in high latitudes (paleolatitude estimate N 63-70°) of the Northern Hemisphere. In spite of the presumed Early Cretaceous age based on freshwater mollusks, the Teete mammal assemblage has a distinctive Jurassic appearance, being most similar to the Middle-Late Jurassic mammal assemblages known from Siberia, Russia and Xinjiang, China. The smooth transition from Jurassic to Cretaceous biota in Northern Asia is best explained by stable environmental conditions.

Cranial biomechanics in basal urodeles: the Siberian salamander (Salamandrella keyserlingii) and its evolutionary and developmental implications.

Developmental changes in salamander skulls, before and after metamorphosis, affect the feeding capabilities of these animals. How changes in cranial morphology and tissue properties affect the function of the skull are key to decipher the early evolutionary history of the crown-group of salamanders. Here, 3D cranial biomechanics of the adult Salamandrella keyserlingii were analyzed under different tissue properties and ossification sequences of the cranial skeleton. This helped unravel that: (a) Mechanical properties of tissues (as bone, cartilage or connective tissue) imply a consensus between the stiffness required to perform a function versus the fixation (and displacement) required with the surrounding skeletal elements. (b) Changes on the ossification pattern, producing fontanelles as a result of bone loss or failure to ossify, represent a trend toward simplification potentially helping to distribute stress through the skull, but may also imply a major destabilization of the skull. (c) Bone loss may be originated due to biomechanical optimization and potential reduction of developmental costs. (d) Hynobiids are excellent models for biomechanical reconstruction of extinct early urodeles.

First nimravid skull from Asia.

Maofelis cantonensis gen. and sp. nov. is described based on a complete cranium from the middle-upper Eocene Youganwo Formation of Maoming Basin, Guangdong Province, China. The new taxon has characters diagnostic for Nimravidae such as a short cat-like skull, short palate, ventral surface of petrosal dorsal to that of basioccipital, serrations on the distal carina of canine, reduced anterior premolars, and absence of posterior molars (M2-3). It is plesiomorphic nimravid taxon similar to Nimravidae indet. from Quercy (France) in having the glenoid pedicle and mastoid process without ventral projections, a planar basicranium in which the lateral rim is not ventrally buttressed, and P1 present. The upper canine is less flattened than in other Nimravidae. Maofelis cantonensis gen. and sp. nov. exemplifies the earliest stage of development of sabertooth specialization characteristic of Nimravidae. This taxon, together with other middle-late Eocene nimravid records in South Asia, suggests origin and initial diversification of Nimravidae in Asia. We propose that this group dispersed to North America in the late Eocene and to Europe in the early Oligocene. The subsequent Oligocene diversification of Nimravidae took place in North America and Europe, while in Asia this group declined in the Oligocene, likely because of the earlier development of open habitats on that continent.

The paleoenvironments of azhdarchid pterosaurs localities in the Late Cretaceous of Kazakhstan.

Five pterosaur localities are currently known from the Late Cretaceous in the northeastern Aral Sea region of Kazakhstan. Of these, one is Turonian-Coniacian in age, the Zhirkindek Formation (Tyulkili), and four are Santonian in age, all from the early Campanian Bostobe Formation (Baibishe, Akkurgan, Buroinak, and Shakh Shakh). All so far collected and identifiable Late Cretaceous pterosaur bones from Kazakhstan likely belong to Azhdarchidae: Azhdarcho sp. (Tyulkili); Aralazhdarchobostobensis (Shakh Shakh); and Samrukianessovi (Akkurgan). These latter two taxa, both from the Bostobe Formation might be synonyms. Azhdarcho sp. from the Zhirkindek Formation lived in a tropical-to-subtropical relatively humid climate on the shore of an estuarine basin connected to the Turgai Sea. Known fossils were collected in association with brackish-water bivalves and so the overall paleoenvironment of this pterosaur was likely an estuarine marsh as indicated by the dominance of conifers and low relative counts of ferns and angiosperms. Aralazhdarchobostobensis, from the Bostobe Formation, lived on a coastal fluvial plain along the Turgai Sea. This paleoenvironment was either floodplain (Akkurgan, Buroinak, and Shakh Shakh) or estuarine (Baibishe). In the Santonian - early Campanian, shallow waters near this coastal plain were sites for the intensive accumulation of phosphates under upwelling conditions caused by strong winds from the ancient Asian landmass. These winds also caused significant aridization of the climate during this time. We speculate that pterosaurs may have been attracted to this area by the abundant resources in the bio-productive estuaries and nearshore upwelling waters.

Long bone histology of the stem salamander Kokartus honorarius (Amphibia: Caudata) from the Middle Jurassic of Kyrgyzstan.

Kokartus honorarius from the Middle Jurassic (Bathonian) of Kyrgyzstan is one of the oldest salamanders in the fossil record, characterized by a mixture of plesiomorphic morphological features and characters shared with crown-group salamanders. Here we present a detailed histological analysis of its long bones. The analysis of a growth series demonstrates a significant histological maturation during ontogeny, expressed by the progressive appearance of longitudinally oriented primary vascular canals, primary osteons, growth marks, remodelling features in primary bone tissues, as well as progressive resorption of the calcified cartilage, formation of endochondral bone and development of cartilaginous to bony trabeculae in the epiphyses. Apart from the presence of secondary osteons, the long bone histology of Kokartus is very similar to that of miniaturized temnospondyls, other Jurassic stem salamanders, miniaturized seymouriamorphs and modern crown-group salamanders. We propose that the presence of secondary osteons in Kokartus honorarius is a plesiomorphic feature, and the loss of secondary osteons in the long bones of crown-group salamanders as well as in those of miniaturized temnospondyls is the result of miniaturization processes. Hitherto, all stem salamander long bong histology (Kokartus, Marmorerpeton and 'salamander A') has been generally described as having paedomorphic features (i.e. the presence of Katschenko's Line and a layer of calcified cartilage), these taxa were thus most likely neotenic forms. The absence of clear lines of arrested growth and annuli in long bones of Kokartus honorarius suggests that the animals lived in an environment with stable local conditions.

The spinal cord supports of vertebrae in the crown-group salamanders (Caudata, Urodela).

The development of spinal cord supports (bony thickenings which extend into the vertebral canal of vertebrae) in primitive (Salamandrella keyserlingii) and derived (Lissotriton vulgaris) salamanders were described. The spinal cord supports develop as the protuberances of periostal bone of the neural arches in the anteroproximal part of the septal collagenous fibers which connect a transverse myoseptum with the notochord and spinal cord, in the septal bundle inside the vertebral canal. Spinal cord supports were also found in some teleostean (Salmo salar, Oncorhynchus mykiss) and dipnoan (Protopterus sp.) fishes. The absence of the spinal cord supports in vertebrates with cartilaginous vertebrae (lampreys, chondrichthyan, and chondrostean fishes) corresponds to the fact that the spinal cord supports are bone structures. The absence of the spinal cord supports in frogs correlates with the lack of the well developed septal bundles inside the vertebral canal. The spinal cord supports are, presumably, a synapomorphic character for salamanders which originated independently of those observed in teleostean and dipnoan fishes.