The last ceratosaur of Asia: a new noasaurid from the Early Cretaceous Great Siberian Refugium.

The noasaurid ceratosaur Kiyacursor longipes gen. et sp. nov. is described based on a fragmentary skeleton including cervical vertebra, pectoral girdle, humerus and hind limbs from the Lower Cretaceous (Aptian) Ilek Formation at Shestakovo 1 locality in Western Siberia, Russia. This is the first ceratosaur from the Early Cretaceous of Asia, extending the stratigraphic range of Ceratosauria by 40 Myr on that continent. Kiyacursor shares unique hind limb proportions with Elaphrosaurus and Limusaurus, suggesting improved cursorial ability. These taxa show an ostrich-like specialization of the pes, with a large third metatarsal and greatly reduced second metatarsal. By contrast, all other fast running non-avian theropod dinosaurs have an arctometatarsalian pes, with the third metatarsal strongly reduced proximally. The new taxon lived in the Early Cretaceous ecosystem containing a number of other Jurassic relics, such as stem salamanders, protosuchian and shartegosuchid crocodyliforms, tritylodontid synapsids and docodontan mammaliaforms.

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.

A derived dryolestid mammal indicates possible insular endemism in the Late Jurassic of Germany.

The Langenberg Quarry near Bad Harzburg has yielded the first Jurassic stem therian mammal of Germany, recovered from Kimmeridgian (Late Jurassic) near shore deposits of a palaeo-island within the Lower Saxony Basin of the European archipelago. The new stem therian is represented by one lower and three upper molars. Hercynodon germanicus gen. et sp. nov. is attributed to the Dryolestidae, a group of pretribosphenic crown mammals that was common in western Laurasia from the Middle Jurassic to the Early Cretaceous. The new taxon is characterised by small size, a reduced cusp pattern in the upper molars lacking a metacone, and enhancement of the shearing crests paracrista and metacrista. Phylogenetic analysis identified Hercynodon gen. nov. as sister taxon of Crusafontia from the Lower Cretaceous (Barremian) of Spain. Both taxa belong to an endemic European clade of dryolestids, including also Achyrodon and Phascolestes from the earliest Cretaceous (Berriasian) of England. Despite its greater geological age, Hercynodon gen. nov. is the most derived representative of that clade, indicated by the complete reduction of the metacone. The discrepancy between derived morphology and geological age may be explained by an increased rate of character evolution in insular isolation. Other insular phenomena have earlier been observed in vertebrates from the Langenberg Quarry, such as dwarfism in the small sauropod Europasaurus, and possible gigantism in the morganucodontan mammaliaform Storchodon and the pinheirodontid multituberculate mammal Teutonodon which grew unusually large.

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.

First rebbachisaurid sauropod dinosaur from Asia.

Dzharatitanis kingi gen. et sp. nov. is based on an isolated anterior caudal vertebra (USNM 538127) from the Upper Cretaceous (Turonian) Bissekty Formation at Dzharakuduk, Uzbekistan. Phylogenetic analysis places the new taxon within the diplodocoid clade Rebbachisauridae. This is the first rebbachisaurid reported from Asia and one of the youngest rebbachisaurids in the known fossil record. The caudal is characterized by a slightly opisthocoelous centrum, 'wing-like' transverse processes with large but shallow PRCDF and POCDF, and the absence of a hyposphenal ridge and of TPRL and TPOL. The neural spine has high SPRL, SPDL, SPOL, and POSL and is pneumatized. The apex of neural spine is transversely expanded and bears triangular lateral processes. The new taxon shares with Demandasaurus and the Wessex rebbachisaurid a high SPDL on the lateral side of the neural spine, separated from SPRL and SPOL. This possibly suggests derivation of Dzharatitanis from European rebbachisaurids. This is the second sauropod group identified in the assemblage of non-avian dinosaurs from the Bissekty Formation, in addition to a previously identified indeterminate titanosaurian.

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.

The oldest record of Alvarezsauridae (Dinosauria: Theropoda) in the Northern Hemisphere.

Procoelous caudal vertebrae, a carpometacarpus with a hypertrophied metacarpal II, and robust proximal and ungual phalanges of manual digit II of a small theropod dinosaur from the Upper Cretaceous (Turonian) Bissekty Formation at Dzharakuduk, Uzbekistan, show unequivocal synapomorphies of the clade Alvarezsauridae and thus are referred to it. The caudal vertebrae have a unique longitudinal canal within the neural arch. The carpometacarpus, with metacarpal III occupying about one third of the width of the carpometacarpus, shows the most plesiomorphic stage of the evolution of the forelimb among known alvarezsaurids. The proximal phalanx of manual digit II differs from the corresponding bone in Parvicursorinae in having a less asymmetrical proximal articular surface without a dorsal process and short ventral ridges. The ungual phalanx of manual digit II has laterally open ventral foramina. The Bissekty alvarezsaurid possibly represents a basal parvicursorine and is the stratigraphically oldest known alvarezsaurid in Asia known to date.

Origin of host-parasite associations of Marsupialges misonnei (Acariformes: Psoroptidae)-a parasitological detective story.

Host associations of permanent ectoparasitic mite Marsupialges misonnei Fain, 1963 (Acariformes: Psoroptidae: Marsupialginae) are analyzed. This species was first recorded from an ethanol-preserved museum specimen of Caluromys philander (Linnaeus, 1758) (Didelphimorphia: Didelphidae) originating from French Guiana. We discovered specimens of M. misonnei from both species known in the carnivore genus Nasua (Carnivora: Procyonidae): N. narica (Linnaeus, 1766) from Panama (collected in the field) and N. nasua (Linnaeus, 1766) from Brazil (collected from dry museum specimen). Two alternative hypotheses about an initial host of this mite (bare-tailed woody opossum or coatis) are discussed. We argue that M. misonnei was originally parasitic on Nasua spp. and occasionally contaminated C. philander from these hosts in the collecting process.

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.

New tyrannosaur from the mid-Cretaceous of Uzbekistan clarifies evolution of giant body sizes and advanced senses in tyrant dinosaurs.

Tyrannosaurids--the familiar group of carnivorous dinosaurs including Tyrannosaurus and Albertosaurus--were the apex predators in continental ecosystems in Asia and North America during the latest Cretaceous (ca. 80-66 million years ago). Their colossal sizes and keen senses are considered key to their evolutionary and ecological success, but little is known about how these features developed as tyrannosaurids evolved from smaller basal tyrannosauroids that first appeared in the fossil record in the Middle Jurassic (ca. 170 million years ago). This is largely because of a frustrating 20+ million-year gap in the mid-Cretaceous fossil record, when tyrannosauroids transitioned from small-bodied hunters to gigantic apex predators but from which no diagnostic specimens are known. We describe the first distinct tyrannosauroid species from this gap, based on a highly derived braincase and a variety of other skeletal elements from the Turonian (ca. 90-92 million years ago) of Uzbekistan. This taxon is phylogenetically intermediate between the oldest basal tyrannosauroids and the latest Cretaceous forms. It had yet to develop the giant size and extensive cranial pneumaticity of T. rex and kin but does possess the highly derived brain and inner ear characteristic of the latest Cretaceous species. Tyrannosauroids apparently developed huge size rapidly during the latest Cretaceous, and their success in the top predator role may have been enabled by their brain and keen senses that first evolved at smaller body size.

Evolutionary transition of dental formula in Late Cretaceous eutherian mammals.

Kulbeckia kulbecke, stem placental mammal from the Late Cretaceous of Uzbekistan, shows a transitional stage of evolution in the dental formula from five to four premolars. A non-replaced dP3/dp3 may occur as individual variation. In other specimens, the lower premolars are crowded with no space for development of dp3. As is evident from the CT scanning of one juvenile specimen, the development of dp3 started in a late ontogenetic stage and was confined to the pulp cavity of the developing p2. This dp3 would have been resorbed in a later ontogenetic stage, as the roots of p2 formed. The initial stage of reduction of the third premolar can be traced to stem therians (Juramaia and Eomaia), which have both dP3 and P3 present in the adult dentition. Further delay in the development of dP3/dp3 led to the loss of the permanent P3/p3 (a possible synapomorphy for Eutheria). The dP3/dp3 was present during most of the adult stages in the Late Cretaceous stem placentals Zhelestidae and Gypsonictops. This tooth is totally absent in basal taxa of Placentalia, which normally have at most four premolars.

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.

Review of taxonomy, geographic distribution, and paleoenvironments of Azhdarchidae (Pterosauria).

The taxonomy, geographic distribution, and paleoenvironmental context of azhdarchid pterosaurs are reviewed. All purported pteranodontid, tapejarid, and azhdarchid specimens from the Cenomanian Kem Kem beds of Morocco are referred to a single azhdarchid taxon, Alanqa saharica. The four proposed autapomorphies of Eurazhdarcho langendorfensis from the lower Maastrichtian Sebes Formation of Romania are based on misinterpretations of material and this taxon is likely a subjective junior synonym of Hatzegopteryx thambema. Among 54 currently reported azhdarchid occurrences (51 skeletal remains and 3 tracks) 13% are from lacustrine deposits, 17% from fluvial plain deposits, 17% from coastal plain deposits, 18% from estuarine and lagoonal deposits, and 35% from costal marine deposits. Azhdarchids likely inhabited a variety of environments, but were abundant near large lakes and rivers and most common in nearshore marine paleoenvironments.

A new phylogeny for basal Trechnotheria and Cladotheria and affinities of South American endemic Late Cretaceous mammals.

The endemic South American mammals Meridiolestida, considered previously as dryolestoid cladotherians, are found to be non-cladotherian trechnotherians related to spalacotheriid symmetrodontans based on a parsimony analysis of 137 morphological characters among 44 taxa. Spalacotheriidae is the sister taxon to Meridiolestida, and the latter clade is derived from a primitive spalacolestine that migrated to South America from North America at the beginning of the Late Cretaceous. Meridiolestida survived until the early Paleocene (Peligrotherium) and early Miocene (Necrolestes) in South America, and their extinction is probably linked to the increasing competition with metatherian and eutherian tribosphenic mammals. The clade Meridiolestida plus Spalacotheriidae is the sister taxon to Cladotheria and forms a new clade Alethinotheria. Alethinotheria and its sister taxon Zhangheotheria, new clade (Zhangheotheriidae plus basal taxa), comprise Trechnotheria. Cladotheria is divided into Zatheria (plus stem taxa, including Amphitherium) and Dryolestida, including Dryolestidae and a paraphyletic array of basal dryolestidans (formerly classified as "Paurodontidae"). The South American Vincelestes and Groebertherium are basal dryolestidans.

A new basal hadrosauroid dinosaur from the Late Cretaceous of Uzbekistan and the early radiation of duck-billed dinosaurs.

Levnesovia transoxiana gen. et sp. nov., from the Late Cretaceous (Middle-Late Turonian) of Uzbekistan, is the oldest well-documented taxon referable to Hadrosauroidea sensu Godefroit et al. It differs from a somewhat younger and closely related Bactrosaurus from Inner Mongolia (China) by a tall sagittal crest on the parietals and the absence of club-shaped dorsal neural spines in adult specimens. Levnesovia, Bactrosaurus and possibly Gilmoreosaurus represent the earliest radiation of Hadrosauroidea, which took place during the Cenomanian-Turonian and possibly in North America. The second, Santonian-age radiation of Hadrosauroidea included Aralosaurus, Hadrosauridae and lineages leading to Tanius (Campanian) and Telmatosaurus (Maastrichtian). Hadrosauridae appears to be monophyletic, but Hadrosaurinae and Lambeosaurinae originated in North America and Asia, respectively.

Turanoceratops tardabilis--the first ceratopsid dinosaur from Asia.

Turanoceratops tardabilis from the Upper Cretaceous (Turonian) Bissekty Formation of Dzharakuduk, Uzbekistan, represents the first definite ceratopsid dinosaur recorded from Asia. Reexamination of the original and study of newly collected material indicate that Turanoceratops has well-developed supraorbital horns and apparently lacks a nasal horn. Turanoceratops is more derived than the more or less coeval Zuniceratops from the Moreno Hill Formation of New Mexico in the presence of double-rooted teeth and of two or three teeth in each vertical dental file.

A previously unrecognized group of Middle Jurassic triconodontan mammals from Central Asia.

Ferganodon narynensis gen. et sp. nov. is represented by a lower molariform tooth from the Middle Jurassic (Callovian) Balabansai Svita in Kyrgyzstan. The new genus is allied with Klamelia zhaopengi Chow and Rich 1984 from the Middle Jurassic Shishugou Formation in Xinjiang, northwest China to the new family Klameliidae based on parallelogram-shaped lower molariforms, imbricating rather than interlocking of cusps e-d-f, by a peculiar distolabial cingulid cusp, and by vertical folding of the enamel on the labial crown side. The new family Klameliidae fam. nov. is most similar to Gobiconodontidae by the structure of the molariform teeth and represents a previously unrecognized radiation of eutriconodontan mammals possibly endemic to Central Asia.

An aegialodontid upper molar and the evolution of mammal dentition.

The most obvious key synapomorphy of the therian mammals is the tribosphenic pattern of their molars. Tribosphenic teeth are capable of both shearing and grinding, which substantially increase effectiveness of food processing and, in turn, permit evolution of a wide range of dietary specializations. Functional tribospheny developed repeatedly during mammalian evolution but was successful only in the Boreosphenida. The earliest stage in the development of boreosphenidan tribospheny has remained poorly understood, being documented only by lower molars of aegialodontids. Here, we report a known upper molar of an aegialodontid mammal, Kielantherium, from the Early Cretaceous of Mongolia.

First jurassic choristodera from Asia.

Although choristoderes have a good Lower Cretaceous record in Asia, they have never previously been recorded from Jurassic deposits. Here we describe fragmentary vertebral material referable to Choristodera indet. from the Middle Jurassic Balabansai Svita of the Fergana Valley, Kyrgyzstan. This provides a significant range extension for the group in Asia and shows that choristoderes already had a Pan-Laurasian distribution in the Jurassic.