A new alvarezsaurid dinosaur (Theropoda, Alvarezsauria) from the Upper Cretaceous Baruungoyot Formation of Mongolia provides insights for bird-like sleeping behavior in non-avian dinosaurs.

Alvarezsauria is a group of early-branching maniraptoran theropods that are distributed globally from the Late Jurassic to the latest Cretaceous. Despite recent increases in the fossil record of this group, the scarcity of complete specimens still restricts interpreting their detailed anatomy, ecology, and evolution. Here, we report a new taxon of derived alvarezsaur, Jaculinykus yaruui gen. et sp. nov., from the Late Cretaceous of Mongolia, which represents a nearly complete and articulated skeleton. Our phylogenetic analysis reveals that Jaculinykus belongs to the sub-clade of Alvarezsauridae, Parvicursorinae, and forms a mononphyletic group with Mononykus and Shuvuuia. Its well-preserved manus has only two fingers, composed of a hypertrophied digit I and greatly reduced digit II, which implies an intermediate condition between the tridactyl manus of Shuvuuia and monodactyl manus of Linhenykus. This highlights a previously unrecognized variation in specialization of alvarezsaurid manus. Notably, the preserved posture of the specimen exhibits a stereotypical avian-like sleeping position seen in the troodontids Mei and Sinornithoides. Evidence of this behavior in the alvarezsaur Jaculinykus suggests that stereotypically avian sleeping postures are a maniraptoran synapomorphy, providing more evidence of bird-like traits being distributed broadly among avian ancestors.

Chronic fracture and osteomyelitis in a large-bodied ornithomimosaur with implications for the identification of unusual endosteal bone in the fossil record.

Paleopathological diagnoses provide key information on the macroevolutionary origin of disease as well as behavioral and physiological inferences that are inaccessible via direct observation of extinct organisms. Here we describe the external gross morphology and internal architecture of a pathologic right second metatarsal (MMNS VP-6332) of a large-bodied ornithomimid (~432 kg) from the Santonian (Upper Cretaceous) Eutaw Formation in Mississippi, using a combination of X-ray computed microtomography (microCT) and petrographic histological analyses. X-ray microCT imaging and histopathologic features are consistent with multiple complete, oblique to comminuted, minimally displaced mid-diaphyseal cortical fractures that produce a "butterfly" fragment fracture pattern, and secondary osteomyelitis with a bone fistula formation. We interpret this as evidence of blunt force trauma to the foot that could have resulted from intra- or interspecific competition or predator-prey interaction, and probably impaired the function of the metatarsal as a weight-bearing element until the animal's death. Of particular interest is the apparent decoupling of endosteal and periosteal pathological bone deposition in MMNS VP-6332, which produces transverse sections exhibiting homogenously thick endosteal pathological bone in the absence of localized periosteal reactive bone. These distribution and depositional patterns are used as criteria for ruling out a pathological origin in favor of a reproductive one for unusual endosteal bone in fossil specimens. On the basis of MMNS VP-6332, we suggest caution in their use to substantiate a medullary bone identification in extinct archosaurians.

New therizinosaurid dinosaur from the marine Osoushinai Formation (Upper Cretaceous, Japan) provides insight for function and evolution of therizinosaur claws.

The record of therizinosaurs is rich in Asian countries such as Mongolia and China. Fragmentary therizinosaur specimens have been reported from the Lower and Upper Cretaceous deposits in Japan. One of these specimens, from the lower Campanian Osoushinai Formation in Nakagawa Town of Hokkaido Prefecture, was previously identified as a maniraptoran theropod dinosaur, possibly therizinosaur, but its taxonomic status remained unresolved. This study re-examines the specimen and provides a more detailed description and attempts to resolve its taxonomic status. Our study demonstrates that it is a new taxon, Paralitherizinosaurus japonicus gen. et sp. nov., because it shows a unique combination of characters in the metacarpal I and unguals. Our phylogenetic analysis places this new taxon within an unresolved clade of Therizinosauridae in the strict consensus tree. The 50% majority-rule consensus tree shows better resolution within Therizinosauridae, showing an unresolved monophyletic clade of Paralitherizinosaurus, Therizinosaurus, Suzhousaurus, and the Bissekty form. Geometric morphometric analysis suggests that Paralitherizinosaurus unguals most closely resemble Therizinosaurus unguals in being slender and has weak flexor tubercles. This study also shows an evolutionary trend in ungual shape, which associates a decrease in mechanical advantage, development of flexor tubercle, and hypothesized output (product of mechanical advantage and development of flexor tubercle) in derived therizinosaurs, supporting the hook-and-pull function of claws to bring vegetation to its mouth. Paralitherizinosaurus is the youngest therizinosaur from Japan and the first recovered from the marine deposits in Asia. This suggests a long temporal existence of therizinosaurs at the eastern edge of the Asian continent and adaptation of therizinosaurs to coastal environments.

A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria.

Late Cretaceous trends in Asian dinosaur diversity are poorly understood, but recent discoveries have documented a radiation of oviraptorosaur theropods in China and Mongolia. However, little work has addressed the factors that facilitated this diversification. A new oviraptorid from the Late Cretaceous of Mongolia sheds light on the evolution of the forelimb, which appears to have played a role in the radiation of oviraptorosaurs. Surprisingly, the reduced arm has only two functional digits, highlighting a previously unrecognized occurrence of digit loss in theropods. Phylogenetic analysis shows that the onset of this reduction coincides with the radiation of heyuannine oviraptorids, following dispersal from southern China into the Gobi region. This suggests expansion into a new niche in the Gobi region, which relied less on the elongate, grasping forelimbs inherited by oviraptorosaurs. Variation in forelimb length and manus morphology provides another example of niche partitioning in oviraptorosaurs, which may have made possible their incredible diversity in the latest Cretaceous of Asia.

A neoceratopsian dinosaur from the early Cretaceous of Mongolia and the early evolution of ceratopsia.

Ceratopsia is a diverse dinosaur clade from the Middle Jurassic to Late Cretaceous with early diversification in East Asia. However, the phylogeny of basal ceratopsians remains unclear. Here we report a new basal neoceratopsian dinosaur Beg tse based on a partial skull from Baruunbayan, Ömnögovi aimag, Mongolia. Beg is diagnosed by a unique combination of primitive and derived characters including a primitively deep premaxilla with four premaxillary teeth, a trapezoidal antorbital fossa with a poorly delineated anterior margin, very short dentary with an expanded and shallow groove on lateral surface, the derived presence of a robust jugal having a foramen on its anteromedial surface, and five equally spaced tubercles on the lateral ridge of the surangular. This is to our knowledge the earliest known occurrence of basal neoceratopsian in Mongolia, where this group was previously only known from Late Cretaceous strata. Phylogenetic analysis indicates that it is sister to all other neoceratopsian dinosaurs.

Author Correction: First Ornithomimid (Theropoda, Ornithomimosauria) from the Upper Cretaceous Djadokhta Formation of Tögrögiin Shiree, Mongolia.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Author Correction: A New Hadrosaurine (Dinosauria: Hadrosauridae) from the Marine Deposits of the Late Cretaceous Hakobuchi Formation, Yezo Group, Japan.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

A New Hadrosaurine (Dinosauria: Hadrosauridae) from the Marine Deposits of the Late Cretaceous Hakobuchi Formation, Yezo Group, Japan.

A nearly complete skeleton of a new hadrosaurid, Kamuysaurus japonicus gen. et sp. nov., was discovered from the outer shelf deposits of the Upper Cretaceous Hakobuchi Formation of the Yezo Group in Hobetsu area of Mukawa town in Hokkaido, Japan. Kamuysaurus belongs to the sub-clade of Hadrosaurinae, Edmontosaurini, and forms a monophyly with Laiyangosaurus and Kerberosaurus from the northern Far East. Kamuysaurus has a long anterior platform for the nasofrontal sutural surface, which may indicate the presence of a small supracranial crest, similar to a sub-adult form of Brachylophosaurus based on the extension of the nasofrontal sutural surface. The Dispersal Extinction Cladogenesis analysis with the 50% Majority Rule consensus tree suggests that the clade of Kamuysaurus, Laiyangosaurus, and Kerberosaurus may have dispersed into Asia prior to the late Campanian and the potential endemism of this clade during the late Campanian and early Maastrichtian in the northern Far East. The results of both Dispersal Extinction Cladogenesis and Ancestral State Reconstruction analyses imply that the marine-influenced environment in North America during the Campanian may have played an important role for the hadrosaurid diversification in its early evolutionary history.

A new iguanodontian (Dinosauria: Ornithopoda) from the Early Cretaceous of Mongolia.

We describe a new iguanodontian ornithopod, Choyrodon barsboldi gen. et sp. nov. from the Albian-aged Khuren Dukh Formation of Mongolia based on several partial skeletons interpreted to represent a subadult growth stage based on osteohistological features. This new taxon is diagnosed by many autapomorphies of the maxilla, nasal, lacrimal, opisthotic, predentary, and surangular. Choyrodon displays an unusual combination of traits, possessing an open antorbital fenestra (a primitive ornithopod trait) together with derived features such as a downturned dentary and enlarged narial fenestra. Histological imaging suggests that the type specimen of Choyrodon would have been a subadult at the time of death. Phylogenetic analysis of two different character matrices do not posit Choyrodon to be the sister taxon or to be more primitive than the iguanodontian Altirhinus kurzanovi, which is found in the same formation. The only resolved relationship of this new taxon is that it was hypothesized to be a sister-taxon with the North American species Eolambia caroljonesa. Though discovered in the same formation and Choyrodon being smaller-bodied than Altirhinus, it does not appear that the former species is an ontogimorph of the latter. Differences in morphology and results of the phylogenetic analyses support their distinction although more specimens of both species will allow better refinement of their uniqueness.

Author Correction: First Ornithomimid (Theropoda, Ornithomimosauria) from the Upper Cretaceous Djadokhta Formation of Tögrögiin Shiree, Mongolia.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

First Ornithomimid (Theropoda, Ornithomimosauria) from the Upper Cretaceous Djadokhta Formation of Tögrögiin Shiree, Mongolia.

The Upper Cretaceous Djadokhta Formation has been intensively surveyed for its fossil vertebrate fauna for nearly a century. Amongst other theropods, dromaeosaurids and parvicursorines are common in the formation, but ornithomimosaurs are extremely rare. A new ornithomimosaur material was discovered from the Djadokhta Formation, represented by eolian deposits, of the Tögrögiin Shiree locality, Mongolia. This is only the third ornithomimosaur specimen reported from this formation, and includes the astragalus, the calcaneum, the third distal tarsal, and a complete pes. The new material is clearly belonged to Ornithomimidae by its arctometatarsalian foot condition and has the following unique characters; unevenly developed pair of concavities of the third distal tarsal, curved contacts between the proximal ends of second and fourth metatarsals, the elongate fourth digit, and a laterally inclined medial condyle on phalanx IV-1. These diagnostic characters of the Djadokhtan ornithomimosaur indicate that this is a new taxon. Our phylogenetic analysis supports three clades within derived ornithomimosaurs, and the new taxon is placed a member of the derived ornithomimosaurs. The present specimen is the first ornithomimid record from eolian Tögrögiin Shiree locality, and is indicative of their capability to adapt to arid environments.

The first oviraptorosaur (Dinosauria: Theropoda) bonebed: evidence of gregarious behaviour in a maniraptoran theropod.

A monodominant bonebed of Avimimus from the Nemegt Formation of Mongolia is the first oviraptorosaur bonebed described and the only recorded maniraptoran bonebed from the Late Cretaceous. Cranial elements recovered from the bonebed provide insights on the anatomy of the facial region, which was formerly unknown in Avimimus. Both adult and subadult material was recovered from the bonebed, but small juveniles are underrepresented. The taphonomic and sedimentological evidence suggests that the Avimimus bonebed represents a perimortem gregarious assemblage. The near absence of juveniles in the bonebed may be evidence of a transient age-segregated herd or 'flock', but the behaviour responsible for this assemblage is unclear. Regardless, the Avimimus bonebed is the first evidence of gregarious behaviour in oviraptorosaurs, and highlights a potential trend of increasing gregariousness in dinosaurs towards the end of the Mesozoic.

Specializations of the mandibular anatomy and dentition of Segnosaurus galbinensis (Theropoda: Therizinosauria).

Definitive therizinosaurid cranial materials are exceptionally rare, represented solely by an isolated braincase and tooth in the North American taxon Nothronychus mckinleyi, the remarkably complete skull of the Asian taxon Erlikosaurus andrewsi, and the lower hemimandibles of Segnosaurus galbinensis. To date, comprehensive descriptions of the former taxa are published; however, the mandibular materials of S. galbinensis have remained largely understudied since their initial description in 1979. Here we provide a comprehensive description of the well-preserved hemimandibles and dentition of S. galbinensis (MPC-D 100/80), from the Upper Cretaceous Bayanshiree Formation, Gobi Desert, Mongolia. The subrectangular and ventrally displaced caudal hemimandible, extreme ventral deflection of the rostral dentary, and edentulism of the caudal dentary of S. galbinensis are currently apomorphic among therizinosaurians. Unique, unreported dental traits including lingually folded mesial carinae, development of a denticulated triangular facet on the distal carinae near the cervix, and extracarinal accessory denticles, suggest a highly specialized feeding strategy in S. galbinensis. The presence of triple carinae on the distalmost lateral tooth crowns is also unique, although may represent an abnormality. Contrasted with the simplistic dentition of the contemporaneous therizinosaurid E. andrewsi, the dentition of S. galbinensis is indicative of niche partitioning in food acquisition, processing, or resources among known therizinosaurids inhabiting Asian ecosystems in the Late Cretaceous. Although not quantitatively correlated with diet, this suite of specializations is otherwise unique among theropod dinosaurs and supports derived inferences of facultative or obligate herbivory in therizinosaurids, ultimately adding novel information to our understanding of ecomorphology in theropods.

Resolving the long-standing enigmas of a giant ornithomimosaur Deinocheirus mirificus.

The holotype of Deinocheirus mirificus was collected by the 1965 Polish-Mongolian Palaeontological Expedition at Altan Uul III in the southern Gobi of Mongolia. Because the holotype consists mostly of giant forelimbs (2.4 m in length) with scapulocoracoids, for almost 50 years Deinocheirus has remained one of the most mysterious dinosaurs. The mosaic of ornithomimosaur and non-ornithomimosaur characters in the holotype has made it difficult to resolve the phylogenetic status of Deinocheirus. Here we describe two new specimens of Deinocheirus that were discovered in the Nemegt Formation of Altan Uul IV in 2006 and Bugiin Tsav in 2009. The Bugiin Tsav specimen (MPC-D 100/127) includes a left forelimb clearly identifiable as Deinocheirus and is 6% longer than the holotype. The Altan Uul IV specimen (MPC-D 100/128) is approximately 74% the size of MPC-D 100/127. Cladistic analysis indicates that Deinocheirus is the largest member of the Ornithomimosauria; however, it has many unique skeletal features unknown in other ornithomimosaurs, indicating that Deinocheirus was a heavily built, non-cursorial animal with an elongate snout, a deep jaw, tall neural spines, a pygostyle, a U-shaped furcula, an expanded pelvis for strong muscle attachments, a relatively short hind limb and broad-tipped pedal unguals. Ecomorphological features in the skull, more than a thousand gastroliths, and stomach contents (fish remains) suggest that Deinocheirus was a megaomnivore that lived in mesic environments.

An exquisitely preserved troodontid theropod with new information on the palatal structure from the Upper Cretaceous of Mongolia.

Troodontidae is a clade of small-bodied theropod dinosaurs. A new troodontid, Gobivenator mongoliensis gen. et sp. nov., is described based on the most complete skeleton of a Late Cretaceous member of this clade presently known, from the Campanian Djadokhta Formation in the central Gobi Desert. G. mongoliensis is different from other troodontids in possessing a pointed anterior end of the fused parietal and a fossa on the surangular in front of the posterior surangular foramen. The skull was superbly preserved in the specimen and provides detailed information of the entire configuration of the palate in Troodontidae. Overall morphology of the palate in Gobivenator resembles those of dromaeosaurids and Archaeopteryx, showing an apparent trend of elongation of the pterygoid process of the palatine and reduction of the pterygopalatine suture toward the basal Avialae. The palatal configuration suggests that the skull of Gobivenator would have been akinetic but had already acquired prerequisites for later evolution of cranial kinesis in birds, such as the loss of the epipterygoid and reduction in contact areas among bones.