Yoshitsugu Kobayashi

Hokkaido University | Hokudai · Museum

Mongolia has a long history of discoveries and research related to dinosaur eggs. Since the first scientific discoveries in the early 1920s, numerous eggs, eggshells, and nests have been collected by various international expeditions and have contributed significantly to our understanding of reproductive traits in dinosaurs. In this study, we repor...

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...

Here we report on the new discovery of a large track, attributed to a tyrannosaur, from Aniakchak National Monument in southwestern Alaska. The track is from the Late Cretaceous (Campanian: Maastrichtian) Chignik Formation, a cyclic sequence of rocks, approximately 500-600 m thick, representing shallow marine to nearshore marine environments in the...

A voice box (larynx) is unique for tetrapods and plays functional roles in respiration, airway protection, and vocalization. However, in birds and other reptiles, the larynx fossil is extremely rare, and the evolution of this structure remains largely unknown. Here we report the fossil larynx found in non-avian dinosaurs from ankylosaur Pinacosauru...

Adaptive radiation of archosaurs, represented by crocodilians, non‐avian dinosaurs, and birds, since the Mesozoic has been studied mainly based on their major skeletal elements (skull, vertebrae, and limbs). However, little is known about the evolution of their hyolaryngeal apparatus, which is involved with feeding, respiration, and vocalization, b...

Hadrosauridae, consisting of two subfamilies (Hadrosaurinae and Lambeosaurinae), is a successful herbivorous dinosaur group that established a high taxonomic diversity and a cosmopolitan biogeographic distribution during the Late Cretaceous. While its success is often attributed to a highly specialized oral processing system, the foraging strategy...

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 t...

Here we report a new articulated skeleton of Yamaceratops dorngobiensis (MPC-D 100/553) from the Khugenetjavkhlant locality at the Shine Us Khudag (Javkhlant Formation, ?Santonian-Campanian) of the eastern Gobi Desert, Mongolia, which represents the first substantially complete skeleton and the first juvenile individual of this taxon. The specimen...

The partially correlative Alaskan dinosaur-bearing Prince Creek Formation (PCF), North Slope, lower Cantwell Formation (LCF), Denali National Park, and Chignik Formation (CF), Aniakchak National Monument, form an N–S transect that, together, provides an unparalleled opportunity to examine an ancient high-latitude terrestrial ecosystem. The PCF, 75–...

Goniopholididae is a group of basal neosuchian crocodyliforms closely related to Paralligatoridae and Eusuchia that lived during the Jurassic and Early Cretaceous. Goniopholidids have the long, flat snout and secondary palate of modern crocodylians, the acquisition of which is regarded as a key feature in the early evolution of crocodylian body pla...

A new ankylosaurid dinosaur, Tarchia tumanovae sp. nov., has been recovered from the Upper Cretaceous Nemegt Formation of Mongolia. It includes a well-preserved skull, dorsal, sacral, caudal vertebrae, sixteen dorsal ribs, ilia, a partial ischium, free osteoderms, and a tail club. The squamosal horns of T. tumanovae are divided into two layers, the...

Carcharodontosauria is a group of medium to large-sized predatory theropods, distributed worldwide during the Cretaceous. These theropods were probably the apex predators of Asiamerica in the early Late Cretaceous prior to the ascent of tyrannosaurids, although few Laurasian species are known from this time due to a poor rock record. Here, we descr...

Here we describe a partial hadrosaurid skeleton from the marine Maastrichtian Kita-ama Formation in Japan as a new taxon, Yamatosaurus izanagii gen. et sp. nov., based on unique characters in the dentition. Our phylogenetic analysis demonstrates Yamatosaurus izanagii belongs to Hadrosauridae, composed of Hadrosaurus foulkii + (Yamatosaurus izanagii...

Pterosaurs, which lived during the Mesozoic, were the first known vertebrates to evolve powered flight.¹,² Arboreal locomotion has been proposed for some taxa,³,⁴ and even considered to have played a role in the origin of pterosaur flight.⁵,⁶ Even so, there is still need for comprehensive quantitative ecomorphological analyses.³,⁴ Furthermore, skel...

Here we describe a partial hadrosaurid skeleton from the marine Maastrichtian Kita-ama Formation of the Izumi Group in Japan as a new taxon, Yamatosaurus izanagii gen. et sp. nov. Our phylogenetic analysis demonstrates it belongs to the Hadrosauridae, composed of Hadrosaurus foulkii + ( Yamatosaurus izanagii + (Saurolophinae + Lambeosaurinae). Yama...

The gizzard is the only gastrointestinal organ for mechanical processing in birds. Many birds use grit in the gizzard to enhance mechanical processing efficiency. We conducted an experiment to test the factors that affect chicken grit use in 68 male layer chicks of Gallus gallus domesticus, which were divided into two different groups in gizzard mu...

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 t...

Hadrosaurid fossils from the Liscomb Bonebed (Prince Creek Formation, North Slope, Alaska) were the first dinosaur bones discovered from the Arctic. While the Prince Creek Formation hadrosaurids were long identified as Edmontosaurus, a member of the sub-clade Hadrosaurinae, they were recently assigned to a newly-erected taxon, Ugrunaaluk kuukpikens...

Asian hesperornithiforms are extremely rare in contrast to North American records; thus, their diversity in Asia during the Cretaceous is unclear. Maastrichtian hesperornithiform materials have been reported from both fluvial and marine deposits in North America but only from fluvial deposits in Asia. Asian hesperornithiforms from Maastrichtian dep...

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

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

While there are now numerous records of dinosaurs from Cretaceous rocks around the state of Alaska, very few fossil records of terrestrial vertebrates are known from the Meso-zoic rocks of the southwestern part of the state. Here we report the new discovery of extensive occurrences of dinosaur tracks from Aniakchak National Monument of the Alaska P...

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 Laiya...

Colonial nesting behavior has been inferred in a variety of non-avian dinosaurs based on high concentrations of nests preserved in an area, but sedimentologic and taphonomic evidence demonstrating the contemporaneity of the nests is often lacking. A new nesting site discovered in the Upper Cretaceous Javkhlant Formation of the eastern Gobi Desert,...

Crocodyliform tracks are reported from the Upper Cretaceous ?Cenomanian-Santonian) Bayanshiree Formation in southeastern Mongolia. Ten tracks are preserved as natural casts, forming a trackway with a quadrupedal gait pattern with a tail trail. All tracks are short and wide, and dominated by toe traces without plantar impressions. Pes tracks are cha...

The Prince Creek Formation of Alaska, a rock unit that represents lower coastal plain and delta deposits, is one of the most important formations in the world for understanding vertebrate ecology in the Arctic during the Cretaceous. Here we report on an isolated cranial material, supraoccipital, of a lambeosaurine hadrosaurid from the Liscomb Boneb...

Crocodylian systematics has long been confounded by conflicting hypotheses of higher level relationships—although molecular data sets strongly supported the sister‐taxon relationship of Tomistoma and Gavialis, morphological data sets placed Gavialis as sister to all other living taxa. One of the perceived difficulties in interpreting morphological...

We report details of a unique association of hadrosaur and therizinosaur tracks found in the Late Cretaceous lower Cantwell Formation, Denali National Park, central Alaska Range, Alaska. This rock unit is now well-documented as a source of thousands of fossil footprints of vertebrates such as fishes, pterosaurs, and avialan and non-avialan dinosaur...

Hesperornithiformes were toothed, foot-propelled diving birds and among the most widely distributed groups of birds in the Cretaceous (Late Albian to Maastrichtian) in the Northern Hemisphere. The first species of this group, Hesperornis regalis was discovered from the Niobrara Formation (Upper Santonian) in Kansas in 1871. H. regalis had extremely...

Archosaurs (e.g., crocodilians, pterosaurs, and dinosaurs including birds) are the most diverse and successful clade of terrestrial vertebrates. An understanding of the nesting methods and behaviors of both extinct (e.g., non-avian dinosaurs) and extant archosaurs (i.e., crocodilians and birds) is crucial for the advancement of our understanding of...

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.

Dinosaurs thrived and reproduced in various regions worldwide, including the Arctic. In order to understand their nesting in diverse or extreme environments, the relationships between nests, nesting environments, and incubation methods in extant archosaurs were investigated. Statistical analyses reveal that species of extant covered nesters (i.e.,...

Although two major clades of crocodylians (Alligatoroidea and Crocodyloidea) were split during the Cretaceous period, relatively few morphological and functional differences between them have been known. In addition, interaction of multiple morphofunctional systems that differentiated their ecology has barely been assessed. In this study, we examin...

Crocodylians are ectothermic animals, and their past distribution has been greatly influenced by changing climate since their Cretaceous origin. The Pliocene-Pleistocene witnessed a contraction of the crocodylian latitudinal ranges due to rapid cooling with superimposed pronounced orbital-scale climate oscillations. However, a chronologically-conti...

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 eoli...

A new oviraptorid dinosaur from the Late Cretaceous of Ganzhou, bringing oviraptrotid diversity of this region to seven taxa, is described. It is characterized by a distinct cassowary-like crest on the skull, no pleurocoels on the centra from the second through fourth cervical vertebrae, a neck twice as long as the dorsal vertebral column and sligh...

The Upper Cretaceous Nemegt Formation of Mongolia is rich in well-preserved dinosaurs and Ornithomimosauria is one of the most common taxa in the formation. Three ornithomimosaur taxa, Anserimimus planinychus, Deinocheirus mirificus, and Gallimimus bullatus, have been discovered from the formation so far. However, the recently discovered specimens...

Ankylosaur braincase and endocranial morphologies are poorly known. Furthermore, cranial endocasts have been described for fewer than ten taxa so far. The complete inner ear morphology is known for only three species – Euoplocephalus tutus, Kunbarrasaurus ieversi, and Pawpawsaurus campbelli. Here, the first cranial endocast morphologies are present...

A theropod tracksite was discovered in the Nemegt Formation (Maastrichtian) at Bügiin Tsav, Mongolia by the Korea-Mongolia International Dinosaur Project in 2009. A total of 67 tracks (14 trackways [one didactylous, 13 tridactylous] and 12 isolated tracks) belonging to four ichnomorphotypes were mapped on a single horizon. This indicates at least f...

Asian hesperornithiforms are extremely rare in contrast to the much more abundant record from North America. In Asia, these fossil birds are only known from fragmentary materials from Mongolia. Here we describe the skeletal remains of a new hesperornithiform Chupkaornis keraorum gen. et sp. nov. from the Upper Cretaceous Kashima Formation (Coniacia...

The North American fossil record of dinosaur eggshells for the Cretaceous is primarily restricted to formations of the middle (Albian–Cenomanian) and uppermost (Campanian–Maastrichtian) stages, with a large gap in the record for intermediate stages. Here we describe a dinosaur eggshell assemblage from a formation that represents an intermediate and...

Nipponosaurus sachalinensis is the only definitive lambeosaurine hadrosaurid from Sakhalin Island of Russia. Previous studies suggested it was a member of Lambeosaurini (derived lambeosaurines). However, its phylogenetic status within Lambeosaurini remains controversial. In addition, some studies argued the juvenile ontogenetic stage of the holotyp...

Derived members of the enigmatic mammalian order Desmostylia have molars comprising appressed columns whose morphology does not render their function in feeding simple to discern. Here we describe a new genus and species, Ounalashkastylus tomidai, more derived than Cornwallius but less derived than Desmostylus and Vanderhoofius, which develop a hyp...

The horned dinosaur Centrosaurus apertus from the Belly River Group (Campanian) is represented by multiple articulated skulls and skeletons, and is particularly notable for its occurrence in dozens of large-scale monodominant bonebeds, which have been found in the Dinosaur Park Formation across southern Alberta and Saskatchewan. Here we present a d...

We report on new records of pterosaur tracks from Alaska. Recent palaeontological investigations in the Lower Cantwell Formation of Denali National Park, central Alaska Range, Alaska, had resulted previously in the discovery of a single pterosaur manus track. Subsequent and ongoing investigation has shown that the track record for pterosaurs in thi...

The Ganzhou area of Jiangxi Province, southern China is becoming one of the most productive oviraptorosaurian localities in the world. A new oviraptorid dinosaur was unearthed from the uppermost Upper Cretaceous Nanxiong Formation of Ganzhou area. It is characterized by an anterodorsally sloping occiput and quadrate (a feature shared with Citipati)...

Background Osteohistological examinations of fossil vertebrates have utilized a number of proxies, such as counts and spacing of lines of arrested growth (LAGs) and osteocyte lacunar densities (OLD), in order to make inferences related to skeletochronology and mass-specific growth rates. However, many of these studies rely on samplings of isolated...

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 mosa...

Post-natal parental care seems to have evolved numerous times in vertebrates. Among extant amniotes, it is present in crocodilians, birds, and mammals. However, evidence of this behavior is extremely rare in the fossil record and is only reported for two types of dinosaurs, and a varanopid ‘pelycosaur’. Here we report new evidence for post-natal pa...

Ankylosaur braincase and endocranial morphology is in general poorly known, and only the cranial endocast and inner ear of Euoplocephalus, an ankylosaurid from North America, and the cranial endocasts of Sauropelta and another unnamed nodosaurid from Japan have been described in detail. The first 3D reconstructions of the brains and inner ears of t...

Herd structure in Late Cretaceous polar dinosaurs: A remarkable new Email alerting services articles cite this article to receive free e-mail alerts when new www.gsapubs.org/cgi/alerts click Subscribe to subscribe to Geology www.gsapubs.org/subscriptions/ click Permission request to contact GSA http://www.geosociety.org/pubs/copyrt.htm#gsa click of...

Herd structure in Late Cretaceous polar dinosaurs: A remarkable new Email alerting services articles cite this article to receive free e-mail alerts when new www.gsapubs.org/cgi/alerts click Subscribe to subscribe to Geology www.gsapubs.org/subscriptions/ click Permission request to contact GSA http://www.geosociety.org/pubs/copyrt.htm#gsa click of...

The pelvic structure in non-avian archosaurs plays a key role in understanding the evolution of terrestrial locomotor patterns because the pelvis contains major attachment sites for proximal hind limb musculature. In order to investigate patterns of pelvic evolution in archosaurs, this study compiled three pelvic indices, as well as femoral head or...

Two types of small-to medium-sized theropod tracks are described for the first time from the Lower Cantwell Formation (Upper Cretaceous) of Denali National Park in the central Alaska Range, Alaska. The Lower Cantwell comprises some 4000 m of fluvial channel and floodplain basin fill of several sedimentary lithologies. The tracks are assigned to the...

Therizinosauria are an unusual group of theropod dinosaurs, found mostly in the Cretaceous deposits in Mongolia, China and western USA. The basal forms of this group are represented by incomplete or disarticulated material. Here, we report a nearly complete, articulated skeleton of a new basal therizinosaur from the Early Cretaceous Yixian Formatio...

Character states for Jianchangosaurus yixianensis, used in the phylogenetic analysis in this study and synapomorphies at each node in all of most parsimonious trees ( Figure 12 ), obtained by TNT. (DOCX)

Bonebeds can provide a wealth of anatomical, taphonomic, and ontogenetic information about the specimens preserved within them, and can provide evidence for inferred behavior. The material described here represents the first known bonebed of ornithomimids in North America, and the fourth record of an ornithomimosaur bonebed in the world. Partial sk...

Skeletal measurements of CMN 12068, 12069, and 12070. Asterisk (*) indicates fragmentary, distorted or incomplete remains. Dash (-) indicates elements that are not present. (XLS)