Exceptionally preserved stomach contents of a young tyrannosaurid reveal an ontogenetic dietary shift in an iconic extinct predator.

Tyrannosaurids were large carnivorous dinosaurs that underwent major changes in skull robusticity and body proportions as they grew, suggesting that they occupied different ecological niches during their life span. Although adults commonly fed on dinosaurian megaherbivores, the diet of juvenile tyrannosaurids is largely unknown. Here, we describe a remarkable specimen of a juvenile Gorgosaurus libratus that preserves the articulated hindlimbs of two yearling caenagnathid dinosaurs inside its abdominal cavity. The prey were selectively dismembered and consumed in two separate feeding events. This predator-prey association provides direct evidence of an ontogenetic dietary shift in tyrannosaurids. Juvenile individuals may have hunted small and young dinosaurs until they reached a size when, to satisfy energy requirements, they transitioned to feeding on dinosaurian megaherbivores. Tyrannosaurids occupied both mesopredator and apex predator roles during their life span, a factor that may have been key to their evolutionary success.

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.

An ankylosaur larynx provides insights for bird-like vocalization in non-avian dinosaurs.

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 Pinacosaurus grangeri. The larynx of Pinacosaurus is composed of the cricoid and arytenoid like non-avian reptiles, but specialized with the firm and kinetic cricoid-arytenoid joint, prominent arytenoid process, long arytenoid, and enlarged cricoid, as a possible vocal modifier like birds rather than vocal source like non-avian reptiles. Although bird-unique vocal source (syrinx) have never been reported in non-avian dinosaurs, Pinacosaurus could have employed bird-like vocalization with the bird-like large, kinetic larynx. This oldest laryngeal fossil from the Cretaceous dinosaur provides the first step for understanding the vocal evolution in non-avian dinosaurs toward birds.

Evolutionary insights from an anatomical network analysis of the hyolaryngeal apparatus in extant archosaurs (birds and crocodilians).

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, because of poor fossil preservation and the difficulty in determining the musculoskeletal homology of the apparatus. Network analysis is a framework to quantitatively characterize the topological organization of anatomical structures for comparing structural integration and modularity regardless of ambiguous homology. Herein, we modeled the musculoskeletal system of hyolarynx in six species of extant archosaurs and its sister-taxon turtle, and conducted a network analysis using network parameters, modular partition, and bone centrality in a phylogenetic framework. The network parameters reveal that ancestral archosaurs have reduced the numbers of elements and links and acquired complex networks as a whole domain with strong modularity in the hyolarynx. Furthermore, the modular partition and centrality reveal that the hyoids are highly evolvable, while the larynx is constrained and less evolvable. The archosaur hyolarynx exhibits different evolutionary trends: crocodilians with the system integration, basihyal simplification, and ceratobranchial centralization; and birds with the simplicity, weak integration, and modularity of the hyolarynx, laryngeal integration with cricoid centrality, and tongue-module expansion with the acquisition of paraglossal. Four hyolaryngeal bones (ceratobranchial, basihyal, paraglossal, and cricoid) have played important roles in archosaur evolution, and their fossil records are keys to understanding the two major archosaur lineages toward crocodilians and birds.

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 juvenile Yamaceratops (Dinosauria, Ceratopsia) from the Javkhlant Formation (Upper Cretaceous) of Mongolia.

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 includes a nearly complete cranium and large portions of the vertebral column and appendicular skeleton. Its skull is about 2/3 the size of the holotype specimen, based on mandibular length. Its juvenile ontogenetic stage is confirmed by multiple indicators of skeletal and morphological immaturity known in ceratopsians, such as the long-grained surface texture on the long bones, the smooth external surface on the postorbital, open neurocentral sutures of all caudal vertebrae, a large orbit relative to the postorbital and jugal, the low angle of the lacrimal ventral ramus relative to the maxillary teeth row, narrow frontal, and straight ventral edge of the dentary. Osteohistological analysis of MPC-D 100/553 recovered three lines of arrested growth, implying around 3 years of age when it died, and verified this specimen's immature ontogenetic stage. The specimen adds a new autapomorphy of Yamaceratops, the anteroventral margin of the fungiform dorsal end of the lacrimal being excluded from the antorbital fossa. Furthermore, it shows a unique combination of diagnostic features of some other basal neoceratopsians: the ventrally hooked rostral bone as in Aquilops americanus and very tall middle caudal neural spines about or more than four times as high as the centrum as in Koreaceratops hwaseongensis, Montanoceratops cerorhynchus, and Protoceratops andrewsi. The jugal with the subtemporal ramus deeper than the suborbital ramus as in the holotype specimen is also shared with A. americanus, Liaoceratops yanzigouensis, and juvenile P. andrewsi. Adding 38 new scorings into the recent comprehensive data matrix of basal Neoceratopsia and taking into account the ontogenetically variable characters recovered Y. dorngobiensis as the sister taxon to Euceratopsia (Leptoceratopsidae plus Coronosauria). A second phylogenetic analysis with another matrix for Ceratopsia also supported this position. The new phylogenetic position of Y. dorngobiensis is important in ceratopsian evolution, as this taxon represents one of the basalmost neoceratopsians with a broad, thin frill and hyper-elongated middle caudal neural spines while still being bipedal.

A new goniopholidid from the Upper Jurassic Morrison Formation, USA: novel insight into aquatic adaptation toward modern crocodylians.

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 plan and their aquatic adaptation. Here, we report a new species, Amphicotylus milesi, with the description from the best-preserved specimen to date of Goniopholididae from Wyoming, USA. Its posterior extension of the nasopharyngeal passage (pterygoid secondary palate) and the shortening and dorsal deflection of the ceratobranchial suggest that basal neosuchians could raise their gular valve to separate oral and pharyngeal cavities as in modern crocodylians. The anatomy of Amphicotylus milesi sheds light on the acquisition of this new respiratory system in the crocodyliform evolution and their early aquatic adaptation, leading to modern crocodylians.

A new ankylosaurid from the Upper Cretaceous Nemegt Formation of Mongolia and implications for paleoecology of armoured dinosaurs.

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 external dermal layer and the underlying squamosal horn proper. The irregular ventral margin of the base of the upper dermal layer may represent a resorption surface, suggesting that the squamosal horns of some ankylosaurids underwent extreme ontogenetic remodeling. Localized pathologies on the dorsosacral ribs and the tail provide evidence of agonistic behaviour. The tail club knob asymmetry of T. tumanovae resulted from restricted bone growth due to tail club strikes. Furthermore, T. tumanovae had an anteriorly protruded shovel-shaped beak, which is a morphological character of selective feeders. Ankylosaurid diets shifted from low-level bulk feeding to selective feeding during the Baruungoyot and the Nemegt "age" (middle Campanian-lower Maastrichtian). This ankylosaurid niche shifting might have been a response to habitat change and competition with other bulk-feeding herbivores.

A new carcharodontosaurian theropod dinosaur occupies apex predator niche in the early Late Cretaceous of Uzbekistan.

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 describe Ulughbegsaurus uzbekistanensis gen. et sp. nov. from the early Late Cretaceous (Turonian) of Central Asia, which represents the first record of a Late Cretaceous carcharodontosaurian from the region. This new taxon is represented by a large, isolated maxilla from the Bissekty Formation of the Kyzylkum Desert, the Republic of Uzbekistan, a formation yielding a rich and diverse assemblage of dinosaurs and other vertebrates from fragmentary remains. Comparison of the maxilla with that of other allosauroids indicates Ulughbegsaurus was 7.5-8 m in body length and greater than 1000 kg in body mass, suggesting it was the previously unrecognized apex predator of the Bissekty ecosystem while smaller known tryannosauroids and dromaeosaurids were probable mesopredators. The discovery of Ulughbegsaurus records the geologically latest stratigraphic co-occurrence of carcharodontosaurid and tyrannosauroid dinosaurs from Laurasia, and evidence indicates carcharodontosaurians remained the dominant predators relative to tyrannosauroids, at least in Asia, as late as the Turonian.

A new darwinopteran pterosaur reveals arborealism and an opposed thumb.

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, skeletal adaptations correlated to specialized lifestyles are often difficult to recognize and interpret in fossils. Here we report on a new darwinopteran pterosaur that inhabited a unique forest ecosystem from the Jurassic of China. The new species exhibits the oldest record of palmar (or true) opposition of the pollex, which is unprecedented for pterosaurs and represents a sophisticated adaptation related to arboreal locomotion. Principal-coordinate analyses suggest an arboreal lifestyle for the new species but not for other closely related species from the same locality, implying a possible case of ecological niche partitioning. The discovery adds to the known array of pterosaur adaptations and the history of arborealism in vertebrates. It also adds to the impressive early bloom of arboreal communities in the Jurassic of China, shedding light on the history of forest environments.

A new basal hadrosaurid (Dinosauria: Ornithischia) from the latest Cretaceous Kita-ama Formation in Japan implies the origin of hadrosaurids.

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 + (Saurolophinae + Lambeosaurinae)). The coracoid lacks a biceps tubercle as in non-hadrosaurid hadrosauroids, suggesting its presence is a key feature for the clade of Saurolophinae and Lambeosaurinae. The evolutionary rates analysis further supports that shoulder and forelimb features, which are likely to have been involved in locomotion, were important for the early evolution of Hadrosauridae. Our biogeographic analyses show that basal hadrosaurids were widely distributed in Asia and Appalachia, that the clade of Saurolophinae and Lambeosaurinae originated in Asia, and that eastern Asia may have served as a refugium of relict hadrosauroid taxa such as Plesiohadros djadokhtaensis, Tanius sinensis, and Yamatosaurus izanagii during the Late Cretaceous. The contemporaneous occurrence of basal (Yamatosaurus izanagii) and derived (Kamuysaurus japonicus) hadrosaurids during the Maastrichtian in Japan is the first record in Asia. Because of the long geographical distance between these localities, they likely did not co-exist, but instead demonstrate some level of provinciality.

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.

Effects of diet and gizzard muscularity on grit use in domestic chickens.

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 muscularity (high and low). Within each muscularity group, two different diets were provided (herbivory and non-herbivory) to test whether diet and gizzard muscularity affect grit characteristics including amount, size, and shape (circularity, roundness, and solidity) at different stages of digestion (ingested grit, grit in gizzard, and excreted grit). All animals ingested more grit than they excreted, possibly because excreted grit was below the detection size limit of 0.5 mm of the present study. The amounts of grit ingested and remained in the gizzard were larger in herbivorous groups, but these groups excreted less grit. Larger, rougher grit was selectively ingested by all chicks, but size preferences were especially pronounced in the herbivorous groups. Grit in the gizzard tended to be larger in herbivorous groups, but the grit in excreta was smaller, whereas the size of excreted grit was larger in groups with less muscular gizzards. Grit in the gizzard was much smoother than the offered and ingested grit, especially in the herbivorous, muscular gizzard groups. Excreted grit in all groups was smoother than the offered grit. These results show that diet affects the characteristics of ingested grit, grit in the gizzard, and excreted grit, whereas gizzard muscularity affects the characteristics of grit in the gizzard and excreted grit. The use of larger sizes and amounts of grit by herbivorous groups may be a response to the needs of digesting hard, coarse materials. The recovered behavioral flexibility of grit use might reflect the omnivorous nature of Gallus gallus domesticus and may aid smooth dietary shifts. The results also show that the shape of grit remaining in the gizzard does not reflect the initial shape of ingested grit, in contrast to previously published ideas. Instead, the shape of grit in the gizzard more closely reflects the diet and gizzard muscularity of chicks.

Re-examination of the cranial osteology of the Arctic Alaskan hadrosaurine with implications for its taxonomic status.

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 kuukpikensis. However, taxonomic status of the new taxon is ambiguous largely due to the immature nature of the specimens upon which it was based. Here we reexamine cranial elements of the Prince Creek Formation hadrosaurine in order to solve its taxonomic uncertainties. The Prince Creek Formation hadrosaurine possesses a short dorsolateral process of the laterosphenoid, one of the diagnostic characters of Edmontosaurus. The Prince Creek Formation hadrosaurine also shows affinity to Edmontosaurus regalis in the presence of a horizontal shelf of the jugal. Our morphological comparisons with other North American Edmontosaurus specimens and our phylogenetic analyses demonstrate that the Prince Creek Formation hadrosaurine should be re-assigned to Edmontosaurus. Because the Prince Creek Formation Edmontosaurus shows differences with lower latitude Edmontosaurus in a dorsoventrally short maxilla, presence of a secondary ridge on the dentary teeth, and the absence of the transverse ridge between basipterygoid processes of the basisphenoid, we consider that the Prince Creek Formation Edmontosaurus should be regarded as Edmontosaurus sp. until further discoveries of mature hadrosaurines from the Prince Creek Formation Bonebed and/or equivalently juvenile Edmontosaurus specimens from the lower latitudes allow direct comparisons. The retention of the Prince Creek Formation hadrosaurine as Edmontosaurus re-establishes a significant latitudinal distribution for this taxon. Despite the large latitudinal distribution of the taxon, the morphological disparity of Edmontosaurus is small within Hadrosaurinae. The small morphological disparity may be related to the relatively low latitudinal temperature gradient during the latest Cretaceous compared to present day, a gradient which might not have imposed significant pressure for much morphological adaptations across a broad latitudinal range.

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.

Dinosaur ichnology and sedimentology of the Chignik Formation (Upper Cretaceous), Aniakchak National Monument, southwestern Alaska; Further insights on habitat preferences of high-latitude hadrosaurs.

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 Mesozoic 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 Peninsula. These tracks are in the Late Cretaceous (Maastrichtian) Chignik Formation, a cyclic sequence of rocks, approximately 500-600 m thick, representing shallow marine to nearshore marine environments in the lower part and continental alluvial coastal plain environments in the upper part of the section. These rocks are part of the Peninsular Terrane and paleomagnetic reconstructions based on the volcanic rocks of this terrane suggest that the Chignik Formation was deposited at approximately its current latitude which is almost 57° N. Recent field work in Aniakchak National Monument has revealed over 75 new track sites, dramatically increasing the dinosaur record from the Alaska Peninsula. Most of the combined record of tracks can be attributed to hadrosaurs, the plant-eating duck-billed dinosaurs. Tracks range in size from those made by full-grown adults to juveniles. Other tracks can be attributed to armored dinosaurs, meat-eating dinosaurs, and two kinds of fossil birds. The track size of the predatory dinosaur suggests a body approximately 6-7 m long, about the estimated size of the North Slope tyrannosaurid Nanuqsaurus. The larger bird tracks resemble Magnoavipes denaliensis previously described from Denali National Park, while the smaller bird tracks were made by a bird about the size of a modern Willet. Previous interdisciplinary sedimentologic and paleontologic work in the correlative and well-known dinosaur bonebeds of the Prince Creek Formation 1400km-1500km further north in Alaska suggested that high-latitude hadrosaurs preferred distal coastal plain or lower delta plain habitats. The ichnological record being uncovered in the Chignik Formation of southwestern Alaska is showing that the hadrosaur tracks here were also made in distal coastal and delta plain conditions. This similarity may corroborate the habitat preference model for Cretaceous high-latitude dinosaurs proposed for the data gathered from the Prince Creek Formation, and may indicate that at least Beringian hadrosaurids had similar habitat preferences regardless of latitude.

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.

The First Definite Lambeosaurine Bone From the Liscomb Bonebed of the Upper Cretaceous Prince Creek Formation, Alaska, United States.

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 Bonebed of the Prince Creek Formation. The lambeosaurine supraoccipital has well-developed squamosal bosses and a short sutural surface with the exoccipital-opisthotic complex, and is similar to lambeosaurine supraoccipitals from the Dinosaur Park Formation in having anteriorly positioned squamosal bosses. Affinities with Canadian lambeosaurines elucidate more extensive faunal exchange between the Arctic and lower paleolatitudes which was previously suggested by the presence of Edmontosaurus, Pachyrhinosaurus, tyrannosaurids, and troodontids in both regions. The presence of one lambeosaurine and nine hadrosaurine supraoccipitals in the Liscomb Bonebed suggests hadrosaurine dominated faunal structure as in the Careless Creek Quarry of the USA that was also deposited under a near-shore environment. It differs from the lambeosaurine dominant structures of localities in Russia and China interpreted as inland environments. This may suggest that lambeosaurines had less preference for near-shore environments than hadrosaurines in both Arctic and lower paleolatitudes.

Mosaic nature in the skeleton of East Asian crocodylians fills the morphological gap between "Tomistominae" and Gavialinae.

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 character evolution on the molecular tree is the extensive character reversal occurring in Gavialinae, the mechanism of which has yet to be explained. Here, we provide evidence of gavialine-specific atavistic characters from East Asian "tomistomines" Penghusuchus pani and Toyotamaphimeia machikanensis. These taxa exhibit a mosaic assembly of "tomistomine" and gavialine features, which fill the gap between the two longirostrine groups. Although the parsimony analysis of morphological data (69 taxa, 254 characters) still supports the previous morphological hypothesis, the alternative tree that was forced to fit the molecular hypothesis was insignificantly (5/954 steps; 0.52%) longer than the unconstrained tree, suggesting that morphological evolution can also be interpreted on the molecular tree. Although the problem of stratigraphic gaps remains, future studies may be directed to resolving the interrelationships within Gavialoidea, a large longirostrine group of crocodylians, in the molecular tree context.