An extraordinary fossil captures the struggle for existence during the Mesozoic.

Dinosaurs and mammals have coexisted for the last ~ 230 million years. Both groups arose during the Late Triassic and diversified throughout the Mesozoic and into the Cenozoic (the latter in the form of birds). Although they undoubtedly interacted in many ways, direct fossil evidence for their interaction is rare. Here we report a new fossil find from the Lujiatun Member of the Lower Cretaceous Yixian Formation of China, showing a gobiconodontid mammal and psittacosaurid dinosaur locked in mortal combat. We entertain various hypothesized explanations for this association, but the balance of the evidence suggests that it represents a predation attempt on the part of the smaller mammal, suddenly interrupted by, and preserved within, a lahar-type volcanic debris flow. Mesozoic mammals are usually depicted as having lived in the shadows of their larger dinosaurian contemporaries, but this new fossil convincingly demonstrates that mammals could pose a threat even to near fully-grown dinosaurs. The Yixian Formation-and the Chinese fossil Jehol Biota more broadly-have played a particularly important role in revealing the diversity of small-bodied dinosaurs and other fauna. We anticipate that the volcanically derived obrution deposits specific to the Lujiatun Member will likewise continue to yield evidence for biotic interactions otherwise unknown from the rest of the fossil record.

Rare juvenile material constrains estimation of skeletal allometry in Gryposaurus notabilis (Dinosauria: Hadrosauridae).

In studying the skeletal allometry of any vertebrate, it is important to sample the ontogenetic extremes to ensure the accuracy of parameter estimation; this is particularly true for fossil taxa, where sampling of ontogenetic series is incomplete and sporadic. Previous studies have examined allometry in the skull of the duck-billed dinosaur Gryposaurus notabilis, but these did not include individuals smaller than ~65% the maximum known size (based on linear dimensions). Here, we report on the two smallest known examples of this species (a mostly complete skeleton and a partial skull), which are ~37% the known maximal size of G. notabilis. Osteohistology indicates that these represent individuals ~2 years of age. Allometric analysis demonstrates that most aspects of the skull of G. notabilis grew isometrically, although the height of the nasal arch grew with positive allometry. Early in the ontogeny of G. notabilis, the dentary teeth possessed secondary ridges, which were lost later in life. This finding has important bearing on hadrosaurid tooth taxonomy. The limb proportions of G. notabilis largely grew isometrically (or with weak negative allometry, at most), like some other hadrosaurids, suggesting that the species did not undergo a gait shift with increasing age (unless it occurred very early in ontogeny). We argue that the lack of significant locomotory performance compensation exhibited by young hadrosaurids helps to explain why they apparently formed small, mutualistic aggregations, presumably for protection from large predators.

"Dragons" on the landscape: Modeling the abundance of large carnivorous dinosaurs of the Upper Jurassic Morrison Formation (USA) and the Upper Cretaceous Dinosaur Park Formation (Canada).

Counts of the number of skeletal specimens of "adult" megaherbivores and large theropods from the Morrison and Dinosaur Park formations-if not biased by taphonomic artifacts-suggest that the big meat-eaters were more abundant, relative to the number of big plant-eaters, than one would expect on the basis of the relative abundance of large carnivores and herbivores in modern mammalian faunas. Models of megaherbivore population density (number of individuals per square kilometer) that attempt to take into account ecosystem productivity, the size structure of megaherbivore populations, and individual megaherbivore energy requirements, when combined with values of the large theropod/megaherbivore abundance ratio, suggest that large theropods may have been more abundant on the landscape than estimates extrapolated from the population density versus body mass relationship of mammalian carnivores. Models of the meat production of megaherbivore populations and the meat requirements of "adult" large theropods suggest that herbivore productivity would have been insufficient to support the associated number of individuals of "adult" large theropods, unless the herbivore production/biomass ratio was substantially higher, and/or the large theropod meat requirement markedly lower, than expectations based on modern mammals. Alternatively, or in addition to one or both of these other factors, large theropods likely included dinosaurs other than megaherbivores as significant components of their diet.

The appendicular myology of Stegoceras validum (Ornithischia: Pachycephalosauridae) and implications for the head-butting hypothesis.

In this study, we use an exceptional skeleton of the pachycephalosaur Stegoceras validum (UALVP 2) to inform a comprehensive appendicular muscle reconstruction of the animal, with the goal of better understanding the functional morphology of the pachycephalosaur postcranial skeleton. We find that S. validum possessed a conservative forelimb musculature, particularly in comparison to early saurischian bipeds. By contrast, the pelvic and hind limb musculature are more derived, reflecting peculiarities of the underlying skeletal anatomy. The iliotibialis, ischiocaudalis, and caudofemoralis muscles have enlarged attachment sites and the caudofemoralis has greater leverage owing to the distal displacement of the fourth trochanter along the femur. These larger muscles, in combination with the wide pelvis and stout hind limbs, produced a stronger, more stable pelvic structure that would have proved advantageous during hypothesized intraspecific head-butting contests. The pelvis may have been further stabilized by enlarged sacroiliac ligaments, which stemmed from the unique medial iliac flange of the pachycephalosaurs. Although the pubis of UALVP 2 is not preserved, the pubes of other pachycephalosaurs are highly reduced. The puboischiofemoralis musculature was likely also reduced accordingly, and compensated for by the aforementioned improved pelvic musculature.

Hadrosauroid eggs and embryos from the Upper Cretaceous (Maastrichtian) of Jiangxi Province, China.

BACKGROUND: Dinosaur eggs containing embryos are rare, limiting our understanding of dinosaur development. Recently, a clutch of subspherical dinosaur eggs was discovered while blasting for a construction project in the Upper Cretaceous red beds (Hekou Formation) of the Ganzhou Basin, Jiangxi Province, China. At least two of the eggs contain identifiable hadrosauroid embryos, described here for the first time. RESULTS: The eggs, attributable to Spheroolithidae indet., are thin-walled and small (~ 660 mL) compared to those of Lambeosaurinae. The shape of the embryonic squamosal is reminiscent of that seen in the Late Cretaceous hadrosauroids Levnesovia transoxiana, Tanius sinensis, and Nanningosaurus dashiensis, suggestive of possible affinities. CONCLUSION: The small size of the eggs and embryos, similar to those of Hadrosaurinae, indicates that the larger eggs and hatchlings typical of Lambeosaurinae are evolutionarily derived.

A new model of forelimb ecomorphology for predicting the ancient habitats of fossil turtles.

Various morphological proxies have been used to infer habitat preferences among fossil turtles and their early ancestors, but most are tightly linked to phylogeny, thereby minimizing their predictive power. One particularly widely used model incorporates linear measurements of the forelimb (humerus + ulna + manus), but in addition to the issue of phylogenetic correlation, it does not estimate the likelihood of habitat assignment. Here, we introduce a new model that uses intramanual measurements (digit III metacarpal + non-ungual phalanges + ungual) to statistically estimate habitat likelihood and that has greater predictive strength than prior estimators. Application of the model supports the hypothesis that stem-turtles were primarily terrestrial in nature and recovers the nanhsiungchelyid Basilemys (a fossil crown-group turtle) as having lived primarily on land, despite some prior claims to the contrary.

Ontogeny of a sexually selected structure in an extant archosaur Gavialis gangeticus (Pseudosuchia: Crocodylia) with implications for sexual dimorphism in dinosaurs.

Despite strong evidence for sexual selection in various display traits and other exaggerated structures in large extinct reptiles, such as dinosaurs, detecting sexual dimorphism in them remains difficult. Their relatively small sample sizes, long growth periods, and difficulties distinguishing the sexes of fossil specimens mean that there are little compelling data on dimorphism in these animals. The extant gharial (Gavialis gangeticus) is a large and endangered crocodylian that is sexually dimorphic in size, but males also possesses a sexually selected structure, the ghara, which has an osteological correlate in the presence of a fossa associated with the nares. This makes the species a unique model for potentially assessing dimorphism in fossil lineages, such as dinosaurs and pterosaurs, because it is a large, slow-growing, egg-laying archosaur. Here we assess the dimorphism of G. gangeticus across 106 specimens and show that the presence of a narial fossa diagnoses adult male gharials. Males are larger than females, but the level of size dimorphism, and that of other cranial features, is low and difficult to detect without a priori knowledge of the sexes, even with this large dataset. By extension, dimorphism in extinct reptiles is very difficult to detect in the absence of sex specific characters, such as the narial fossa.

The internal cranial anatomy of Champsosaurus (Choristodera: Champsosauridae): Implications for neurosensory function.

Although isolated Champsosaurus remains are common in Upper Cretaceous sediments of North America, the braincase of these animals is enigmatic due to the fragility of their skulls. Here, two well-preserved specimens of Champsosaurus (CMN 8920 and CMN 8919) are CT scanned to describe their neurosensory structures and infer sensory capability. The anterior portion of the braincase was poorly ossified and thus does not permit visualization of a complete endocast; however, impressions of the olfactory stalks indicate that they were elongate and likely facilitated good olfaction. The posterior portion of the braincase is ossified and morphologically similar to that of other extinct diapsids. The absence of an otic notch and an expansion of the pars inferior of the inner ear suggests Champsosaurus was limited to detecting low frequency sounds. Comparison of the shapes of semicircular canals with lepidosaurs and archosauromorphs demonstrates that the semicircular canals of Champsosaurus are most similar to those of aquatic reptiles, suggesting that Champsosaurus was well adapted for sensing movement in an aquatic environment. This analysis also demonstrates that birds, non-avian archosauromorphs, and lepidosaurs possess significantly different canal morphologies, and represents the first morphometric analysis of semicircular canals across Diapsida.

Computed tomography analysis of the cranium of Champsosaurus lindoei and implications for the choristoderan neomorphic ossification.

Choristoderes are extinct neodiapsid reptiles that are well known for their unusual cranial anatomy, possessing an elongated snout and expanded temporal arches. Although choristodere skulls are well described externally, their internal anatomy remains unknown. An internal description was needed to shed light on peculiarities of the choristodere skull, such as paired gaps on the ventral surface of the skull that may pertain to the fenestra ovalis, and a putative neomorphic ossification in the lateral wall of the braincase. Our goals were: (i) to describe the cranial elements of Champsosaurus lindoei in three dimensions; (ii) to describe paired gaps on the ventral surface of the skull to determine if these are indeed the fenestrae ovales; (iii) to illustrate the morphology of the putative neomorphic bone; and (iv) to consider the possible developmental and functional origins of the neomorph. We examined the cranial anatomy of the choristodere Champsosaurus lindoei (CMN 8920) using high-resolution micro-computed tomography scanning. We found that the paired gaps on the ventral surface of the skull do pertain to the fenestrae ovales, an unusual arrangement that may be convergent with some plesiosaurs, some aistopods, and some urodeles. The implications of this morphology in Champsosaurus are unknown and will be the subject of future work. We found that the neomorphic bone is a distinct ossification, but is not part of the wall of the brain cavity or the auditory capsule. Variation in the developmental pathways of cranial bones in living amniotes was surveyed to determine how the neomorphic bone may have developed. We found that the chondrocranium and splanchnocranium show little to no variation across amniotes, and the neomorphic bone is therefore most likely to have developed from the dermatocranium; however, the stapes is a pre-existing cranial element that is undescribed in choristoderes and may be homologous with the neomorphic bone. If the neomorphic bone is not homologous with the stapes, the neomorph likely developed from the dermatocranium through incomplete fusion of ossification centres from a pre-existing bone, most likely the parietal. Based on the apparent morphology of the neomorph in Coeruleodraco, the neomorph was probably too small to play a significant structural role in the skull of early choristoderes and it may have arisen through non-adaptive means. In neochoristoderes, such as Champsosaurus, the neomorph was likely recruited to support the expanded temporal arches.

A Problematic Tyrannosaurid (Dinosauria: Theropoda) Skeleton and Its Implications for Tyrannosaurid Diversity in the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta.

Several published censuses have noted the presence of two tyrannosaurids, Daspletosaurus sp. and Albertosaurus sarcophagus, within the Upper Cretaceous Horseshoe Canyon Formation of Alberta. Although A. sarcophagus is known from more than a dozen major discoveries in these strata, Daspletosaurus sp. is known from just a single problematic skeleton (lacking most of the skull) of a young individual. Here we describe and figure this skeleton, and marshal a variety of osteohistologic, morphometric, and phylogenetic methods to accurately determine its taxonomic status. Although none of these methods individually provides convincing evidence regarding the affinities of the specimen, together (and including other historical and biostratigraphic considerations) they strongly imply that the skeleton instead pertains to a young A. sarcophagus. In this way, we show that only a single species of tyrannosaurid is definitively present in the Horseshoe Canyon Formation, greatly simplifying interpretations of tyrannosaurid evolution and ecology in this setting. Anat Rec, 303:673-690, 2020. © 2019 The Authors. The Anatomical Record published by Wiley Periodicals, Inc. on behalf of American Association of Anatomists.

Competition structured a Late Cretaceous megaherbivorous dinosaur assemblage.

Modern megaherbivore community richness is limited by bottom-up controls, such as resource limitation and resultant dietary competition. However, the extent to which these same controls impacted the richness of fossil megaherbivore communities is poorly understood. The present study investigates the matter with reference to the megaherbivorous dinosaur assemblage from the middle to upper Campanian Dinosaur Park Formation of Alberta, Canada. Using a meta-analysis of 21 ecomorphological variables measured across 14 genera, contemporaneous taxa are demonstrably well-separated in ecomorphospace at the family/subfamily level. Moreover, this pattern is persistent through the approximately 1.5 Myr timespan of the formation, despite continual species turnover, indicative of underlying structural principles imposed by long-term ecological competition. After considering the implications of ecomorphology for megaherbivorous dinosaur diet, it is concluded that competition structured comparable megaherbivorous dinosaur communities throughout the Late Cretaceous of western North America.

Supplementary cranial description of the types of Edmontosaurus regalis (Ornithischia: Hadrosauridae), with comments on the phylogenetics and biogeography of Hadrosaurinae.

The cranial anatomy of the flat-skulled hadrosaurine Edmontosaurus regalis (Ornithischia: Hadrosauridae) is extensively described here, based on the holotype and paratype collected from the middle part of the Horseshoe Canyon Formation in southern Alberta. Focus is given to previously undocumented features of ontogenetic and phylogenetic importance. This description facilitates overall osteological comparisons between E. regalis and other hadrosaurids (especially E. annectens), and revises the diagnosis of E. regalis, to which a new autapomorphy (the dorsal half of the jugal anterior process bearing a sharp posterolateral projection into the orbit) is added. We consider the recently named Ugrunaaluk kuukpikensis from the upper Campanian/lower Maastrichtian of Alaska a nomen dubium, and conservatively regard the Alaskan material as belonging to Edmontosaurus sp.. A phylogenetic analysis of Hadrosauroidea using maximum parsimony further corroborates the sister-taxon relationship between E. regalis and E. annectens. In the strict consensus tree, Hadrosaurus foulkii occurs firmly within the clade comprising all non-lambeosaurine hadrosaurids, supporting the taxonomic scheme that divides Hadrosauridae into Hadrosaurinae and Lambeosaurinae. Within Edmontosaurini, Kerberosaurus is posited as the sister taxon to the clade of Shantungosaurus + Edmontosaurus. The biogeographic reconstruction of Hadrosaurinae in light of the time-calibrated cladogram and probability calculation of ancestral areas for all internal nodes reveals a significantly high probability for the North American origin of the clade. However, the Laramidia-Appalachia dispersals around the Santonian-Campanian boundary, inferred from the biogeographic scenario for the North American origin of Hadrosaurinae, are in conflict with currently accepted paleogeographic models. By contrast, the Asian origin of Hadrosaurinae with its relatively low probability resulting from the biogeographic analysis is worth seriously considering, despite the lack of fossil material from the Santonian and lower Campanian of Asia. Extra fossil collecting in appropriate geographic locations and stratigraphic intervals of Asia and Europe will help to clarify the biogeographic dynamics of hadrosaurine dinosaurs in the near future.

Spiclypeus shipporum gen. et sp. nov., a Boldly Audacious New Chasmosaurine Ceratopsid (Dinosauria: Ornithischia) from the Judith River Formation (Upper Cretaceous: Campanian) of Montana, USA.

This study reports on a new ceratopsid, Spiclypeus shipporum gen et sp. nov., from the lower Coal Ridge Member of the Judith River Formation in Montana, USA, which dates to ~76 Ma (upper Campanian). The species is distinguished by rugose dorsal contacts on the premaxillae for the nasals, laterally projecting postorbital horncores, fully fused and anteriorly curled P1 and P2 epiparietals, and a posterodorsally projecting P3 epiparietal. The holotype specimen is also notable for its pathological left squamosal and humerus, which show varied signs of osteomyelitis and osteoarthritis. Although the postorbital horncores of Spiclypeus closely resemble those of the contemporaneous 'Ceratops', the horncores of both genera are nevertheless indistinguishable from those of some other horned dinosaurs, including Albertaceratops and Kosmoceratops; 'Ceratops' is therefore maintained as a nomen dubium. Cladistic analysis recovers Spiclypeus as the sister taxon to the clade Vagaceratops + Kosmoceratops, and appears transitional in the morphology of its epiparietals. The discovery of Spiclypeus adds to the poorly known dinosaur fauna of the Judith River Formation, and suggests faunal turnover within the formation.

The functional and palaeoecological implications of tooth morphology and wear for the megaherbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta, Canada.

Megaherbivorous dinosaurs were exceptionally diverse on the Late Cretaceous island continent of Laramidia, and a growing body of evidence suggests that this diversity was facilitated by dietary niche partitioning. We test this hypothesis using the fossil megaherbivore assemblage from the Dinosaur Park Formation (upper Campanian) of Alberta as a model. Comparative tooth morphology and wear, including the first use of quantitative dental microwear analysis in the context of Cretaceous palaeosynecology, are used to infer the mechanical properties of the foods these dinosaurs consumed. The phylliform teeth of ankylosaurs were poorly adapted for habitually processing high-fibre plant matter. Nevertheless, ankylosaur diets were likely more varied than traditionally assumed: the relatively large, bladed teeth of nodosaurids would have been better adapted to processing a tougher, more fibrous diet than the smaller, cusp-like teeth of ankylosaurids. Ankylosaur microwear is characterized by a preponderance of pits and scratches, akin to modern mixed feeders, but offers no support for interspecific dietary differences. The shearing tooth batteries of ceratopsids are much better adapted to high-fibre herbivory, attested by their scratch-dominated microwear signature. There is tentative microwear evidence to suggest differences in the feeding habits of centrosaurines and chasmosaurines, but statistical support is not significant. The tooth batteries of hadrosaurids were capable of both shearing and crushing functions, suggestive of a broad dietary range. Their microwear signal overlaps broadly with that of ankylosaurs, and suggests possible dietary differences between hadrosaurines and lambeosaurines. Tooth wear evidence further indicates that all forms considered here exhibited some degree of masticatory propaliny. Our findings reveal that tooth morphology and wear exhibit different, but complimentary, dietary signals that combine to support the hypothesis of dietary niche partitioning. The inferred mechanical and dietary patterns appear constant over the 1.5 Myr timespan of the Dinosaur Park Formation megaherbivore chronofauna, despite continual species turnover.

Skull ecomorphology of megaherbivorous dinosaurs from the dinosaur park formation (upper campanian) of Alberta, Canada.

Megaherbivorous dinosaur coexistence on the Late Cretaceous island continent of Laramidia has long puzzled researchers, owing to the mystery of how so many large herbivores (6-8 sympatric species, in many instances) could coexist on such a small (4-7 million km(2)) landmass. Various explanations have been put forth, one of which-dietary niche partitioning-forms the focus of this study. Here, we apply traditional morphometric methods to the skulls of megaherbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta to infer the ecomorphology of these animals and to test the niche partitioning hypothesis. We find evidence for niche partitioning not only among contemporaneous ankylosaurs, ceratopsids, and hadrosaurids, but also within these clades at the family and subfamily levels. Consubfamilial ceratopsids and hadrosaurids differ insignificantly in their inferred ecomorphologies, which may explain why they rarely overlap stratigraphically: interspecific competition prevented their coexistence.

Feeding height stratification among the herbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta, Canada.

BACKGROUND: Herbivore coexistence on the Late Cretaceous island continent of Laramidia has been a topic of great interest, stemming from the paradoxically high diversity and biomass of these animals in relation to the relatively small landmass available to them. Various hypotheses have been advanced to account for these facts, of which niche partitioning is among the most frequently invoked. However, despite its wide acceptance, this hypothesis has not been rigorously tested. This study uses the fossil assemblage from the Dinosaur Park Formation of Alberta as a model to investigate whether niche partitioning facilitated herbivorous dinosaur coexistence on Laramidia. Specifically, the question of feeding height stratification is examined in light of the role it plays in facilitating modern ungulate coexistence. RESULTS: Most herbivorous dinosaur species from the Dinosaur Park Formation were restricted to feeding no higher than approximately 1 m above the ground. There is minimal evidence for feeding height partitioning at this level, with ceratopsids capable of feeding slightly higher than ankylosaurs, but the ecological significance of this is ambiguous. Hadrosaurids were uniquely capable of feeding up to 2 m quadrupedally, or up to 5 m bipedally. There is no evidence for either feeding height stratification within any of these clades, or for change in these ecological relationships through the approximately 1.5 Ma record of the Dinosaur Park Formation. CONCLUSIONS: Although we cannot reject the possibility, we find no good evidence that feeding height stratification, as revealed by reconstructed maximum feeding heights, played an important role in facilitating niche partitioning among the herbivorous dinosaurs of Laramidia. Most browsing pressure was concentrated in the herb layer, although hadrosaurids were capable of reaching shrubs and low-growing trees that were out of reach from ceratopsids, ankylosaurs, and other small herbivores, effectively dividing the herbivores in terms of relative abundance. Sympatric hadrosaurids may have avoided competing with one another by feeding differentially using bipedal and quadrupedal postures. These ecological relationships evidently proved to be evolutionarily stable because they characterize the herbivore assemblage of the Dinosaur Park Formation through time. If niche partitioning served to facilitate the rich diversity of these animals, it may have been achieved by other means in addition to feeding height stratification. Consideration of other feeding height proxies, including dental microwear and skull morphology, may help to alleviate problems of underdetermination identified here.