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Recent years have seen increasing scientific interest in whether neuron counts can act as correlates of diverse biological phenomena. Lately, Herculano-Houzel (2023) argued that fossil endocasts and comparative neurological data from extant sauropsids allow to reconstruct telencephalic neuron counts in Mesozoic dinosaurs and pterosaurs, which might act as proxies for behaviors and life history traits in these animals. According to this analysis, large theropods such as Tyrannosaurus rex were long-lived, exceptionally intelligent animals equipped with "macaque- or baboon-like cognition", whereas sauropods and most ornithischian dinosaurs would have displayed significantly smaller brains and an ectothermic physiology. Besides challenging established views on Mesozoic dinosaur biology, these claims raise questions on whether neuron count estimates could benefit research on fossil animals in general. Here, we address these findings by revisiting Herculano-Houzel's (2023) work, identifying several crucial shortcomings regarding analysis and interpretation. We present revised estimates of encephalization and telencephalic neuron counts in dinosaurs, which we derive from phylogenetically informed modeling and an amended dataset of endocranial measurements. For large-bodied theropods in particular, we recover significantly lower neuron counts than previously proposed. Furthermore, we review the suitability of neurological variables such as neuron numbers and relative brain size to predict cognitive complexity, metabolic rate and life history traits in dinosaurs, coming to the conclusion that they are flawed proxies for these biological phenomena. Instead of relying on such neurological estimates when reconstructing Mesozoic dinosaur biology, we argue that integrative studies are needed to approach this complex subject., (© 2024 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

The Late Cretaceous of western North America supported diverse dinosaur assemblages, though understanding patterns of dinosaur diversity, evolution, and extinction has been historically limited by unequal geographic and temporal sampling. In particular, the existence and extent of faunal endemism along the eastern coastal plain of Laramidia continues to generate debate, and finer scale regional patterns remain elusive. Here, we report a new centrosaurine ceratopsid, Lokiceratops rangiformis , from the lower portion of the McClelland Ferry Member of the Judith River Formation in the Kennedy Coulee region along the Canada-USA border. Dinosaurs from the same small geographic region, and from nearby, stratigraphically equivalent horizons of the lower Oldman Formation in Canada, reveal unprecedented ceratopsid richness, with four sympatric centrosaurine taxa and one chasmosaurine taxon. Phylogenetic results show that Lokiceratops , together with Albertaceratops and Medusaceratops , was part of a clade restricted to a small portion of northern Laramidia approximately 78 million years ago. This group, Albertaceratopsini, was one of multiple centrosaurine clades to undergo geographically restricted radiations, with Nasutuceratopsini restricted to the south and Centrosaurini and Pachyrostra restricted to the north. High regional endemism in centrosaurs is associated with, and may have been driven by, high speciation rates and diversity, with competition between dinosaurs limiting their geographic range. High speciation rates may in turn have been driven in part by sexual selection or latitudinally uneven climatic and floral gradients. The high endemism seen in centrosaurines and other dinosaurs implies that dinosaur diversity is underestimated and contrasts with the large geographic ranges seen in most extant mammalian megafauna., Competing Interests: Andrew A. Farke is an Academic Editor for PeerJ. Brock A. Sisson is owner of Fossilogic LLC, which produces cast replicas of Lokiceratops elements., (© 2024 Loewen et al.)

Caenagnathidae is a clade of derived, Late Cretaceous oviraptorosaurian theropods from Asia and North America. Because their remains are rare and often fragmentary, caenagnathid diversity is poorly understood. Anzu wyliei is the only caenagnathid species currently described from the late Maastrichtian Hell Creek Formation of the USA and is also among the largest and most completely preserved North American caenagnathids. Smaller, less complete caenagnathid material has long been known from the Hell Creek Formation, but it is unclear whether these are juvenile representatives of Anzu or if they represent distinct, unnamed taxa. Here, we describe a relatively small caenagnathid hindlimb from the Hell Creek Formation, and conduct osteohistological analysis to assess its maturity. Histological data and morphological differences from Anzu wyliei and other caenagnathids allow us to conclude that this specimen represents a new species of caenagnathid from the Hell Creek Formation, with a smaller adult body size than Anzu. This new taxon is also distinct from other small caenagnathid material previously described from the area, potentially indicating the coexistence of three distinct caenagnathid species in the Hell Creek Formation. These results show that caenagnathid diversity in the Hell Creek ecosystem has been underestimated., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2024 Atkins-Weltman et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

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., (© 2022 The Authors. The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)

The early centrosaurine ceratopsid, Wendiceratops pinhornensis, was discovered in Alberta, Canada in a medium density monodominant bonebed from the Oldman Formation (mid-Campanian, ~79 Ma). The bonebed contains abundant, well-preserved, adult-sized and some juvenile-sized postcranial material, allowing for the first description of a number of elements of the postcrania of this basal centrosaurine ceratopsid. The postcranial elements described are generally consistent with postcrania described for more derived centrosaurine taxa. However the rectangular-shaped distal terminus of the ischium previously considered to be an apomorphy of Wendiceratops. is shown to also be present in Medusaceratops, and thus may be a synapomorphy of basal centrsaurines. The bonebed represents a lag deposit within a mudstone-bearing overbank facies and contains individuals from multiple age classes. It contains over 95% ceratopsid remains, with all identifiable elements referable to Wendiceratops. The elements are completely disarticulated, but have undergone little weathering or abrasion (both Stage 0), although the ends of long bones and processes capped by cartilage in life frequently exhibit evidence of wet rot and breakage by hydrological reworking after decomposition. The taphonomy of the bonebed is consist with other monodominant centrosaurine bonebeds that have been interpreted as mass death assemblages preserving evidence of gregarious (herding) behavior. At approximately 79 million years old, the Wendiceratops bonebed is approximately two million years older than other ceratopsid bonebeds indicating that this bonebed is the oldest documented evidence of herding behavior in a ceratopsid., (© 2022 American Association for Anatomy.)

Despite the long history of research in the late Campanian Judith River Formation in northern Montana, most of the vertebrate fossils are represented by fragmentary remains, making precise taxonomic identifications difficult. Contrary to this, the partially contemporaneous Dinosaur Park Formation, Alberta, Canada is known for its tremendous fossil preservation, permitting rigorous studies of dinosaur diversity, evolution, and biostratigraphy. Hadrosaurids comprise one of the most abundant dinosaur clades in the Dinosaur Park Formation, but taxonomic affinities of hadrosaurid specimens remain poorly understood in the Judith River Formation. Corythosaurus is the most common hadrosaurid in the Dinosaur Park Formation and, to date, has been restricted to this formation. This study reports the first definitive Corythosaurus specimens from the Judith River Formation, which were discovered on two private ranches in northern Montana. The attribution of the most complete skeleton to Corythosaurus is indicated by: wide crest-snout angle, presence of premaxilla-nasal fontanelle, dorsoventrally expanded nasal, laterally exposed ophthalmic canal of the laterosphenoid, and tall neural spines. A second specimen preserves a large ilium that can be positively identified as Corythosaurus based on its associated skull, which is now in private hands. The specimens were recovered from the Coal Ridge Member of the Judith River Formation, which is approximately time equivalent to the Dinosaur Park Formation. Thus, the discovery of Corythosaurus in the Judith River Formation extends the biogeographic range of this genus and establishes a framework for future interformational biostratigraphic studies of Late Cretaceous dinosaur faunas in North America., (© 2022 American Association for Anatomy.)

Lambeosaurine hadrosaurids exhibited extreme modifications to the skull, where the premaxillae, nasals, and prefrontals were modified to form their iconic supracranial crests. This morphology contrasts with their sister group, Hadrosaurinae, which possessed the plesiomorphic arrangement of bones. Although studies have discussed differences between lambeosaurine and hadrosaurine skull morphology and ontogeny, there is little information detailing suture modifications through ontogeny and evolution. Suture morphology is of particular interest due to its correlation with the mechanical loading of the skull in extant vertebrates. We quantify and contrast the morphology of calvarial sutures in iguanodontians and ontogenetic series of Corythosaurus and Gryposaurus to test whether the evolution of lambeosaurine crests impacted the mechanical loading of the skull. We found that suture interdigitation (SI) increases through ontogeny in hadrosaurids, although this increase is more extreme in Corythosaurus than Gryposaurus, and overall suture complexity (i.e., overall shape) remained constant. Lambeosaurines also have higher SI than other iguanodontians, even in crestless juveniles, suggesting that increased sinuosity is unrelated to the structural support of the crest. Hadrosaurines and basal iguanodontians did not differ. Similarly, lambeosaurines have more complexly shaped sutures than hadrosaurines and basal iguanodontians, while the latter two groups do not differ. Taken together, these results suggest that lambeosaurine calvarial sutures are more interdigitated than other iguanodontians, and although suture sinuosity increased through ontogeny, the suture shape remained constant. These ontogenetic and evolutionary patterns suggest that increased suture complexity in lambeosaurines coincided with crest evolution, and corresponding modifications to their facial skeleton altered the distribution of stress while feeding., (© 2023 The Authors. Evolution & Development published by Wiley Periodicals LLC.)

Large theropod dinosaurs are often reconstructed with their marginal dentition exposed because of the enormous size of their teeth and their phylogenetic association to crocodylians. We tested this hypothesis using a multiproxy approach. Regressions of skull length and tooth size for a range of theropods and extant varanid lizards confirm that complete coverage of theropod dinosaur teeth with extraoral tissues (gingiva and labial scales) is both plausible and consistent with patterns observed in living ziphodont amniotes. Analyses of dental histology from crocodylians and theropod dinosaurs, including Tyrannosaurus rex , further indicate that the most likely condition was complete coverage of the marginal dentition with extraoral tissue when the mouth was closed. This changes our perceptions about the appearance and oral configuration of these iconic predators and has broad implications for our interpretations of other terrestrial animals with large teeth.

Ankylosaurid dinosaurs were heavily armoured herbivores with tails modified into club-like weapons. These tail clubs have widely been considered defensive adaptations wielded against predatory theropod dinosaurs. Here we argue instead that ankylosaurid tail clubs were sexually selected structures used primarily for intraspecific combat. We found pathological osteoderms (armour plates) in the holotype specimen of Zuul crurivastator , which are localized to the flanks in the hip region rather than distributed randomly across the body, consistent with injuries inflicted by lateral tail-swinging and ritualized combat. We failed to find convincing evidence for predation as a key selective pressure in the evolution of the tail club. High variation in tail club size through time, and delayed ontogenetic growth of the tail club further support the sexual selection hypothesis. There is little doubt that the tail club could have been used in defence when needed, but our results suggest that sexual selection drove the evolution of this impressive weapon. This changes the prevailing view of ankylosaurs, suggesting they were behaviorally complex animals that likely engaged in ritualized combat for social dominance as in other ornithischian dinosaurs and mammals.

The Late Cretaceous (Maastrichtian) Ruth Mason Dinosaur Quarry (RMDQ) represents a monodominant Edmontosaurus annectens bonebed from the Hell Creek Formation of South Dakota and has been determined as a catastrophic death assemblage likely belonging to a single population, providing an ideal sample to investigate hadrosaurid growth and population dynamics. For this study, size-frequency distributions were constructed from linear measurements of long bones (humeri, femora, tibiae) from RMDQ that revealed five relatively distinct size classes along a generally right-skewed distribution, which is consistent with a catastrophic assemblage. To test the relationship between morphological size ranges and ontogenetic age classes, subsets from each size-frequency peak were transversely thin-sectioned at mid-diaphysis to conduct an ontogenetic age assessment based on growth marks and observations of the bone microstructure. When combining these independent datasets, growth marks aligned with size-frequency peaks, with the exclusion of the overlapping subadult-adult size range, indicating a strong size-age relationship in early ontogeny. A growth curve analysis of tibiae indicated that E. annectens exhibited a similar growth trajectory to the Campanian hadrosaurid Maiasaura, although attaining a much larger asymptotic body size by about 9 years of age, further suggesting that the clade as a whole may have inherited a similar growth strategy. This rich new dataset for E. annectens provides new perspectives on other hypotheses of hadrosaurid life history. When the RMDQ population was compared with size distributions from other hadrosaurid bonebed assemblages, juveniles (categorized as ages one and two) were either completely absent from or heavily underrepresented in the samples, providing support for the hypothesized segregation between juvenile and adult hadrosaurids. Osteohistological comparison with material from polar and temperate populations of Edmontosaurus revealed that previous conclusions correlating osteohistological growth patterns with the strength of environmental stressors were a result of sampling non-overlapping ontogenetic growth stages., (© 2022 Anatomical Society.)

Other than repaired fractures, osteoarthritis, and periosteal reaction, the vertebrate fossil record has limited evidence of non-osseous diseases. This difficulty in paleontological diagnoses stems from (1) the inability to conduct medical testing, (2) soft-tissue pathologic structures are less likely to be preserved, and (3) many osseous lesions are not diagnostically specific. However, here reported for the first time is an avian-style respiratory disorder in a non-avian dinosaur. This sauropod presents irregular bony pathologic structures stemming from the pneumatic features in the cervical vertebrae. As sauropods show well-understood osteological correlates indicating that respiratory tissues were incorporated into the post-cranial skeleton, and thus likely had an 'avian-style' form of respiration, it is most parsimonious to identify these pathologic structures as stemming from a respiratory infection. Although several extant avian infections produce comparable symptoms, the most parsimonious is airsacculitis with associated osteomyelitis. From actinobacterial to fungal in origin, airsacculitis is an extremely prevalent respiratory disorder in birds today. While we cannot pinpoint the specific infectious agent that caused the airsacculitis, this diagnosis establishes the first fossil record of this disease. Additionally, it allows us increased insight into the medical disorders of dinosaurs from a phylogenetic perspective and understanding what maladies plagued the "fearfully great lizards"., (© 2022. The Author(s).)

The Wapiti Formation of northwest Alberta and northeast British Columbia, Canada, preserves an Upper Cretaceous terrestrial vertebrate fauna that is latitudinally situated between those documented further north in Alaska and those from southern Alberta and the contiguous U.S.A. Therefore, the Wapiti Formation is important for identifying broad patterns in vertebrate ecology, diversity, and distribution across Laramidia during the latest Cretaceous. Tracksites are especially useful as they provide a range of palaeoecological, palaeoenvironmental, and behavioural data that are complementary to the skeletal record. Here, we describe the Tyrants Aisle locality, the largest in-situ tracksite known from the Wapiti Formation. The site occurs in the lower part of Unit 4 of the formation (~72.5 Ma, upper Campanian), exposed along the southern bank of the Redwillow River. More than 100 tracks are documented across at least three distinct track-bearing layers, which were deposited on an alluvial floodplain. Hadrosaurid tracks are most abundant, and are referable to Hadrosauropodus based on track width exceeding track length, broad digits, and rounded or bilobed heel margins. We suggest the hadrosaurid trackmaker was Edmontosaurus regalis based on stratigraphic context. Tyrannosaurids, probable troodontids, possible ornithomimids, and possible azhdarchid pterosaurs represent minor but notable elements of the ichnofauna, as the latter is unknown from skeletal remains within the Wapiti Formation, and all others are poorly represented. Possible social behaviour is inferred for some of the hadrosaurid and small theropod-like trackmakers based on trackway alignment, suitable spacing and consistent preservation. On a broad taxonomic level (i.e., family or above), ichnofaunal compositions indicate that hadrosaurids were palaeoecologically dominant across Laramidia during the late Campanian within both high-and low-latitude deposits, although the role of depositional environment requires further testing., Competing Interests: The authors have declared that no competing interests exist.

Schroeder et al . (Reports, 26 February 2021, p. 941) reported a size gap among predatory dinosaur species. We argue that the supporting dataset is skewed toward Late Cretaceous North America and that the gap was likely absent during other intervals in most geographic regions. We urge broader consideration of this hypothesis, with quantitative evaluation of preservational and dataset biases.

During the latest Cretaceous, the European Archipelago was characterized by highly fragmented landmasses hosting putative dwarfed, insular dinosaurs, claimed as fossil evidence of the "island rule". The Villaggio del Pescatore quarry (north-eastern Italy) stands as the most informative locality within the palaeo-Mediterranean region and represents the first, multi-individual Konservat-Lagerstätte type dinosaur-bearing locality in Italy. The site is here critically re-evaluated as early Campanian in age, thus preceding the final fragmentation stages of the European Archipelago, including all other European localities preserving hypothesized dwarfed taxa. New skeletal remains allowed osteohistological analyses on the hadrosauroid Tethyshadros insularis indicating subadult features in the type specimen whereas a second, herein newly described, larger individual is likely somatically mature. A phylogenetic comparative framework places the body-size of T. insularis in range with other non-hadrosaurid Eurasian hadrosauroids, rejecting any significant evolutionary trend towards miniaturisation in this clade, confuting its 'pygmy' status, and providing unmatched data to infer environmentally-driven body-size trends in Mesozoic dinosaurs., (© 2021. The Author(s).)

Background: The origin of powered avian flight was a locomotor innovation that expanded the ecological potential of maniraptoran dinosaurs, leading to remarkable variation in modern birds (Neornithes). The avian sternum is the anchor for the major flight muscles and, despite varying widely in morphology, has not been extensively studied from evolutionary or functional perspectives. We quantify sternal variation across a broad phylogenetic scope of birds using 3D geometric morphometrics methods. Using this comprehensive dataset, we apply phylogenetically informed regression approaches to test hypotheses of sternum size allometry and the correlation of sternal shape with both size and locomotory capabilities, including flightlessness and the highly varying flight and swimming styles of Neornithes., Results: We find evidence for isometry of sternal size relative to body mass and document significant allometry of sternal shape alongside important correlations with locomotory capability, reflecting the effects of both body shape and musculoskeletal variation. Among these, we show that a large sternum with a deep or cranially projected sternal keel is necessary for powered flight in modern birds, that deeper sternal keels are correlated with slower but stronger flight, robust caudal sternal borders are associated with faster flapping styles, and that narrower sterna are associated with running abilities. Correlations between shape and locomotion are significant but show weak explanatory power, indicating that although sternal shape is broadly associated with locomotory ecology, other unexplored factors are also important., Conclusions: These results display the ecological importance of the avian sternum for flight and locomotion by providing a novel understanding of sternum form and function in Neornithes. Our study lays the groundwork for estimating the locomotory abilities of paravian dinosaurs, the ancestors to Neornithes, by highlighting the importance of this critical element for avian flight, and will be useful for future work on the origin of flight along the dinosaur-bird lineage., (© 2021. The Author(s).)

Inferring the body mass of fossil taxa, such as non-avian dinosaurs, provides a powerful tool for interpreting physiological and ecological properties, as well as the ability to study these traits through deep time and within a macroevolutionary context. As a result, over the past 100 years a number of studies advanced methods for estimating mass in dinosaurs and other extinct taxa. These methods can be categorized into two major approaches: volumetric-density (VD) and extant-scaling (ES). The former receives the most attention in non-avian dinosaurs and advanced appreciably over the last century: from initial physical scale models to three-dimensional (3D) virtual techniques that utilize scanned data obtained from entire skeletons. The ES approach is most commonly applied to extinct members of crown clades but some equations are proposed and utilized in non-avian dinosaurs. Because both approaches share a common goal, they are often viewed in opposition to one another. However, current palaeobiological research problems are often approach specific and, therefore, the decision to utilize a VD or ES approach is largely question dependent. In general, biomechanical and physiological studies benefit from the full-body reconstruction provided through a VD approach, whereas large-scale evolutionary and ecological studies require the extensive data sets afforded by an ES approach. This study summarizes both approaches to body mass estimation in stem-group taxa, specifically non-avian dinosaurs, and provides a comparative quantitative framework to reciprocally illuminate and corroborate VD and ES approaches. The results indicate that mass estimates are largely consistent between approaches: 73% of VD reconstructions occur within the expected 95% prediction intervals of the ES relationship. However, almost three quarters of outliers occur below the lower 95% prediction interval, indicating that VD mass estimates are, on average, lower than would be expected given their stylopodial circumferences. Inconsistencies (high residual and per cent prediction deviation values) are recovered to a varying degree among all major dinosaurian clades along with an overall tendency for larger deviations between approaches among small-bodied taxa. Nonetheless, our results indicate a strong corroboration between recent iterations of the VD approach based on 3D specimen scans suggesting that our current understanding of size in dinosaurs, and hence its biological correlates, has improved over time. We advance that VD and ES approaches have fundamentally (metrically) different advantages and, hence, the comparative framework used and advocated here combines the accuracy afforded by ES with the precision provided by VD and permits the rapid identification of discrepancies with the potential to open new areas of discussion., (© 2020 Cambridge Philosophical Society.)

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.

The holotype of the dromaeosaurid Saurornitholestes langstoni was described in 1978 on the basis of fewer than 30 associated cranial and postcranial bones of a single individual from Dinosaur Provincial Park. Four additional partial skeletons of Saurornitholestes were recovered from Campanian (Upper Cretaceous) beds of Alberta and Montana over the next 25 years, although reasonably complete skeletons remained elusive, and virtually nothing was known about the skull. The lack of truly diagnostic material has been problematic, and the relationships of Saurornitholestes to other dromaeosaurids have been difficult to resolve because of the incomplete knowledge of its anatomy. In 2014, an almost complete skeleton, including the skull, was collected less than a kilometer from where the holotype had been found. Although similar in body size to Velociraptor, the facial region of the skull is relatively shorter, taller, and wider. The nasals are pneumatic. The premaxillary teeth are distinctive, and teeth previously identified in the Dinosaur Park Formation as Zapsalis abradens can now be identified as the second premaxillary tooth of S. langstoni. Morphology and wear patterns suggest that these may have been specialized for preening feathers. Many traits define a Campanian North American clade, Saurornitholestinae, that is distinct from an Asian clade that includes Velociraptor (Velociraptorinae). This new information on the skull allows a more complete evaluation of its systematic position within the Dromaeosauridae and supports the suggestion of at least two major faunal interchanges between Asia and North America during the Cretaceous. Anat Rec, 303:691-715, 2020. © 2019 American Association for Anatomy., (© 2019 American Association for Anatomy.)

A new genus and species of non-hadrosaurid hadrosauroid, Gobihadros mongoliensis, is described from a virtually complete and undeformed skull and postcranial skeleton, as well as extensive referred material, collected from the Baynshire Formation (Cenomanian-Santonian) of the central and eastern Gobi Desert, Mongolia. Gobihadros mongoliensis is the first non-hadrosaurid hadrosauroid from the Late Cretaceous of central Asia known from a complete, articulated skull and skeleton. The material reveals the skeletal anatomy of a proximate sister taxon to Hadrosauridae in remarkable detail. Gobihadros is similar to Bactrosaurus johnsoni and Gilmoreosaurus mongoliensis, but can be distinguished from them in several autapomorphic traits, including the maximum number (three) of functional dentary teeth per tooth position, a premaxillary oral margin with a 'double-layer morphology', and a sigmoidal dorsal outline of the ilium with a well-developed, fan-shaped posterior process. All of these characters in Gobihadros are inferred to be convergent in Hadrosauridae. Phylogenetic analysis positions Gobihadros mongoliensis as a Bactrosaurus-grade hadrosauromorph hadrosauroid. Its relationship with Maastrichtian hadrosaurids from Asia (e.g., Saurolophus angustirostris, Kerberosaurus manakini, Wulagasaurus dongi, Kundurosaurus nagornyi) are sufficiently distant to indicate that these latter taxa owe their distribution to migration from North America across Beringia, rather than having a common Asian origin with Go. mongoliensis., Competing Interests: The authors have declared that no competing interests exist.

Sauropod dinosaurs were the largest terrestrial vertebrates; yet despite a robust global fossil record, the paucity of cranial remains complicates attempts to understand their paleobiology. An assemblage of small diplodocid sauropods from the Upper Jurassic Morrison Formation of Montana, USA, has produced the smallest diplodocid skull yet discovered. The ~24 cm long skull is referred to cf. Diplodocus based on the presence of several cranial and vertebral characters. This specimen enhances known features of early diplodocid ontogeny including a short snout with narrow-crowned teeth limited to the anterior portion of the jaws and more spatulate teeth posteriorly. The combination of size plus basal and derived character expression seen here further emphasizes caution when naming new taxa displaying the same, as these may be indicative of immaturity. This young diplodocid reveals that cranial modifications occurred throughout growth, providing evidence for ontogenetic dietary partitioning and recapitulation of ancestral morphologies.

The teeth of putatively carnivorous dinosaurs are often blade-shaped with well-defined serrated cutting edges (Figure 1). These ziphodont teeth are often easily differentiated based on the morphology and density of the denticles [1, 2]. A tearing function has been proposed for theropod denticles in general [3], but the functional significance of denticle phenotypic variation has received less attention. In particular, the unusual hooked denticles found in troodontids suggest a different feeding strategy or diet compared to other small theropods. We used a two-pronged approach to investigate the function of denticle shape variation across theropods with both congruent body shapes and sizes (e.g., dromaeosaurids versus troodontids) and highly disparate body shapes and sizes (e.g., troodontids versus tyrannosaurids), using microwear and finite element analyses (Figure 1). We found that many toothed coelurosaurian theropods employed a puncture-and-pull feeding movement, in which parallel scratches form while biting down into prey and oblique scratches form as the head is pulled backward with the jaws closed. In finite element simulations, theropod teeth had the lowest stresses when bite forces were aligned with the oblique family of microwear scratches. Different denticle morphologies performed differently under a variety of simulated biting angles: Dromaeosaurus and Saurornitholestes were well-adapted for handling struggling prey, whereas troodontid teeth were more likely to fail at non-optimal bite angles. Troodontids may have favored softer, smaller, or immobile prey., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

A partial skeleton from the Little Snowy Mountains of central Montana is the first referable specimen of the Morrison Formation macronarian sauropod Camarasaurus. This specimen also represents the northernmost occurrence of a sauropod in the Morrison. Histological study indicates that, although the specimen is relatively small statured, it is skeletally mature; this further emphasizes that size is not a undeviating proxy to maturity in dinosaurs, and that morphologies associated with an individual's age and stature may be more nebulous in sauropods.

Background: The Belly River Group of southern Alberta is one of the best-sampled Late Cretaceous terrestrial faunal assemblages in the world. This system provides a high-resolution biostratigraphic record of terrestrial vertebrate diversity and faunal turnover, and it has considerable potential to be a model system for testing hypotheses of dinosaur palaeoecological dynamics, including important aspects of palaeoecommunity structure, trophic interactions, and responses to environmental change. Vertebrate fossil microsites (assemblages of small bones and teeth concentrated together over a relatively short time and thought to be representative of community composition) offer an unparalleled dataset to better test these hypotheses by ameliorating problems of sample size, geography, and chronostratigraphic control that hamper other palaeoecological analyses. Here, we assembled a comprehensive relative abundance dataset of microsites sampled from the entire Belly River Group and performed a series of analyses to test the influence of environmental factors on site and taxon clustering, and assess the stability of faunal assemblages both temporally and spatially. We also test the long-held idea that populations of large dinosaur taxa were particularly sensitive to small-scale environmental gradients, such as the paralic (coastal) to alluvial (inland) regimes present within the time-equivalent depositional basin of the upper Oldman and lower Dinosaur Park Formations., Results: Palaeoenvironment (i.e. reconstructed environmental conditions, related to relative amount of alluvial, fluvial, and coastal influence in associated sedimentary strata) was found to be strongly associated with clustering of sites by relative-abundance faunal assemblages, particularly in relation to changes in faunal assemblage composition and marine-terrestrial environmental transitions. Palaeogeography/palaeolandscape were moderately associated to site relative abundance assemblage clustering, with depositional setting and time (i.e. vertical position within stratigraphic unit) more weakly associated. Interestingly, while vertebrate relative abundance assemblages as a whole were strongly correlated with these marine-terrestrial transitions, the dinosaur fauna does not appear to be particularly sensitive to them., Conclusions: This analysis confirms that depositional setting (i.e. the sediment type/sorting and associated characteristics) has little effect on faunal assemblage composition, in contrast to the effect of changes in the broader palaeoenvironment (e.g. upper vs. lower coastal plain, etc.), with marine-terrestrial transitions driving temporal faunal dynamics within the Belly River Group. The similarity of the dinosaur faunal assemblages between the time-equivalent portions of the Dinosaur Park Formation and Oldman Formation suggests that either these palaeoenvironments are more similar than characterized in the literature, or that the dinosaurs are less sensitive to variation in palaeoenvironment than has often been suggested. A lack of sensitivity to subtle environmental gradients casts doubt on these forces acting as a driver of putative endemism of dinosaur populations in the Late Cretaceous of North America.

Background: Hadrosaurid dinosaurs, dominant Late Cretaceous herbivores, possessed complex dental batteries with up to 300 teeth in each jaw ramus. Despite extensive interest in the adaptive significance of the dental battery, surprisingly little is known about how the battery evolved from the ancestral dinosaurian dentition, or how it functioned in the living organism. We undertook the first comprehensive, tissue-level study of dental ontogeny in hadrosaurids using several intact maxillary and dentary batteries and compared them to sections of other archosaurs and mammals. We used these comparisons to pinpoint shifts in the ancestral reptilian pattern of tooth ontogeny that allowed hadrosaurids to form complex dental batteries., Results: Comparisons of hadrosaurid dental ontogeny with that of other amniotes reveals that the ability to halt normal tooth replacement and functionalize the tooth root into the occlusal surface was key to the evolution of dental batteries. The retention of older generations of teeth was driven by acceleration in the timing and rate of dental tissue formation. The hadrosaurid dental battery is a highly modified form of the typical dinosaurian gomphosis with a unique tooth-to-tooth attachment that permitted constant and perfectly timed tooth eruption along the whole battery., Conclusions: We demonstrate that each battery was a highly dynamic, integrated matrix of living replacement and, remarkably, dead grinding teeth connected by a network of ligaments that permitted fine scale flexibility within the battery. The hadrosaurid dental battery, the most complex in vertebrate evolution, conforms to a surprisingly simple evolutionary model in which ancestral reptilian tissue types were redeployed in a unique manner. The hadrosaurid dental battery thus allows us to follow in great detail the development and extended life history of a particularly complex food processing system, providing novel insights into how tooth development can be altered to produce complex dentitions, the likes of which do not exist in any living vertebrate.

The causes, rate, and selectivity of the end-Cretaceous mass extinction continue to be highly debated [1-5]. Extinction patterns in small, feathered maniraptoran dinosaurs (including birds) are important for understanding extant biodiversity and present an enigma considering the survival of crown group birds (Neornithes) and the extinction of their close kin across the end-Cretaceous boundary [6]. Because of the patchy Cretaceous fossil record of small maniraptorans [7-12], this important transition has not been closely examined in this group. Here, we test the hypothesis that morphological disparity in bird-like dinosaurs was decreasing leading up to the end-Cretaceous mass extinction, as has been hypothesized in some dinosaurs [13, 14]. To test this, we examined tooth morphology, an ecological indicator in fossil reptiles [15-19], from over 3,100 maniraptoran teeth from four groups (Troodontidae, Dromaeosauridae, Richardoestesia, and cf. Aves) across the last 18 million years of the Cretaceous. We demonstrate that tooth disparity, a proxy for variation in feeding ecology, shows no significant decline leading up to the extinction event within any of the groups. Tooth morphospace occupation also remains static over this time interval except for increased size during the early Maastrichtian. Our data provide strong support that extinction within this group occurred suddenly after a prolonged period of ecological stability. To explain this sudden extinction of toothed maniraptorans and the survival of Neornithes, we propose that diet may have been an extinction filter and suggest that granivory associated with an edentulous beak was a key ecological trait in the survival of some lineages., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

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 sutures of the skulls of vertebrates are generally open early in life and slowly close as maturity is attained. The assumption that all vertebrates follow this pattern of progressive sutural closure has been used to assess maturity in the fossil remains of non-avian dinosaurs. Here, we test this assumption in two members of the Extant Phylogenetic Bracket of the Dinosauria, the emu, Dromaius novaehollandiae and the American alligator, Alligator mississippiensis, by investigating the sequence and timing of sutural fusion in their skulls. As expected, almost all the sutures in the emu skull progressively close (i.e., they get narrower) and then obliterate during ontogeny. However, in the American alligator, only two sutures out of 36 obliterate completely and they do so during embryonic development. Surprisingly, as maturity progresses, many sutures of alligators become wider in large individuals compared to younger, smaller individuals. Histological and histomorphometric analyses on two sutures and one synchondrosis in an ontogenetic series of American alligator confirmed our morphological observations. This pattern of sutural widening might reflect feeding biomechanics and dietary changes through ontogeny. Our findings show that progressive sutural closure is not always observed in extant archosaurs, and therefore suggest that cranial sutural fusion is an ambiguous proxy for assessing maturity in non-avian dinosaurs.

Tooth morphology and development can provide valuable insights into the feeding behaviour and evolution of extinct organisms. The teeth of Theropoda, the only clade of predominantly predatory dinosaurs, are characterized by ziphodonty, the presence of serrations (denticles) on their cutting edges. Known today only in varanid lizards, ziphodonty is much more pervasive in the fossil record. Here we present the first model for the development of ziphodont teeth in theropods through histological, SEM, and SR-FTIR analyses, revealing that structures previously hypothesized to prevent tooth breakage instead first evolved to shape and maintain the characteristic denticles through the life of the tooth. We show that this novel complex of dental morphology and tissues characterizes Theropoda, with the exception of species with modified feeding behaviours, suggesting that these characters are important for facilitating the hypercarnivorous diet of most theropods. This adaptation may have played an important role in the initial radiation and subsequent success of theropods as terrestrial apex predators.

The fossil record of ceratopsid dinosaurs between the occurrence of their proximate sister taxa in the Turonian and the beginning of their well-documented radiation from the late Campanian of North America onwards (approximately 90 and 77 Ma) is poor, with only seven taxa described from this early period in their evolution. We describe a new taxon of a highly adorned basal centrosaurine, Wendiceratops pinhornensis gen. et sp. nov., from the lower part of the Oldman Formation (middle Campanian, approximately 78-79 Ma), Alberta, Canada. Over 200 bones derived from virtually all parts of the skeleton, including multiple well-preserved specimens of the diagnostic parietosquamosal frill, were collected from a medium-density monodominant bonebed, making the new taxon one of the best-represented early ceratopsids. The new taxon is apomorphic in having epiparietals at loci 2 and 3 developed as broad-based, pachyostotic processes that are strongly procurved anterodorsally to overhang the posterior and lateral parietal rami, and an ischium with a broad, rectangular distal terminus. Although the morphology of the nasal is incompletely known, Wendiceratops is inferred to have a large, upright nasal horn located close to the orbits, which represents the oldest occurrence of this feature in Ceratopsia. Given the phylogenetic position of the new taxon within Centrosaurinae, a enlarged nasal horn is hypothesized to have arisen independently at least twice in ceratopsid evolution.

The anterior cervical vertebrae form the skeletal connection between the cranial and postcranial skeletons in higher tetrapods. As a result, the morphology of the atlas-axis complex is likely to be shaped by selection pressures acting on either the head or neck. The neoceratopsian (Reptilia:Dinosauria) syncervical represents one of the most highly modified atlas-axis regions in vertebrates, being formed by the complete coalescence of the three most anterior cervical vertebrae. In ceratopsids, the syncervical has been hypothesized to be an adaptation to support a massive skull, or to act as a buttress during intraspecific head-to-head combat. Here, we test these functional/adaptive hypotheses within a phylogenetic framework and critically examine the previously proposed methods for quantifying relative head size in the fossil record for the first time. Results indicate that neither the evolution of cranial weaponry nor large head size correlates with the origin of cervical fusion in ceratopsians, and we, therefore, reject both adaptive hypotheses for the origin of the syncervical. Anterior cervical fusion has evolved independently in a number of amniote clades, and further research on extant groups with this peculiar anatomy is needed to understand the evolutionary basis for cervical fusion in Neoceratopsia., (© 2015 The Author(s). Evolution © 2015 The Society for the Study of Evolution.)

Spectacularly preserved non-avian dinosaurs with integumentary filaments/feathers have revolutionized dinosaur studies and fostered the suggestion that the dinosaur common ancestor possessed complex integumentary structures homologous to feathers. This hypothesis has major implications for interpreting dinosaur biology, but has not been tested rigorously. Using a comprehensive database of dinosaur skin traces, we apply maximum-likelihood methods to reconstruct the phylogenetic distribution of epidermal structures and interpret their evolutionary history. Most of these analyses find no compelling evidence for the appearance of protofeathers in the dinosaur common ancestor and scales are usually recovered as the plesiomorphic state, but results are sensitive to the outgroup condition in pterosaurs. Rare occurrences of ornithischian filamentous integument might represent independent acquisitions of novel epidermal structures that are not homologous with theropod feathers., (© 2015 The Author(s) Published by the Royal Society. All rights reserved.)

The evolution of avian feathers has recently been illuminated by fossils and the identification of genes involved in feather patterning and morphogenesis. However, molecular studies have focused mainly on protein-coding genes. Using comparative genomics and more than 600,000 conserved regulatory elements, we show that patterns of genome evolution in the vicinity of feather genes are consistent with a major role for regulatory innovation in the evolution of feathers. Rates of innovation at feather regulatory elements exhibit an extended period of innovation with peaks in the ancestors of amniotes and archosaurs. We estimate that 86% of such regulatory elements and 100% of the nonkeratin feather gene set were present prior to the origin of Dinosauria. On the branch leading to modern birds, we detect a strong signal of regulatory innovation near insulin-like growth factor binding protein (IGFBP) 2 and IGFBP5, which have roles in body size reduction, and may represent a genomic signature for the miniaturization of dinosaurian body size preceding the origin of flight., (© The Author 2014. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.)

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 bones from single individuals. These analyses do not take individual variation into account, and as a result may lead to misleading inferences of the physiology of extinct organisms. This study uses a multi-element, multi-individual sampling of ornithomimid dinosaurs to test the amount of individual variation in the aforementioned osteohistological indicators. Based on these results we also assess the conclusions of previous studies that tested paleohistological hypotheses using isolated elements., Results: LAG number was found to be consistent within the hind limb bones of each individual, with the exception of the fibula, which preserves one additional LAG. Considerable differences in LAG spacing were found between elements of the sampled individuals, with larger variation found in elements of the foot compared with the femur, fibula, and tibia. Osteocyte lacunar density ranged between 29000 and 42000 osteocyte lacunae per mm(3), and was found to vary more between hind limb bones of an individual and within bones, than between the average values of individuals., Conclusions: The variation between hind limb elements in LAG number and LAG spacing suggests that direct comparisons of these elements may be misleading, and that LAG spacing is not a reliable proxy for mass-specific growth rates of an individual. Sampling of multiple bones should be performed as an internal check of model-based LAG retro-calculation and growth equations. The observation that osteocyte lacunar density varies more between individual bone elements than between average individual values suggests that the choice of sampled element can greatly influence the result, and care should be taken to not bias interpretations of the physiology of fossil tetrapods.

Large-scale adaptive radiations might explain the runaway success of a minority of extant vertebrate clades. This hypothesis predicts, among other things, rapid rates of morphological evolution during the early history of major groups, as lineages invade disparate ecological niches. However, few studies of adaptive radiation have included deep time data, so the links between extant diversity and major extinct radiations are unclear. The intensively studied Mesozoic dinosaur record provides a model system for such investigation, representing an ecologically diverse group that dominated terrestrial ecosystems for 170 million years. Furthermore, with 10,000 species, extant dinosaurs (birds) are the most speciose living tetrapod clade. We assembled composite trees of 614-622 Mesozoic dinosaurs/birds, and a comprehensive body mass dataset using the scaling relationship of limb bone robustness. Maximum-likelihood modelling and the node height test reveal rapid evolutionary rates and a predominance of rapid shifts among size classes in early (Triassic) dinosaurs. This indicates an early burst niche-filling pattern and contrasts with previous studies that favoured gradualistic rates. Subsequently, rates declined in most lineages, which rarely exploited new ecological niches. However, feathered maniraptoran dinosaurs (including Mesozoic birds) sustained rapid evolution from at least the Middle Jurassic, suggesting that these taxa evaded the effects of niche saturation. This indicates that a long evolutionary history of continuing ecological innovation paved the way for a second great radiation of dinosaurs, in birds. We therefore demonstrate links between the predominantly extinct deep time adaptive radiation of non-avian dinosaurs and the phenomenal diversification of birds, via continuing rapid rates of evolution along the phylogenetic stem lineage. This raises the possibility that the uneven distribution of biodiversity results not just from large-scale extrapolation of the process of adaptive radiation in a few extant clades, but also from the maintenance of evolvability on vast time scales across the history of life, in key lineages., Competing Interests: The authors have declared that no competing interests exist.

Dromaeosaurids from the Maastrichtian of North America have a poor fossil record and are known largely from isolated teeth, which have typically been referred to taxa based on more complete material from earlier Campanian strata. An almost complete maxilla with well-preserved dentition and an associated dentary from the Hell Creek Formation of Montana are used to establish a new dromaeosaurid taxon in the latest Maastrichtian, immediately prior to the end-Cretaceous extinction event. Acheroraptor temertyorum gen. et sp. nov. is differentiated from other dromaeosaurids on the basis of a hypertrophied postantral wall that projects posteriorly into the antorbital fenestra, a maxillary fenestra positioned low in the antorbital fossa and directly posterior to the promaxillary fenestra, and distinctive dentition with marked apicobasal ridges. The new material allows a dromaeosaurid from the Maastrichtian of North America to be placed within a phylogenetic framework for the first time. Phylogenetic analysis suggests Acheroraptor is a velociraptorine that is more closely related to Asian dromaeosaurids, including Tsaagan and Velociraptor, than it is to Dromaeosaurus, Saurornitholestes, or any other taxon from North America. As part of the Lancian Tyrannosaurus-Triceratops fauna, A. temertyorum is the latest occurring dromaeosaurid. Its relationships and occurrence suggest a complex historical biogeographic scenario that involved multiple, bi-directional faunal interchanges between Asia and North America during the Late Cretaceous.

Taphonomic biases dictate how organisms are represented in the fossil record, but their effect on studies of vertebrate diversity dynamics is poorly studied. In contrast to the high diversity and abundance of small-bodied animals in extant ecosystems, small-bodied dinosaurs are less common than their large-bodied counterparts, but it is unclear whether this reflects unique properties of dinosaurian ecosystems or relates to taphonomic biases. A new, fully domed pachycephalosaurid dinosaur, Acrotholus audeti, from the Santonian of Alberta predates incompletely domed taxa, and provides important new information on pachycephalosaur evolution and the completeness of the ornithischian fossil record. Here we provide the first empirical evidence that the diversity of small-bodied ornithischian dinosaurs is strongly underestimated based on ghost lineages and the high proportion of robust and diagnostic frontoparietal domes compared with other pachycephalosaur fossils. This suggests preservational biases have a confounding role in attempts to decipher vertebrate palaeoecology and diversity dynamics through the Mesozoic.

Brain and nasal cavity endocasts of four corythosaurian lambeosaurines (Dinosauria: Ornithischia) were investigated to test hypotheses of cranial crest function related to sensorineural systems. Endocasts were generated through computed tomography and three-dimensional rendering and visualization software. The sample comprises a range of ontogenetic stages from the taxa Lambeosaurus, Corythosaurus, and Hypacrosaurus. Results show that the morphology of brain endocasts differs little from that of hadrosaurines. The strikingly convoluted nasal vestibule of Hypacrosaurus altispinus, when interpreted in the context of lambeosaurine phylogeny, suggests selective pressure for nasal cavity function independent from changes in the external shape of the crest and associated visual display function. The plesiomorphically small olfactory bulbs and apparently small olfactory region of the nasal cavity argues against the hypothesis that increased olfactory acuity played a causal role in crest evolution. The elongate cochlea of the inner ear reveals that hearing in lambeosaurines emphasized low frequencies consistent with the hypothesized low-frequency calls made by the crests under the resonation model of crest function. The brain is relatively large in lambeosaurines compared with many other large dinosaurs, and the cerebrum is relatively larger than that of all non-hadrosaurian ornithischians and large theropods, but compares favorably with hadrosaurine hadrosaurids as well as some maniraptoran theropods. It is concluded that the large brains of lambeosaurines are consistent with the range of social behaviors inferred when the crest is interpreted as an intraspecific signaling structure., ((c) 2009 Wiley-Liss, Inc.)