Parental feeding in the dinosaur Lufengosaurus revealed through multidisciplinary comparisons with altricial and precocious birds.

Previous studies arguing for parental care in dinosaurs have been primarily based on fossil accumulations of adults and hatchlings, perinatal and post-hatchlings in nests and nest areas, and evidence of brooding, the majority of which date to the Late Cretaceous. Similarly, the general body proportions of preserved embryonic skeletons of the much older Early Jurassic Massospondylus have been used to suggest that hatchlings were unable to forage for themselves. Here, we approach the question of parental care in dinosaurs by using a combined morphological, chemical, and biomechanical approach to compare early embryonic and hatchling bones of the Early Jurassic sauropodomorph Lufengosaurus with those of extant avian taxa with known levels of parental care. We compare femora, the main weight-bearing limb bone, at various embryonic and post-embryonic stages in a precocious and an altricial extant avian dinosaur with those of embryonic and hatchling Lufengosaurus, and find that the rate and degree of bone development in Lufengosaurus is closer to that of the highly altricial Columba (pigeon) than the precocious Gallus (chicken), providing strong support for the hypothesis that Lufengosaurus was fully altricial. We suggest that the limb bones of Lufengosaurus hatchlings were not strong enough to forage for themselves and would likely need parental feeding.

A Spanish saltasauroid titanosaur reveals Europe as a melting pot of endemic and immigrant sauropods in the Late Cretaceous.

A new lithostrotian titanosaur, Qunkasaura pintiquiniestra gen. et sp. nov., is described based on a single partial skeleton from the late Campanian-early Maastrichtian fossil-site of Lo Hueco (Cuenca, Spain). This new taxon is supported by an exclusive combination of characters that highlights strong convergences with members of the South American Aeolosaurini. Qunkasaura allows to reorganise the complex phylogenetic relationships of the increasingly diverse finicretaceous sauropods of Europe. Phylogenetic analyses places Qunkasaura within Saltasauridae and possibly Opisthocoelicaudiinae, together with Abditosaurus. A new clade is established, Lohuecosauria, including Saltasaurus, Lohuecotitan, their most recent common ancestor and all its descendants. Two distinct Ibero-Armorican Campanian-Maastrichtian saltasauroid lineages are recognised: (i) Lirainosaurinae that is exclusive from Europe, and (ii) a saltasaurid lineage with possible opisthocoelicaudiine affinities, with a Laurasian distribution. Lirainosaurinae was a relict lineage including possible dwarf forms that evolved in isolation after reaching Europe before the Late Cretaceous through the Apulian route. The occurrence of opisthocoelicaudiines in Europe may be the result of a Late Cretaceous interchange between Europe and Asia. No evidence of insular dwarfism is found in the Ibero-Armorican opisthocoelicaudiines suggesting that they may have been newcomers to the area that arrived before the 'Maastrichtian Dinosaur Turnover' in southwestern Europe.

Lost, hidden, broken, cut-estimating and interpreting the shapes and masses of damaged assemblages of plesiosaur gastroliths.

Background: Gastroliths are stones of uncertain purpose that are commonly found inside the rib cages of plesiosaur fossils worldwide. Gastroliths from four Alberta (Canada) plesiosaurs were studied to determine both their shapes and masses, and their mass fractions relative to body mass. One animal's set of gastroliths was 100% complete and fully visible, but the others showed varying degrees of loss, damage or obscuration, so estimations of their original states were needed. Methods: The studied animals were: Albertonectes vanderveldei, Fluvionectes sloanae, Nichollssaura borealis and Wapuskanectes betsynichollsae. The animals come from three different palaeoenvironments: open marine, near shore marine, and fluvial. Gastrolith shapes were classified as either xiphoid, cylindrical, discoidal or spherical based on observed and/or estimated dimensions. Although not all methods could be applied in all cases, gastrolith shapes and masses were estimated four different ways: (1) direct measurement and weighing of a subset and predicting the properties of the remaining obscured and hidden stones; (2) measuring triaxial ellipsoid dimensions of free stones to calculate volumes and multiplying by the mass density of chert; (3) measuring two visible triaxial dimensions of embedded stones, estimating the hidden third dimension three different ways, and then determining volumes and masses by calculation; and (4) predicting the density and mass of a densely packed cluster of small gastroliths using geometrical arguments. Results: Total gastrolith mass never exceeded 0.2% of body mass in any plesiosaur, and is consistent with the idea that the amounts of gastroliths recovered with plesiosaurs would be ineffective as ballast. The largest plesiosaur in the sample had the largest single gastrolith and total gastrolith mass increases with body size. The shape characteristics of the gastroliths were different for different environments, but compositionally they are dominated by black cherts. A possible common source for the gastroliths was identified for the two geographically close and near-contemporanous Nichollssaura and Wapuskanectes.

The dentition of the Late Jurassic dwarf sauropod Europasaurus holgeri from northern Germany: ontogeny, function, and implications for a rhamphotheca-like structure in Sauropoda.

The basal macronarian sauropod Europasaurus holgeri is known only from the Late Jurassic of the Langenberg Quarry near Goslar, Lower Saxony, Germany. Europasaurus has been identified as an insular dwarf and shows a clear resemblance to Camarasaurus and Giraffatitan. This study provides a detailed description of the dentition of Europasaurus based on an array of fossils outstanding in their abundance, variety of preservation, and ontogenetic range. Dental morphology for the replacement and functional dentitions, the tooth replacement pattern, and implications for food intake are described for the Europasaurus dentition, which is characterized by broad-crowned teeth. Characteristic features for Europasaurus are the presence of denticles on replacement teeth, the wrinkled enamel, and large wear facets both on the apex and on the carinae of the tooth crowns. The partially articulated skull SNHM-2207-R and isolated tooth rows DfmMh/FV 580.1 and DfmMh/FV 896.7 suggest the presence of strong connective tissue partially covering the teeth. This connective tissue would have provided stability and protection for the teeth. Evidence for this connective tissue include exposed tooth necks, in-situ teeth with strongly resorbed roots which no longer would have been connected to the jaw bone, and wrinkled enamel and its surface pattern. The same features can be observed in other sauropod taxa as well. We therefore suggest that eusauropods in general possessed this connective tissue structure, which may be an autapomorphy of the group. Possibly, this hypothetical structure is homologous to the rhamphotheca in birds and some non-avian theropods, which, however rarely, show such a close integration of keratinous tissue and teeth that we hypothesize here.

A lost tooth in the jungle: revisiting the sole dinosaurian record from northern Brazil.

The record of Mesozoic reptiles in the Northern region of Brazil is extremely limited, with the only definite occurrence consisting of two ziphodont teeth recovered from an oil well core in the municipality of Nova Olinda do Norte, Amazonas state, from strata of the Alter do Chão Formation (Cenomanian) of the Amazonas Basin. In this study, we aim to reevaluate MCT.R.514 using the most recent methodologies available for the identification of isolated theropod teeth. The cladistic analyses recovered MCT.R.514 as a possible metriacanthosaurid or non carcharodontosaurid allosauroid while the LDAs showed affinities between the specimen and Abelisauridae, Piatniktzysauridae, and Tyrannosauridae. MCT.R.514 was assigned as an abelisaurid due to its overall morphology that lacked metriacanthosaurid synapomorphies (i.e. spiraling mesial carina, strongly labially displaced distal carina, and well-marked mesiolingual longitudinal groove), while maintaining homoplastic features between both groups (i.e. irregular enamel texture, a lingually biconcave cross section, and a straight distal margin). This, together with a mostly Laurasian distribution of Metriacanthosauridae favors the assignment of MCT.R.514 as an abelisaurid as the most parsimonious hypothesis in this occasion. The identification of the specimen as an abelisaurid further expands the still scarce "Mid'' Cretaceous record of this clade in Brazil.

Modelling take-off moment arms in an ornithocheiraean pterosaur.

Take-off is a vital part of powered flight which likely constrains the size of birds, yet extinct pterosaurs are known to have reached far larger sizes. Three different hypothesised take-off motions (bipedal burst launching, bipedal countermotion launching, and quadrupedal launching) have been proposed as explanations for how pterosaurs became airborne and circumvented this proposed morphological limit. We have constructed a computational musculoskeletal model of a 5 m wingspan ornithocheiraean pterosaur, reconstructing thirty-four key muscles to estimate the muscle moment arms throughout the three hypothesised take-off motions. Range of motion constrained hypothetical kinematic sequences for bipedal and quadrupedal take-off motions were modelled after extant flying vertebrates. Across our simulations we did not find higher hindlimb moment arms for bipedal take-off motions or noticeably higher forelimb moment arms in the forelimb for quadrupedal take-off motions. Despite this, in all our models we found the muscles utilised in the quadrupedal take-off have the largest total launch applicable moment arms throughout the entire take-off sequences and for the take-off pose. This indicates the potential availability of higher leverage for a quadrupedal take-off than hypothesised bipedal motions in pterosaurs pending further examination of muscle forces.

The postcranial anatomy of Moschorhinus kitchingi (Therapsida: Therocephalia) from the Karoo Basin of South Africa.

Therocephalia are an important clade of non-mammalian therapsids that evolved a diverse array of morphotypes and body sizes throughout their evolutionary history. The postcranial anatomy of therocephalians has largely been overlooked, but remains important towards understanding aspects of their palaeobiology and phylogenetic relationships. Here, we provide the first postcranial description of the large akidnognathid eutherocephalian Moschorhinus kitchingi by examining multiple specimens from fossil collections in South Africa. We also compare the postcranial anatomy with previously described therocephalian postcranial material and provide an updated literature review to ensure a reliable foundation of comparison for future descriptive work. Moschorhinus shares all the postcranial features of eutherocephalians that differentiate them from early-diverging therocephalians, but is differentiated from other eutherocephalian taxa by aspects concerning the scapula, interclavicle, sternum, manus, and femur. The novel anatomical data from this contribution shows that Moschorhinus possessed a stocky bauplan with a particularly robust scapula, humerus, and femur. These attributes, coupled with the short and robust skull bearing enlarged conical canines imply that Moschorhinus was well equipped to grapple with and subdue prey items. Additionally, the combination of these attributes differ from those of similarly sized coeval gorgonopsians, which would have occupied a similar niche in late Permian ecosystems. Moreover, Moschorhinus was the only large carnivore known to have survived the Permo-Triassic mass extinction. Thus, the subtle but important postcranial differences may suggest a type of niche partitioning in the predator guild during the Permo-Triassic mass extinction interval.

Endocranial development in non-avian dinosaurs reveals an ontogenetic brain trajectory distinct from extant archosaurs.

Modern birds possess highly encephalized brains that evolved from non-avian dinosaurs. Evolutionary shifts in developmental timing, namely juvenilization of adult phenotypes, have been proposed as a driver of head evolution along the dinosaur-bird transition, including brain morphology. Testing this hypothesis requires a sufficient developmental sampling of brain morphology in non-avian dinosaurs. In this study, we harness brain endocasts of a postnatal growth series of the ornithischian dinosaur Psittacosaurus and several other immature and mature non-avian dinosaurs to investigate how evolutionary changes to brain development are implicated in the origin of the avian brain. Using three-dimensional characterization of neuroanatomical shape across archosaurian reptiles, we demonstrate that (i) the brain of non-avian dinosaurs underwent a distinct developmental trajectory compared to alligators and crown birds; (ii) ornithischian and non-avialan theropod dinosaurs shared a similar developmental trajectory, suggesting that their derived trajectory evolved in their common ancestor; and (iii) the evolutionary shift in developmental trajectories is partly consistent with paedomorphosis underlying overall brain shape evolution along the dinosaur-bird transition; however, the heterochronic signal is not uniform across time and neuroanatomical region suggesting a highly mosaic acquisition of the avian brain form.

Trophic evolution in ornithopod dinosaurs revealed by dental wear.

Ornithopod dinosaurs evolved numerous craniodental innovations related to herbivory. Nonetheless, the relationship between occlusion, tooth wear rate, and tooth replacement rate has been neglected. Here, we reconstruct tooth wear rates by measuring tooth replacement rates and tooth wear volumes, and document their dental microwear. We demonstrate that total tooth volume and rates of tooth wear increased steadily during ornithopod evolution, with deeply-nested taxa wearing up to 3360 mm3 of tooth volume/day. Increased wear resulted in asymmetric tooth crown formation with uneven von Ebner line increment width by the Late Jurassic, and in faster tooth replacement rates in multiple lineages by the mid-Cretaceous. Microwear displays a contrasting pattern, with decreasing complexity and pit percentages in deeply-nested and later-occurring taxa. We hypothesize that early ornithopods were browsers and/or frugivores but deeply nested iguanodontians were bulk-feeders, eating tougher, less nutritious plants; these trends correlate with increasing body mass and longer gut passage times.

Possible eucynodont (Synapsida: Cynodontia) tracks from a lacustrine facies in the Lower Jurassic Moenave Formation of southwestern Utah.

Eight fossil tetrapod footprints from lake-shore deposits in the Lower Jurassic Moenave Formation at the St. George Dinosaur Discovery Site (SGDS) in southwestern Utah cannot be assigned to the prevalent dinosaurian (Anomoepus, Eubrontes, Gigandipus, Grallator, Kayentapus) or crocodyliform (Batrachopus) ichnotaxa at the site. The tridactyl and tetradactyl footprints are incomplete, consisting of digit- and digit-tip-only imprints. Seven of the eight are likely pes prints; the remaining specimen is a possible manus print. The pes prints have digit imprint morphologies and similar anterior projections and divarication angles to those of Brasilichnium, an ichnotaxon found primarily in eolian paleoenvironments attributed to eucynodont synapsids. Although their incompleteness prevents clear referral to Brasilichnium, the SGDS tracks nevertheless suggest a eucynodont track maker and thus represent a rare, Early Mesozoic occurrence of such tracks outside of an eolian paleoenvironment.

The complexity of tracking stegosaurs and their gregarious behavior.

Stegosaur tracks were unknown until the identification of Deltapodus more than 20 years ago. Currently, the Iberian Peninsula, especially Teruel Province, is one of the areas globally with the most occurrences of these tracks. However, their identification, based on the global record, is problematic due to their similarities with sauropod tracks. A review of the largest number of analyzed Deltapodus tracks globally, including the holotype of D. ibericus and a description of new occurrences, has been carried out. Our research shows substantial morphological variations, but all the studied tracks can be considered D. ibericus based on the manus morphology and the morphometric data. These variations are related to substrate differences and/or different dynamic foot postures (possibly ontogenetically related) during locomotion, as evidenced by changes within the same trackway. We provide detailed comparisons via 3D modeling with sauropod tracks, and our data show that they generally have proportionally longer manus and wider pes because of the differences in the metapodial bones. The scarcity of stegosaur trackways in the fossil record has prevented the identification of gregarious behavior in this group of herbivorous dinosaurs. Two of the studied tracksites show evidence of this behavior, being the only examples among stegosaurs described thus far in the fossil record.

Geometric morphometric analysis of an ontogenetic cranial series of the Permian dicynodont Diictodon feliceps.

The Karoo Basin of South Africa is renowned for its abundance and diversity of therapsid fossils. Among the most ubiquitous and persistent of the Permian fauna is the small herbivorous dicynodont Diictodon feliceps. Intraspecific variation in Diictodon is historically confounding, and while ontogeny is frequently cited as a potential source of variation, observable developmental changes have never been calibrated. The present study revisits this issue, comparing three-dimensional landmark configurations of 82 Diictodon crania to investigate the association between shape, size and dimorphism. Beyond the statistically significant relationship between shape and allometry, our results determine the shape differences between juvenile and adult skulls of Diictodon, aligned with common craniofacial features documented in other tetrapod taxa. Functionally, these changes are attributed to development of the jaw musculature for feeding on larger, tougher plant matter during later ontogeny. Cranial morphological variation owing to sexual dimorphism is negligible, but distinct differences are noted in the allometric trajectories of each morphotype. A component of non-allometric variation cannot be accounted for, and we propose that this represents natural variation, rather than an artefact of taphonomic deformation.

Paleobiological implications of chevron pathology in the sauropodomorph Plateosaurus trossingensis from the Upper Triassic of SW Germany.

Paleopathology, the study of diseases and injuries from the fossil record, allows for a unique view into the life of prehistoric animals. Pathologies have nowadays been described in nearly all groups of fossil vertebrates, especially dinosaurs. Despite the large number of skeletons, pathologies had never been reported in the sauropodomorph Plateosaurus trossingensis. Here we describe the first pathologies of Plateosaurus using two individuals with pathologies in the chevrons of the tail, from the Upper Triassic of Trossingen, SW Germany. The two specimens each contain three consecutive pathological chevrons. Our results show that the pathologies were caused by external trauma in one individual and potentially tendinous trauma in the other. Healing of the lesions allowed survival of both animals. Using additional pathological specimens found in other collections and from multiple localities, we observe that 14.8% of all individuals of Plateosaurus contain pathologies within their chevrons, suggesting it was a vulnerable bone.

The first deep-snouted tyrannosaur from Upper Cretaceous Ganzhou City of southeastern China.

Tyrannosaurids were the most derived group of Tyrannosauroidea and are characterized by having two body plans: gracile, long-snouted and robust, deep-snouted skulls. Both groups lived sympatrically in central Asia. Here, we report a new deep-snouted tyrannosaurid, Asiatyrannus xui gen. et sp. nov., from the Upper Cretaceous of Ganzhou City, southeastern China, which has produced the large-bodied and long-snouted Qianzhousaurus. Based on histological analysis, the holotype of Asiatyrannus xui is not a somatically mature adult, but it already passed through the most rapid growth stages. Asiatyrannus is a small to medium-sized tyrannosaurine, with a skull length of 47.5 cm and an estimated total body length of 3.5-4 m; or around half the size of Qianzhousaurus and other large-bodied tyrannosaurines in similar growth stages. Asiatyrannus and Qianzhousaurus are sympatric tyrannosaurid genera in the Maastrichtian of southeastern China. Asiatyrannus differs from Qianzhousaurus in that it has a proportionally deeper snout, longer premaxilla, deeper maxilla, and deeper dentary, and the cornual process of the lacrimal is inflated without developing a discrete horn. The different skull proportions and body sizes suggest that Asiatyrannus and Qianzhousaurus likely had different feeding strategies and occupied different ecological niches.

Early Cretaceous troodontine troodontid (Dinosauria: Theropoda) from the Ohyamashimo Formation of Japan reveals the early evolution of Troodontinae.

A new troodontid dinosaur, Hypnovenator matsubaraetoheorum gen. et sp. nov., is described based on an articulated postcranial skeleton recovered from the fluvial deposits of the Albian Ohyamashimo Formation of the Sasayama Group in Tambasasayama City, Hyogo Prefecture, Japan. Hypnovenator is distinguished from other troodontids by four autapomorphies and a combination of additional features. Our phylogenetic analysis positions Hypnovenator as the oldest and one of the most basal troodontines, forming a clade with Gobivenator mongoliensis. The discovery of Hypnovenator suggests that small-bodied maniraptorans with a sleeping posture were common not only in environments with volcanic and eolian events or alluvial systems but also in fluvial systems. Geometric morphometric analysis of manual ungual phalanges shows that manual ungual phalanges I and III of Hypnovenator exhibit considerable morphological variation but are functionally similar, which differs from those of non-troodontine troodontids, reflecting the transition of manual motion within Troodontinae. Hypnovenator also has mosaic features in the pes related to cursoriality. This study reveals that asymmetrical arctometatarsus occurred by the Albian, and some morphological changes, such as shorter digit IV than digit III and non-ungual phalanges of digits III with roller joints and digit IV with weakly ginglymoid articulation, arose during the early Late Cretaceous.

Unraveling sauropod diversity in the Portezuelo Formation of Patagonia through a comprehensive analysis of new and existing material.

The Portezuelo Formation preserves an outstanding record of the upper Turonian - lower Coniacian. Despite the discovery of a significant quantity of sauropod fossil material from the formation, only two species have been formally described to date: Malarguesaurus florenciae and Futalognkosaurus dukei. Here we present new sauropod material mostly composed of non-articulated caudal vertebrae (MCF-PVPH 916 and 917) that belong to two titanosauriforms on the basis of the following features: anterior caudal vertebrae with procoelous-opisthoplatyan articulations, transverse processes that reach the posterior articular face of the centrum and neural spines with a transverse width of around 50% of their anteroposterior length; anterior and middle caudal vertebrae with the neural arch restricted to the anterior half of the centrum; middle caudal centra with circular cross-section. Phylogenetic analysis recovers the new material in close relation to Malarguesaurus within a monophyletic clade at the base of Somphospondyli. This clade shares large pedicel height with a vertical anterior border on the middle caudal vertebrae, a vertical orientation of the neural spines on the distalmost middle caudal vertebrae and proximalmost posterior caudal vertebrae, and subequal relative lengths of the proximal ulnar condylar processes. The specimens presented here are distinct not only from Futalognkosaurus, but also from other indeterminate titanosaurian remains from the same formation. However, there are no significant differences between the specimen MCF-PVPH 917 and Malarguesaurus, but there are differences between the posterior caudal vertebrae of MCF-PVPH 916 and Malarguesaurus, so they could be considered different species. Whilst we err on the side of caution in not naming new taxa here, the two specimens significantly expand what we know about sauropods in the Turonian-Coniacian ecosystems of Patagonia, which will continue to do so as more material is discovered.

A new stegosaur (Dinosauria: Ornithischia) from the Middle Jurassic of Gansu Province, China.

Stegosaurs are a minor but iconic clade of ornithischian dinosaurs, yet due to a poor fossil record, their early evolution is poorly understood. Here, we describe a new stegosaur, Baiyinosaurus baojiensis, gen. et sp. nov. from the Middle Jurassic Wangjiashan Formation of the Pingchuan District, Baiyin City, Gansu Province, China. The frontal of Baiyinosaurus possesses a unique characteristic among Stegosauria: it is wider than long and contributes to both the medial and anterior margins of the supratemporal fenestra. The character combinations of dorsal vertebrae of Baiyinosaurus are also different to other stegosaurs: its neural arches are not greatly elongated, its parapophyses are well developed, and its neural spines are axially expanded in lateral. The features of the frontal and vertebrae of Baiyinosaurus are reminiscent of basally branching thyreophorans, indicating that Baiyinosaurus is transitional in morphology between early thyreophorans and early-diverging stegosaurs. Systematic analysis shows that Baiyinosaurus is an early-diverging stegosaur.

Earliest evidence of avian primary feather moult.

Feather moulting is a crucial process in the avian life cycle, which evolved to maintain plumage functionality. However, moulting involves both energetic and functional costs. During moulting, plumage function temporarily decreases between the shedding of old feathers and the full growth of new ones. In flying taxa, a gradual and sequential replacement of flight feathers evolved to maintain aerodynamic capabilities during the moulting period. Little is known about the moult strategies of non-avian pennaraptoran dinosaurs and stem birds, before the emergence of crown lineage. Here, we report on two Early Cretaceous pygostylian birds from the Yixian Formation (125 mya), probably referable to Confuciusornithiformes, exhibiting morphological characteristics that suggest a gradual and sequential moult of wing flight feathers. Short primary feathers interpreted as immature are symmetrically present on both wings, as is typical among extant flying birds. Our survey of the enormous collection of the Tianyu Museum confirms previous findings that evidence of active moult in non-neornithine pennaraptorans is rare and likely indicates a moult cycle greater than one year. Documenting moult in Mesozoic feathered dinosaurs is critical for understanding their ecology, locomotor ability and the evolution of this important life-history process in birds.

Intraspecific variation in the pterosaur Rhamphorhynchus muensteri-implications for flight and socio-sexual signaling.

Pterosaurs were the first powered flying vertebrates, with a fossil record that stretches back to about 230 million years before present. Most species are only known from one to three specimens, which are most often fragmentary. However, Rhamphorhynchus muensteri is known from numerous excellent specimens, including multiple specimens with soft tissue preservation. As such, Rhamphorhynchus muensteri is one of the only pterosaurs amenable to analysis for intraspecific variation. It has been previously predicted that elements directly involved in the flight apparatus, such as those of the forelimb, will be more highly constrained in their proportions than other parts of the skeleton. We investigated the degree of variation seen in elements and body parts of Rhamphorhynchus, which represents the best model system among pterosaurs for testing these expectations of intraspecific variation. We recover evidence for high levels of constraint throughout the appendicular and axial elements (head, neck, torso, tail, forelimbs, hindlimbs), suggesting that all were important for flight. We further find that tail variation increases among the largest specimens, suggesting reduced constraint and/or stronger sexual selection on the tail in more mature individuals.

Lokiceratops rangiformis gen. et sp. nov. (Ceratopsidae: Centrosaurinae) from the Campanian Judith River Formation of Montana reveals rapid regional radiations and extreme endemism within centrosaurine dinosaurs.

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.