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.

A giant tyrannosaur from the Campanian-Maastrichtian of southern North America and the evolution of tyrannosaurid gigantism.

Tyrannosaurid dinosaurs dominated as predators in the Late Cretaceous of Laurasia, culminating in the evolution of the giant Tyrannosaurus rex, both the last and largest tyrannosaurid. Where and when Tyrannosaurini (T. rex and kin) originated remains unclear. Competing hypotheses place tyrannosaurin origins in Asia, or western North America (Laramidia). We report a new tyrannosaurin, Tyrannosaurus mcraeensis, from the Campanian-Maastrichtian Hall Lake Formation of New Mexico, based on a fossil previously referred to T. rex. T. mcraeensis predates T. rex by ~ 6-7 million years, yet rivaled it in size. Phylogenetic analysis recovers T. mcraeensis as sister to T. rex and suggests Tyrannosaurini originated in southern Laramidia. Evolution of giant tyrannosaurs in southern North America, alongside giant ceratopsians, hadrosaurs, and titanosaurs suggests large-bodied dinosaurs evolved at low latitudes in North America.

Cranial anatomy of Allosaurus jimmadseni, a new species from the lower part of the Morrison Formation (Upper Jurassic) of Western North America.

Allosaurus is one of the best known theropod dinosaurs from the Jurassic and a crucial taxon in phylogenetic analyses. On the basis of an in-depth, firsthand study of the bulk of Allosaurus specimens housed in North American institutions, we describe here a new theropod dinosaur from the Upper Jurassic Morrison Formation of Western North America, Allosaurus jimmadseni sp. nov., based upon a remarkably complete articulated skeleton and skull and a second specimen with an articulated skull and associated skeleton. The present study also assigns several other specimens to this new species, Allosaurus jimmadseni, which is characterized by a number of autapomorphies present on the dermal skull roof and additional characters present in the postcrania. In particular, whereas the ventral margin of the jugal of Allosaurus fragilis has pronounced sigmoidal convexity, the ventral margin is virtually straight in Allosaurus jimmadseni. The paired nasals of Allosaurus jimmadseni possess bilateral, blade-like crests along the lateral margin, forming a pronounced nasolacrimal crest that is absent in Allosaurus fragilis.

A mid-Cretaceous tyrannosauroid and the origin of North American end-Cretaceous dinosaur assemblages.

Late Cretaceous dinosaur assemblages of North America-characterized by gigantic tyrannosaurid predators, and large-bodied herbivorous ceratopsids and hadrosaurids-were highly successful from around 80 million years ago (Ma) until the end of the 'Age of Dinosaurs' 66 Ma. However, the origin of these iconic faunas remains poorly understood because of a large, global sampling gap in the mid-Cretaceous, associated with an extreme sea-level rise. We describe the most complete skeleton of a predatory dinosaur from this gap, which belongs to a new tyrannosauroid theropod from the Middle Turonian (~92 Ma) of southern Laramidia (western North America). This taxon, Suskityrannus hazelae gen. et sp. nov., is a small-bodied species phylogenetically intermediate between the oldest, smallest tyrannosauroids and the gigantic, last-surviving tyrannosaurids. The species already possesses many key features of the tyrannosaurid bauplan, including the phylogenetically earliest record of an arctometatarsalian foot in tyrannosauroids, indicating that the group developed enhanced cursorial abilities at a small body size. Suskityrannus is part of a transitional Moreno Hill (that is, Zuni) dinosaur assemblage that includes dinosaur groups that became rare or were completely absent in North America around the final 15 Myr of the North American Cretaceous before the end-Cretaceous mass extinction, as well as small-bodied forebears of the large-bodied clades that dominated at this time.

Patterns of divergence in the morphology of ceratopsian dinosaurs: sympatry is not a driver of ornament evolution.

Establishing the origin and function of unusual traits in fossil taxa provides a crucial tool in understanding macroevolutionary patterns over long periods of time. Ceratopsian dinosaurs are known for their exaggerated and often elaborate horns and frills, which vary considerably between species. Many explanations have been proposed for the origin and evolution of these 'ornamental' traits, from predator defence to socio-sexual dominance signalling and, more recently, species recognition. A key prediction of the species recognition hypothesis is that two or more species possessing divergent ornamental traits should have been at least partially sympatric. For the first time to our knowledge, we test this hypothesis in ceratopsians by conducting a comparison of the morphological characters of 46 species. A total of 350 ceratopsian cladistic characters were categorized as either 'internal', 'display' (i.e. ornamental) or 'non display'. Patterns of diversity of these characters were evaluated across 1035 unique species pairs. Display characters were found to diverge rapidly overall, but sympatric species were not found to differ significantly in their ornamental disparity from non-sympatric species, regardless of phylogenetic distance. The prediction of the species recognition hypothesis, and thus the idea that ornamentation evolved as a species recognition mechanism, has no statistical support among known ceratopsians.

Microbial-caddisfly bioherm association from the Lower Cretaceous Shinekhudag Formation, Mongolia: Earliest record of plant armoring in fossil caddisfly cases.

Caddisfly larvae construct underwater protective cases using surrounding materials, thus providing information on environmental conditions in both modern and ancient systems. Microbial bioherms associated with caddisfly cases are found in the Berriassian-Hauterivian (~140-130 Ma) Shinekhudag Formation of Mongolia, and yield new insights into aspects of lacustrine paleoecosystems and paleoenvironments. This formation contains the earliest record of plant-armored caddisfly cases and a rare occurrence of microbial-caddisfly association from the Mesozoic. The bioherms are investigated within the context of stratigraphic correlations, depositional environment interpretations, and basin-evolution models of the sedimentary fill. The bioherms form 0.5-2.0 m diameter mound-shaped bodies and are concentrated within a single, oil shale-bound stratigraphic interval. Each bioherm is composed of up to 40% caddisfly cases along with stromatolites of millimeter-scale, micritic laminations. Petrographic analyses reveal these bioherms are composed of non-systematic associations of columnar and oncoidal microbialites, constructed around colonies of caddisfly cases. The cases are straight to curved, slightly tapered, and tube-shaped, with a progressively increasing length and width trend (7-21 mm by 1.5-2.5 mm). Despite these variations, the case architectures reveal similar construction materials; the particles used for cases are dominated by plant fragments, ostracod valves, carbonate rocks, and rare mica and feldspar grains. Allochems within the bioherms include ooids, ostracods, plant fragments, rare gastropods, feldspar grains bound in micritic matrices, and are consolidated by carbonate dominated cements. The combination of microbial-caddisfly association, plant fragment case particles, and ooids/oncoids are indicative of a shallow, littoral lake setting. Stratigraphic juxtaposition of nearshore bioherms and the bounding distal oil-shale facies suggests that the bioherms developed in an underfilled lake basin, resulting from an abrupt and short-lived lake desiccation event. Lake chemistry is believed to have been relatively alkaline, saline to hypersaline, and rich in Ca, Mg, and HCO3 ions. Through analyzing bioherm characteristics, caddisfly case architecture, carbonate microfacies, and stratigraphic variability, we infer larger-scale processes that controlled basin development during their formation.

A New Centrosaurine Ceratopsid, Machairoceratops cronusi gen et sp. nov., from the Upper Sand Member of the Wahweap Formation (Middle Campanian), Southern Utah.

The Upper Cretaceous (middle-late Campanian) Wahweap Formation of southern Utah contains the oldest diagnostic evidence of ceratopsids (to date, all centrosaurines) in North America, with a number of specimens recovered from throughout a unit that spans between 81 and 77 Ma. Only a single specimen has been formally named, Diabloceratops eatoni, from the lower middle member of the formation. Machairoceratops cronusi gen. et sp. nov., a new centrosaurine ceratopsid from the upper member of the Wahweap Formation, is here described based on cranial material representing a single individual recovered from a calcareous mudstone. The specimen consists of two curved and elongate orbital horncores, a left jugal, a nearly complete, slightly deformed braincase, the left squamosal, and a mostly complete parietal ornamented by posteriorly projected, anterodorsally curved, elongate spikes on either side of a midline embayment. The fan-shaped, stepped-squamosal is diagnostic of Centrosaurinae, however, this element differs from the rectangular squamosal in Diabloceratops. Machairoceratops also differs in the possession of two anterodorsally (rather than laterally) curved epiparietal ornamentations on either side of a midline embayment that are distinguished by a posteromedially-oriented sulcus along the entire length of the epiparietal. Additionally, the parietosquamosal frill is lacking any other epiossifications along its periphery. Machairoceratops shares a triangular (rather than round) frill and spike-like epiparietal loci (p1) ornamentation with the stratigraphically lower Diabloceratops. Both parsimony and Bayesian phylogenetic analyses place Machairoceratops as an early-branching centrosaurine. However, the parsimony-based analysis provides little resolution for the position of the new taxon, placing it in an unresolved polytomy with Diabloceratops. The resultant Bayesian topology yielded better resolution, aligning Machairoceratops as the definitive sister taxon to a clade formed by Diabloceratops and Albertaceratops. Considered together, both phylogenetic methods unequivocally place Machairoceratops as an early-branching centrosaurine, and given the biostratigraphic position of Machairoceratops, these details increase the known ceratopsid diversity from both the Wahweap Formation and the southern portion of Laramidia. Finally, the unique morphology of the parietal ornamentation highlights the evolutionary disparity of frill ornamentation near the base of Centrosaurinae.

A new chasmosaurine from northern Laramidia expands frill disparity in ceratopsid dinosaurs.

A new taxon of chasmosaurine ceratopsid demonstrates unexpected disparity in parietosquamosal frill shape among ceratopsid dinosaurs early in their evolutionary radiation. The new taxon is described based on two apomorphic squamosals collected from approximately time equivalent (approximately 77 million years old) sections of the upper Judith River Formation, Montana, and the lower Dinosaur Park Formation of Dinosaur Provincial Park, Alberta. It is referred to Chasmosaurinae based on the inferred elongate morphology. The typical chasmosaurine squamosal forms an obtuse triangle in dorsal view that tapers towards the posterolateral corner of the frill. In the dorsal view of the new taxon, the lateral margin of the squamosal is hatchet-shaped with the posterior portion modified into a constricted narrow bar that would have supported the lateral margin of a robust parietal. The new taxon represents the oldest chasmosaurine from Canada, and the first pre-Maastrichtian ceratopsid to have been collected on both sides of the Canada-US border, with a minimum north-south range of 380 km. This squamosal morphology would have given the frill of the new taxon a unique dorsal profile that represents evolutionary experimentation in frill signalling near the origin of chasmosaurine ceratopsids and reinforces biogeographic differences between northern and southern faunal provinces in the Campanian of North America.

The basal nodosaurid ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (lower Albian) Escucha Formation of northeastern Spain.

Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic recognized character that define nodosaurid ankylosaurs. These new specimens comprise a new genus and species of nodosaurid ankylosaur and represent the single most complete taxon of ankylosaur from the Cretaceous of Europe. These two specimens were examined and compared to all other known ankylosaurs. Comparisons of these specimens document that Europelta carbonensis n. gen., n. sp. is a nodosaur and is the sister taxon to the Late Cretaceous nodosaurids Anoplosaurus, Hungarosaurus, and Struthiosaurus, defining a monophyletic clade of European nodosaurids- the Struthiosaurinae.

Tyrant dinosaur evolution tracks the rise and fall of Late Cretaceous oceans.

The Late Cretaceous (∼95-66 million years ago) western North American landmass of Laramidia displayed heightened non-marine vertebrate diversity and intracontinental regionalism relative to other latest Cretaceous Laurasian ecosystems. Processes generating these patterns during this interval remain poorly understood despite their presumed role in the diversification of many clades. Tyrannosauridae, a clade of large-bodied theropod dinosaurs restricted to the Late Cretaceous of Laramidia and Asia, represents an ideal group for investigating Laramidian patterns of evolution. We use new tyrannosaurid discoveries from Utah--including a new taxon which represents the geologically oldest member of the clade--to investigate the evolution and biogeography of Tyrannosauridae. These data suggest a Laramidian origin for Tyrannosauridae, and implicate sea-level related controls in the isolation, diversification, and dispersal of this and many other Late Cretaceous vertebrate clades.

A remarkable short-snouted horned dinosaur from the Late Cretaceous (late Campanian) of southern Laramidia.

The fossil record of centrosaurine ceratopsids is largely restricted to the northern region of western North America (Alberta, Montana and Alaska). Exceptions consist of single taxa from Utah (Diabloceratops) and China (Sinoceratops), plus otherwise fragmentary remains from the southern Western Interior of North America. Here, we describe a remarkable new taxon, Nasutoceratops titusi n. gen. et sp., from the late Campanian Kaiparowits Formation of Utah, represented by multiple specimens, including a nearly complete skull and partial postcranial skeleton. Autapomorphies include an enlarged narial region, pneumatic nasal ornamentation, abbreviated snout and elongate, rostrolaterally directed supraorbital horncores. The subrectangular parietosquamosal frill is relatively unadorned and broadest in the mid-region. A phylogenetic analysis indicates that Nasutoceratops is the sister taxon to Avaceratops, and that a previously unknown subclade of centrosaurines branched off early in the group's history and persisted for several million years during the late Campanian. As the first well-represented southern centrosaurine comparable in age to the bulk of northern forms, Nasutoceratops provides strong support for the provincialism hypothesis, which posits that Laramidia-the western landmass formed by inundation of the central region of North America by the Western Interior Seaway-hosted at least two coeval dinosaur communities for over a million years of late Campanian time.

New horned dinosaurs from Utah provide evidence for intracontinental dinosaur endemism.

BACKGROUND: During much of the Late Cretaceous, a shallow, epeiric sea divided North America into eastern and western landmasses. The western landmass, known as Laramidia, although diminutive in size, witnessed a major evolutionary radiation of dinosaurs. Other than hadrosaurs (duck-billed dinosaurs), the most common dinosaurs were ceratopsids (large-bodied horned dinosaurs), currently known only from Laramidia and Asia. Remarkably, previous studies have postulated the occurrence of latitudinally arrayed dinosaur "provinces," or "biomes," on Laramidia. Yet this hypothesis has been challenged on multiple fronts and has remained poorly tested. METHODOLOGY/PRINCIPAL FINDINGS: Here we describe two new, co-occurring ceratopsids from the Upper Cretaceous Kaiparowits Formation of Utah that provide the strongest support to date for the dinosaur provincialism hypothesis. Both pertain to the clade of ceratopsids known as Chasmosaurinae, dramatically increasing representation of this group from the southern portion of the Western Interior Basin of North America. Utahceratops gettyi gen. et sp. nov.-characterized by short, rounded, laterally projecting supraorbital horncores and an elongate frill with a deep median embayment-is recovered as the sister taxon to Pentaceratops sternbergii from the late Campanian of New Mexico. Kosmoceratops richardsoni gen. et sp. nov.-characterized by elongate, laterally projecting supraorbital horncores and a short, broad frill adorned with ten well developed hooks-has the most ornate skull of any known dinosaur and is closely allied to Chasmosaurus irvinensis from the late Campanian of Alberta. CONCLUSIONS/SIGNIFICANCE: Considered in unison, the phylogenetic, stratigraphic, and biogeographic evidence documents distinct, co-occurring chasmosaurine taxa north and south on the diminutive landmass of Laramidia. The famous Triceratops and all other, more nested chasmosaurines are postulated as descendants of forms previously restricted to the southern portion of Laramidia. Results further suggest the presence of latitudinally arrayed evolutionary centers of endemism within chasmosaurine ceratopsids during the late Campanian, the first documented occurrence of intracontinental endemism within dinosaurs.

A new Basal sauropodomorph dinosaur from the Lower Jurassic Navajo sandstone of Southern Utah.

BACKGROUND: Basal sauropodomorphs, or 'prosauropods,' are a globally widespread paraphyletic assemblage of terrestrial herbivorous dinosaurs from the Late Triassic and Early Jurassic. In contrast to several other landmasses, the North American record of sauropodomorphs during this time interval remains sparse, limited to Early Jurassic occurrences of a single well-known taxon from eastern North America and several fragmentary specimens from western North America. METHODOLOGY/PRINCIPAL FINDINGS: On the basis of a partial skeleton, we describe here a new basal sauropodomorph dinosaur from the Lower Jurassic Navajo Sandstone of southern Utah, Seitaad ruessi gen. et sp. nov. The partially articulated skeleton of Seitaad was likely buried post-mortem in the base of a collapsed dune foreset. The new taxon is characterized by a plate-like medial process of the scapula, a prominent proximal expansion of the deltopectoral crest of the humerus, a strongly inclined distal articular surface of the radius, and a proximally and laterally hypertrophied proximal metacarpal I. CONCLUSIONS/SIGNIFICANCE: Phylogenetic analysis recovers Seitaad as a derived basal sauropodomorph closely related to plateosaurid or massospondylid 'prosauropods' and its presence in western North America is not unexpected for a member of this highly cosmopolitan clade. This occurrence represents one of the most complete vertebrate body fossil specimens yet recovered from the Navajo Sandstone and one of the few basal sauropodomorph taxa currently known from North America.