Historically, ecomorphological inferences regarding theropod (i.e. 'predatory') dinosaurs were guided by an assumption that they were singularly hypercarnivorous. A recent plethora of maniraptoran discoveries has produced evidence challenging this notion. Here, we report on a new species of maniraptoran theropod, Nothronychus graffami sp. nov. Relative completeness of this specimen permits a phylogenetic reassessment of Therizinosauria-the theropod clade exhibiting the most substantial anatomical evidence of herbivory. In the most comprehensive phylogenetic study of the clade conducted to date, we recover Therizinosauria as the basalmost maniraptoran lineage. Using concentrated changes tests, we present evidence for correlated character evolution among herbivorous and hypercarnivorous taxa and propose ecomorphological indicators for future interpretations of diet among maniraptoran clades. Maximum parsimony optimizations of character evolution within our study indicate an ancestral origin for dietary plasticity and facultative herbivory (omnivory) within the clade. These findings suggest that hypercarnivory in paravian dinosaurs is a secondarily derived dietary specialization and provide a potential mechanism for the invasion of novel morpho-and ecospace early in coelurosaurian evolution-the loss of obligate carnivory and origin of dietary opportunism.

BackgroundDuring 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 FindingsHere 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/SignificanceConsidered 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.

BackgroundTroodontids are a predominantly small-bodied group of feathered theropod dinosaurs notable for their close evolutionary relationship with Avialae. Despite a diverse Asian representation with remarkable growth in recent years, the North American record of the clade remains poor, with only one controversial species—Troodon formosus—presently known from substantial skeletal remains.Methodology/Principal FindingsHere we report a gracile new troodontid theropod—Talos sampsoni gen. et sp. nov.—from the Upper Cretaceous Kaiparowits Formation, Utah, USA, representing one of the most complete troodontid skeletons described from North America to date. Histological assessment of the holotype specimen indicates that the adult body size of Talos was notably smaller than that of the contemporary genus Troodon. Phylogenetic analysis recovers Talos as a member of a derived, latest Cretaceous subclade, minimally containing Troodon, Saurornithoides, and Zanabazar. MicroCT scans reveal extreme pathological remodeling on pedal phalanx II-1 of the holotype specimen likely resulting from physical trauma and subsequent infectious processes.Conclusion/Significance Talos sampsoni adds to the singularity of the Kaiparowits Formation dinosaur fauna, which is represented by at least 10 previously unrecognized species including the recently named ceratopsids Utahceratops and Kosmoceratops, the hadrosaurine Gryposaurus monumentensis, the tyrannosaurid Teratophoneus, and the oviraptorosaurian Hagryphus. The presence of a distinct troodontid taxon in the Kaiparowits Formation supports the hypothesis that late Campanian dinosaurs of the Western Interior Basin exhibited restricted geographic ranges and suggests that the taxonomic diversity of Late Cretaceous troodontids from North America is currently underestimated. An apparent traumatic injury to the foot of Talos with evidence of subsequent healing sheds new light on the paleobiology of deinonychosaurians by bolstering functional interpretations of prey grappling and/or intraspecific combat for the second pedal digit, and supporting trackway evidence indicating a minimal role in weight bearing.

Recent fieldwork in the Tropic Shale of southern Utah has resulted in the recovery of several specimens of short-necked plesiosaurs including both polycotylid plesiosauroids and pliosaurids. This report focuses on the former, whereas the latter are discussed in an accompanying paper. Although the Late Cretaceous Cenomanian/Turonian Stage boundary falls within the lower few meters of the Tropic Shale, all but one of the specimens were found in lower Turonian strata based on molluscan assemblages found in direct association with the skeletal material. The plesiosauroid family Polycotylidae is represented by at least three taxa: (1) the 'typical' late Cenomanian/early Turonian Trinacromerum; (2) Eopolycotylus rankini, gen. et sp. nov., a new genus and species that shares synapomorphies with the late Santonian/early Campanian Polycotylus latipinnus; and (3) Palmula quadratus, gen. et sp. nov., another new taxon that shares synapomorphies with unnamed taxa from the late Cenomanian of South Dakota and Japan. Cladistic analysis supports the division of the Polycotylidae into two new subfamilies, here designated the Polycotylinae and the Palmulainae. The Turonian plesiosaur fauna of the Tropic Shale, and consequently the large vertebrate fauna of the Cretaceous Western Interior Seaway, is considerably more diverse than previously realized, and there is no indication that this fauna suffered any negative consequences as a result of global scale oceanographic events, including marine extinctions, that transpired during late Cenomanian-early Turonian time.

Fluctuations in depositional conditions during the onset of severe climate events in Earth history predispose stratigraphic archives to hiatuses, often hindering complete reconstructions of paleoclimate events and their triggers. Several studies have proposed that a hiatus of unknown duration exists at the base of Oceanic Anoxic Event 2 (OAE2) in the North American Western Interior Basin at the base Turonian global boundary stratotype section and point (GSSP) in Pueblo, Colorado, which potentially influences integrated radioisotopic, biostratigraphic, and astrochronologic age models of the Cenomanian-Turonian boundary interval. To quantify the duration of this hiatus, refine the chronology of OAE2, and assess marine geochemical perturbations associated with the onset of the event, we present new 40Ar/39Ar dates from regional bentonites along with a new proximal-distal chemostratigraphic transect of the epeiric Western Interior Basin (WIB), including initial osmium isotope (Osi) and stable carbon isotope (δ13C) data. The new 40Ar/39Ar age determinations confirm and further constrain previous estimates of Cenomanian-Turonian boundary timing. Further, the regional chemostratigraphic synthesis demonstrates the conformity of the OAE2 successions correlated to Pueblo, shows that the duration of the lag between the onset of the Osi and δ13C excursions is ∼60 k.y., and thus constrains the magnitude of the pre-OAE2 hiatus in Pueblo to less than this value. The new astronomically tuned, conformable Osi record across the onset of OAE2 captures a geologically rapid onset of large igneous province volcanism, consistent with other records, such that the addition of CO2 to the ocean-atmosphere system may have driven changes in marine carbonate chemistry. Additionally, the refined chronostratigraphy of OAE2 and the Cenomanian-Turonian boundary in the central WIB improves correlation with other records, such as those in the Eagle Ford Group, Texas. The correlations highlight that discrepancies among OAE2 age models from globally distributed sections commonly stem from differing definitions of the event and uncertainties associated with astronomical tuning, in addition to stratigraphic preservation.