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Interbasinal stratigraphic correlation provides the foundation for all consequent continental-scale geological and paleontological analyses. Correlation requires synthesis of lithostratigraphic, biostratigraphic and geochronologic data, and must be periodically updated to accord with advances in dating techniques, changing standards for radiometric dates, new stratigraphic concepts, hypotheses, fossil specimens, and field data. Outdated or incorrect correlation exposes geological and paleontological analyses to potential error. The current work presents a high-resolution stratigraphic chart for terrestrial Late Cretaceous units of North America, combining published chronostratigraphic, lithostratigraphic, and biostratigraphic data. 40Ar / 39Ar radiometric dates are newly recalibrated to both current standard and decay constant pairings. Revisions to the stratigraphic placement of most units are slight, but important changes are made to the proposed correlations of the Aguja and Javelina formations, Texas, and recalibration corrections in particular affect the relative age positions of the Belly River Group, Alberta; Judith River Formation, Montana; Kaiparowits Formation, Utah; and Fruitland and Kirtland formations, New Mexico. The stratigraphic ranges of selected clades of dinosaur species are plotted on the chronostratigraphic framework, with some clades comprising short-duration species that do not overlap stratigraphically with preceding or succeeding forms. This is the expected pattern that is produced by an anagenetic mode of evolution, suggesting that true branching (speciation) events were rare and may have geographic significance. The recent hypothesis of intracontinental latitudinal provinciality of dinosaurs is shown to be affected by previous stratigraphic miscorrelation. Rapid stepwise acquisition of display characters in many dinosaur clades, in particular chasmosaurine ceratopsids, suggests that they may be useful for high resolution biostratigraphy.

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.

Troodontids 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. Here 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. 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.

The Upper Cretaceous (Campanian Stage) Kaiparowits Formation of southern Utah, USA, preserves abundant plant, invertebrate, and vertebrate fossil taxa. Taken together, these fossils indicate that the ecosystems preserved in the Kaiparowits Formation were characterized by high biodiversity. Hundreds of vertebrate and invertebrate species and over 80 plant morphotypes are recognized from the formation, but insects and their associations with plants are largely undocumented. Here, we describe a new fossil leaf taxon, Catula gettyi gen et. sp. nov. in the family Lauraceae from the Kaiparowits Formation. Catula gettyi occurs at numerous localities in this deposit that represent ponded and distal floodplain environments. The type locality for C . gettyi has yielded 1,564 fossil leaf specimens of this species, which provides the opportunity to circumscribe this new plant species. By erecting this new genus and species, we are able to describe ecological associations on C . gettyi and place these interactions within a taxonomic context. We describe an extensive archive of feeding damage on C . gettyi caused by herbivorous insects, including more than 800 occurrences of insect damage belonging to five functional feeding groups indicating that insect-mediated damage on this taxon is both rich and abundant. Catula gettyi is one of the best-sampled host plant taxa from the Mesozoic Era, a poorly sampled time interval, and its insect damage is comparable to other Lauraceae taxa from the younger Late Cretaceous Hell Creek Flora of North Dakota, USA.

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. 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. 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.

Tyrannosaurids are hypothesized to be gregarious, possibly parasocial carnivores engaging in cooperative hunting and extended parental care. A tyrannosaurid (cf. Teratophoneus curriei ) bonebed in the late Campanian age Kaiparowits Formation of southern Utah, nicknamed the Rainbows and Unicorns Quarry (RUQ), provides the first opportunity to investigate possible tyrannosaurid gregariousness in a taxon unique to southern Laramidia. Analyses of the site’s sedimentology, fauna, flora, stable isotopes, rare earth elements (REE), charcoal content and taphonomy suggest a complex history starting with the deaths and transport of tyrannosaurids into a peri-fluvial, low-energy lacustrine setting. Isotopic and REE analyses of the fossil material yields a relatively homogeneous signature indicating the assemblage was derived from the same source and represents a fauna living in a single ecospace. Subsequent drying of the lake and fluctuating water tables simultaneously overprinted the bones with pedogenic carbonate and structurally weakened them through wet-dry cycling. Abundant charcoal recovered from the primary bone layer indicate a low temperature fire played a role in the site history, possibly triggering an avulsion that exhumed and reburied skeletal material on the margin of a new channel with minimal transport. Possible causes of mortality and concentration of the tyrannosaurids include cyanobacterial toxicosis, fire, and flooding, the latter being the preferred hypothesis. Comparisons of the RUQ site with other North American tyrannosaur bonebeds (Dry Island-Alberta; Daspletosaurus horneri -Montana) suggest all formed through similar processes. Combined with ichnological evidence, these tyrannosaur mass-burial sites could be part of an emerging pattern throughout Laramidia reflecting innate tyrannosaurid behavior such as habitual gregariousness.

The Kaiparowits Formation preserves one of the best fossil records of Cretaceous North America, which provides great insight into the paleoecology. In an effort to investigate the paleohydrology of the Kaiparowits Formation, stable isotope compositions (δ13C, δ18O-carbonate, δ18O-phosphate) of 41 hadrosaur teeth, 27 crocodile teeth, and 35 turtle shell fragments were analyzed. The mean O-isotope compositions of drinking water (δ18Ow) calculated from the O-isotope of bioapatite (phosphate-δ18Op) are −13.76 ± 2.08‰ (SMOW) for hadrosaur, −8.88 ± 2.76‰ (SMOW) for crocodile, and −10.14 ± 2.62‰ (SMOW) for turtle, which strongly reflect niche partitioning. The Kaiparowits formation does not fit the global trend in isotopic compositions of vertebrate skeletal remains from previous studies, which suggests a unique hydrological setting of the Kaiparowits basin. High-elevation runoff from the Mogollon Highlands and sea level fluctuation may have contributed to such a unique paleohydrology.

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.

The most complete known North American enantiornithine was collected in 1992 but never formally described. The so-called “Kaiparowits avisaurid” remains one of the most exceptional Late Cretaceous enantiornithine fossils. We recognize this specimen as a new taxon, Mirarce eatoni (gen. et sp. nov.), and provide a complete anatomical description. We maintain that the specimen is referable to the Avisauridae, a clade previously only known in North America from isolated tarsometatarsi. Information from this specimen helps to clarify evolutionary trends within the Enantiornithes. Its large body size supports previously observed trends toward larger body mass in the Late Cretaceous. However, trends toward increased fusion of compound elements across the clade as a whole are weak compared to the Ornithuromorpha. The new specimen reveals for the first time the presence of remige papillae in the enantiornithines, indicating this feature was evolved in parallel to dromaeosaurids and derived ornithuromorphs. Although morphology of the pygostyle and (to a lesser degree) the coracoid and manus appear to remain fairly static during the 65 million years plus of enantiornithine evolution, by the end of the Mesozoic at least some enantiornithine birds had evolved several features convergent with the Neornithes including a deeply keeled sternum, a narrow furcula with a short hypocleidium, and ulnar quill knobs—all features that indicate refinement of the flight apparatus and increased aerial abilities. We conduct the first cladistic analysis to include all purported avisuarid enantiornithines, recovering an Avisauridae consisting of a dichotomy between North and South American taxa. Based on morphological observations and supported by cladistic analysis, we demonstrate Avisaurus to be paraphyletic and erect a new genus for “ A. gloriae ,” Gettyia gen. nov.

For nearly 60 years, skulls of Parasaurolophus species have been differentiated primarily on the basis of crest shape rather than on unique morphologic characters of other cranial elements. Complicating matters is the fact that crests dramatically change shape throughout ontogeny. Without a complete growth series, it has become difficult to assess the taxonomic distinctness of each species through the lens of allometric growth. Parasaurolophus cyrtocristatus has proven to be especially troublesome to assess because of the poorly preserved nature of the type and only skull. A new, partial skull from the Fossil Forest Member of the Fruitland Formation—the same geologic unit as the type specimen—is the first opportunity to re-diagnose this species as well as redefine the genus with many new traits. An undescribed, short-crested subadult skull from the Kaiparowits Formation of Utah previously assigned to cf. P. cyrtocristatus allows detailed comparisons to be made between the unnamed Utah taxon and the material of this species from the type locality. We find that several characteristics of the squamosal, supraoccipital, and premaxilla shared between the referred skull and the type skull are unique to P. cyrtocristatus (senso stricto) within the genus, irrespective of the overall crest shape. A phylogenetic analysis that includes six new characters posits that P. cyrtocristatus and P. tubicen are sister taxa, and that the latter does not share a closest common ancestor with the long-crested P. walkeri as previously hypothesized. This result helps to explain why both taxa are found in northeastern New Mexico, USA and in sequential geologic units (Fruitland Formation and Kirtland Formation, respectively). Additionally, the exquisitely preserved new skull provides the first opportunity to unequivocally identify the osteological make-up of the Parasaurolophus cranial crest. Unlike in previous reconstructions, the crest composition in Parasaurolophus follows what is seen in other lambeosaurines such as Corythosaurus , where the dorsal process of the premaxilla dominates the crest, with the nasal forming 80% of the ventral paired tubes, and the lateral premaxillary process acting a lateral cover between the dorsal and ventral tubes. The skull of P. cyrtocristatus is still incompletely known, so more complete material will likely reveal new features that further differentiate this species and aid in determining the pace of ornamental crest evolution.