title: New horned dinosaurs from Utah provide evidence for intracontinental dinosaur endemism.
abstract: 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.
  
  
  RESULTS
  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
  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.
authors.affiliation: Utah Museum of Natural History and Department of Geology and Geophysics, University of Utah, Salt Lake City, Utah, USA. ssampson@umnh.utah.edu
authors.forename: Scott D
authors.initials: SD
authors.lastname: Sampson
authors.forename: Mark A
authors.initials: MA
authors.lastname: Loewen
authors.forename: Andrew A
authors.initials: AA
authors.lastname: Farke
authors.forename: Eric M
authors.initials: EM
authors.lastname: Roberts
authors.forename: Catherine A
authors.initials: CA
authors.lastname: Forster
authors.forename: Joshua A
authors.initials: JA
authors.lastname: Smith
authors.forename: Alan L
authors.initials: AL
authors.lastname: Titus
mesh_terms: D000818:Animals; D025061:Dinosaurs / Q000033:anatomy & histology / Q000145:classification*; D005580:Fossils; D009656:North America; D010163:Paleontology; D010802:Phylogeny; D014589:Utah