Alan L. Titus

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 North American therizinosaurid and the role of herbivory in ‘predatory’ dinosaur evolution

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.

PLESIOSAURS FROM THE UPPER CRETACEOUS (CENOMANIAN–TURONIAN) TROPIC SHALE OF SOUTHERN UTAH, PART 2:POLYCOTYLIDAE

Abstract 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.

A New Troodontid Theropod, Talos sampsoni gen. et sp. nov., from the Upper Cretaceous Western Interior Basin of North America

Background 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. Methodology/Principal Findings 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. 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.

PLESIOSAURS FROM THE UPPER CRETACEOUS (CENOMANIAN–TURONIAN) TROPIC SHALE OF SOUTHERN UTAH, PART 1: NEW RECORDS OF THE PLIOSAUR BRACHAUCHENIUS LUCASI

Abstract Recent fieldwork in the Tropic Shale of southern Utah has resulted in the recovery of two new specimens of the short-necked pliosaurid Brachauchenius lucasi Williston, 1903. Both specimens were recovered in Lower Turonian strata a few meters above the Cenomanian-Turonian Stage boundary as determined from molluscan assemblages found in direct association, and both include skull material. The larger of the two also includes pectoral and pelvic elements previously unknown for this taxon. These specimens represent the first records of B. lucasi from the western margin of the Cretaceous Western Interior Seaway; previously recorded specimens were recovered in Kansas and Texas. The Cenomanian through middle Turonian temporal range of B. lucasi, in addition to its geographic range across the seaway, indicates that this taxon suffered few, if any, negative consequences as a result of global scale oceanographic events, including marine extinctions, that transpired during late Cenomanian-early Turonian time.

A gravid fossil turtle from the Upper Cretaceous (Campanian) Kaiparowits Formation, southern Utah

An Adocus turtle collected from the Upper Cretaceous (late Campanian) Kaiparowits Formation in southern Utah contains partial eggs and numerous well-preserved bones beneath the carapace. The bones include the closely associated–articulated elements of the right pectoral girdle, skull and cervical vertebrae. The eggs are about 35 mm in diameter, and the rigid 250–280-μm thick eggshell exhibits needle-like aragonitic crystals that form the narrow, straight shell units. The eggs appear smaller than those within a gravid Adocus from Alberta, and the eggshell is thinner and lacks the feathered structure reported in the Alberta eggs. Differences in the Utah and Alberta Adocus eggs may result from differences in the stage of egg formation or the specimens may represent different Adocus species.