Combat Lesion in Triceratops: Histological and Chemical Diagnosis

In the collective imagination derived from scientific and popular literature, Triceratops often faced each other in combat. Thus, from the second half of the twentieth century, these ceratopsids were described as pugnacious animals. This arises primarily from the interpretation of extracranial fenestrae in ceratopsids being the result of combat trauma. However, the diagnosis of the traumatic nature of these anatomical variants of their neck frill requires evidence of bone healing and remodelling by microscopy analysis. Here, we present the case of the Triceratops horridus known as Big John, which is one of the largest specimens discovered in the Hell Creek Formation (Upper Cretaceous; MT, USA). Its right squamosal bone shows an extrafenestra with irregular margins and signs of inflammation. Microscopy analysis revealed newly formed and healing bone, with histological signs typical of the bone remodelling phase. Chemical analysis revealed sulphur that was derived from glucosaminoglycans and sulphated glycoproteins of the preosseous osteoid substance present in the healing phases of a bone trauma. Histological and microanalytical analyses confirm that the squamosal fenestra of Big John is the result of a traumatic event, which might indeed have occurred during combat with another Triceratops.

The promise of taphonomy as a nomothetic discipline: taphonomic bias in two dinosaur-bearing faunas in North America

The fossil record of dinosaurs is a rich, if biased, one with nearly complete skeletons, partial skeletons, and isolated parts found in diverse, well-studied faunal assemblages around the world. Among the recognized biases are the preferential preservation of large dinosaurs and the systematic underrepresentation of small dinosaurs. Such biases have been quantitatively described in the Upper Cretaceous (Campanian) Dinosaur Park Formation of Alberta, where large, nearly complete dinosaurs were found and described early in collecting history and small, very incomplete dinosaurs were found and described later. This pattern, apparently replicated in the Maastrichtian Hell Creek Formation of Montana, is so striking that it begs the question of whether this is a nomothetic principle for the preservation of dinosaur faunas elsewhere. We tested this hypothesis by analyzing the very well-studied dinosaur fauna of the Upper Jurassic (Kimmeridgian) Morrison Formation of the western United States. The Morrison Formation fails to show any correlation between body size and completeness, order of discovery, or order of description. Both large and small dinosaurs of the Morrison include highly complete as well as highly incomplete taxa, and both large and small dinosaurs were discovered and described early in collecting history as well as more recently. The differences in preservation between the Dinosaur Park Formation and the Morrison Formation are so striking that we posit a Dinosaur Park model of dinosaur fossil preservation and a Morrison model. Future study will show whether either or both represent durable nomothetic models for dinosaur fossil preservation.

Metatarsals of a large caenagnathid cf. Anzu wyliei (Theropoda: Oviraptorosauria) from the Hell Creek Formation in South Dakota, U.S.A.

Isolated metatarsals III and IV of a caenagnathid theropod likely referable to Anzu wyliei are described from a locality of the Hell Creek Formation in northwestern South Dakota of the U.S.A. These bones are missing from the holotype and only partial shafts have been described for a specimen referable to this species. Accordingly, the present description adds further anatomical information on this already well-known species of Caenagnathidae. The present finding also demonstrates the significance of isolated or fragmentary specimens found in multitaxic bone beds.

A systematic reappraisal and quantitative study of the non-marine teleost fishes from the late Maastrichtian of the Western Interior of North America – evidence from vertebrate microfossil localities

The diversity and distribution of non-marine teleost fishes in the Western Interior of North America during the late Maastrichtian is documented based on isolated elements from vertebrate microfossil localities in the Hell Creek Formation of Montana, the Lance Formation of Wyoming, and the Scollard Formation of Alberta. A minimum of 20 taxa are recognized based on >1900 abdominal centra and tooth-bearing elements. These include two elopomorphs, six osteoglossomorphs, three ostariophysans, one esocid, six acanthomorphs, and two taxa of unknown relationships. These assemblages differ from late Campanian assemblages in the absence of the Clupeomorpha and the presence of the Perciformes. Within the Hell Creek Formation, we record patterns in the relative abundances of the most abundant taxa leading up to the Cretaceous/Paleogene (K/Pg) boundary. Most notably, acanthomorphs increased in abundance up-section whereas a group of osteoglossomorphs represented by Coriops and/or Lopadichthys concurrently decreased in abundance. Conversely, some teleosts exhibited more stable or slightly fluctuating relative abundances through the formation (Wilsonichthyidae, Esocidae). These late Maastrichtian teleost assemblages are of higher diversity than an early Eocene assemblage from Wyoming that is preserved under similar taphonomic conditions. This pattern either suggests that lower Cenozoic deposits in the Western Interior are insufficiently sampled or that the K/Pg mass extinction event adversely affected non-marine teleosts.

A chasmosaurine ceratopsid premaxilla from the basal sandstone of the Hell Creek Formation, Montana

A well-preserved large chasmosaurine ceratopsid premaxilla (MOR 1122 7-22-00-1) collected from the basal sandstone of the Cretaceous Hell Creek Formation (HCF) represents one of the stratigraphically lowest ceratopsid occurrences in the formation. The specimen was discovered in 2000, during the excavation of a large Torosaurus latus skull (MOR 1122) which was later hypothesized to represent an advanced growth stage of the more commonly recovered HCF ceratopsid Triceratops. MOR 1122 7-22-00-1 compares favorably with the incomplete premaxillae of the MOR 1122 skull and reveals details of premaxilla morphology from this stratigraphic zone. It preserves large, closely spaced ventromedial foramina, a narrow triangular process, and a thin septal flange at the base of the narial strut. The nasal process is narrow, caudally inclined and has a forked dorsal surface which appears to represent an intermediate between the morphology expressed in the slightly stratigraphically lower ceratopsid Eotriceratops xerinsularis from the Horseshoe Canyon Formation of Alberta and specimens recovered higher in the HCF. MOR 1122 7-22-00-1 expresses a deep recess extending medial to the strut of the triangular process, a feature shared with other HCF ceratopsid specimens but not Eotriceratops or other earlier occurring triceratopsin taxa. The morphology of MOR 1122 7-22-00-1 is consistent with noted stratigraphic trends in HCF ceratopsids and highlights the increased complexity of the narial region in uppermost Cretaceous triceratopsins.

The Hell Creek Formation, Montana: A Stratigraphic Review and Revision Based on a Sequence Stratigraphic Approach

The Upper Maastrichtian fluvial Hell Creek Formation of the Fort Peck Lake area, Montana (and regional equivalents) is notable for its vertebrate fossils and for the K-Pg mass extinction at or near its upper contact. Despite intense study, internal stratigraphy of the Hell Creek Formation is still poorly constrained, hindering study. This work reviews the stratigraphy of the Hell Creek Formation, as currently understood, and proposes important revisions to the recently proposed type section, particularly concerning complexity of the Hell Creek Formation basal contact. This work also subdivides the Montanan Hell Creek Formation into four 4th order depositional sequences, superimposed over a 3rd order marine transgression. Sequence boundaries are defined by four, laterally continuous disconformities formed by pauses in the creation of accommodation space, marked by overlying amalgamated channel complexes, or less commonly, correlative interfluve paleosols. Cyclicity in Montana may be correlative with similar 4th order cyclicity and marine influence documented in North and South Dakota, Alberta, and Saskatchewan. Magnetostratigraphy and new biostratigraphic data support correlation of the upper Montanan sequence with the North Dakotan Cantapeta tongue (and overlying fines) and Canadian Scollard and Frenchman Formations.

Molecular tests support the viability of rare earth elements as proxies for fossil biomolecule preservation

The rare earth element (REE) composition of a fossil bone reflects its chemical alteration during diagenesis. Consequently, fossils presenting low REE concentrations and/or REE profiles indicative of simple diffusion, signifying minimal alteration, have been proposed as ideal candidates for paleomolecular investigation. We directly tested this prediction by conducting multiple biomolecular assays on a well-preserved fibula of the dinosaur Edmontosaurus from the Cretaceous Hell Creek Formation previously found to exhibit low REE concentrations and steeply-declining REE profiles. Gel electrophoresis identified the presence of organic material in this specimen, and subsequent immunofluorescence and enzyme-linked immunosorbant assays identified preservation of epitopes of the structural protein collagen I. Our results thereby support the utility of REE profiles as proxies for soft tissue and biomolecular preservation in fossil bones. Based on considerations of trace element taphonomy, we also draw predictions as to the biomolecular recovery potential of additional REE profile types exhibited by fossil bones.