Jordan C. Mallon

Variation in the Skull of Anchiceratops (Dinosauria, Ceratopsidae) from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta

ABSTRACT Anchiceratops is a chasmosaurine ceratopsid from the Upper Cretaceous Horseshoe Canyon Formation (HCF) of Alberta. It is distinguished primarily by its unique parietosquamosal frill ornamentation and possibly by the presence of a ventrally flexed olfactory bulb of the brain. Although Anchiceratops is known from at least ten partial skulls, only two of these have been formally described. These skulls are not stratigraphically segregated, but they differ markedly in their proportions (e.g., supraorbital horncore and frill dimensions), causing previous authors to account for this disparity with reference to either interspecific or sexual differences. Both of these hypotheses assume that variation in Anchiceratops is dimorphic; however, this assumption has never been tested with reference to all available material. The present study describes all material from the HCF that can be positively attributed to Anchiceratops, and tests the assumption of dimorphism by subjecting this material to a series of morphometric analyses. We find no compelling evidence for dimorphism in Anchiceratops, although sample size is still too small for convincing statistical support. We conclude that there is a single, variable species of Anchiceratops, A. ornatus. Average sedimentation rates for the HCF suggest that A. ornatus is a particularly long-lived species compared with other ceratopsids (∼1.5–2.0 Ma), and the paleoecological implications of this are discussed. A cladistic analysis that includes the new data presented here indicates that Anchiceratops is more closely related to Chasmosaurus than to Triceratops, in contrast with previous studies.

Skull Ecomorphology of Megaherbivorous Dinosaurs from the Dinosaur Park Formation (Upper Campanian) of Alberta, Canada

Megaherbivorous dinosaur coexistence on the Late Cretaceous island continent of Laramidia has long puzzled researchers, owing to the mystery of how so many large herbivores (6–8 sympatric species, in many instances) could coexist on such a small (4–7 million km2) landmass. Various explanations have been put forth, one of which–dietary niche partitioning–forms the focus of this study. Here, we apply traditional morphometric methods to the skulls of megaherbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta to infer the ecomorphology of these animals and to test the niche partitioning hypothesis. We find evidence for niche partitioning not only among contemporaneous ankylosaurs, ceratopsids, and hadrosaurids, but also within these clades at the family and subfamily levels. Consubfamilial ceratopsids and hadrosaurids differ insignificantly in their inferred ecomorphologies, which may explain why they rarely overlap stratigraphically: interspecific competition prevented their coexistence.

The Functional and Palaeoecological Implications of Tooth Morphology and Wear for the Megaherbivorous Dinosaurs from the Dinosaur Park Formation (Upper Campanian) of Alberta, Canada

Megaherbivorous dinosaurs were exceptionally diverse on the Late Cretaceous island continent of Laramidia, and a growing body of evidence suggests that this diversity was facilitated by dietary niche partitioning. We test this hypothesis using the fossil megaherbivore assemblage from the Dinosaur Park Formation (upper Campanian) of Alberta as a model. Comparative tooth morphology and wear, including the first use of quantitative dental microwear analysis in the context of Cretaceous palaeosynecology, are used to infer the mechanical properties of the foods these dinosaurs consumed. The phylliform teeth of ankylosaurs were poorly adapted for habitually processing high-fibre plant matter. Nevertheless, ankylosaur diets were likely more varied than traditionally assumed: the relatively large, bladed teeth of nodosaurids would have been better adapted to processing a tougher, more fibrous diet than the smaller, cusp-like teeth of ankylosaurids. Ankylosaur microwear is characterized by a preponderance of pits and scratches, akin to modern mixed feeders, but offers no support for interspecific dietary differences. The shearing tooth batteries of ceratopsids are much better adapted to high-fibre herbivory, attested by their scratch-dominated microwear signature. There is tentative microwear evidence to suggest differences in the feeding habits of centrosaurines and chasmosaurines, but statistical support is not significant. The tooth batteries of hadrosaurids were capable of both shearing and crushing functions, suggestive of a broad dietary range. Their microwear signal overlaps broadly with that of ankylosaurs, and suggests possible dietary differences between hadrosaurines and lambeosaurines. Tooth wear evidence further indicates that all forms considered here exhibited some degree of masticatory propaliny. Our findings reveal that tooth morphology and wear exhibit different, but complimentary, dietary signals that combine to support the hypothesis of dietary niche partitioning. The inferred mechanical and dietary patterns appear constant over the 1.5 Myr timespan of the Dinosaur Park Formation megaherbivore chronofauna, despite continual species turnover.

Improving the repeatability of low magnification microwear methods using high dynamic range imaging

Abstract The recent advent of low magnification microwear analysis has allowed the efficient study of entire vertebrate faunas using only an optical stereomicroscope. Photographic visualization of microwear by this means has proven difficult, however, and, as a result, few high-resolution photos of low magnification microwear have been published. The repeatability of the method has also been questioned because low magnification microwear analysis involves the visual inspection of microwear features. We show that the use of high dynamic range imaging improves the visualization of microwear features in photographs and that using these photographs as a counting medium increases the repeatability of the method. We also show that counting from the photographs allows us to accurately classify ungulates as browsers, grazers, or mixed feeders.

Jaw mechanics and evolutionary paleoecology of the megaherbivorous dinosaurs from the Dinosaur Park Formation (upper Campanian) of Alberta, Canada

ABSTRACT The question of what role differential jaw mechanics may have played in facilitating dietary niche partitioning among Late Cretaceous megaherbivorous dinosaurs from Laramidia is examined, using the fossil assemblage of the Dinosaur Park Formation as a test case. We use phylogenetic inference to reconstruct the mandibular adductor musculature of these animals, which we then apply to the construction of biomechanical lever models of the mandible to estimate relative bite forces. Our findings reveal predictably weak bite forces in ankylosaurs, and comparatively high bite forces in ceratopsids and hadrosaurids, both of which possessed a mechanical advantage that produced bite forces 2–3 times higher than those forces exerted by the adductor musculature. The impressive jaw mechanism shared by the last two taxa evolved in a stepwise fashion, independently in each lineage. There is tentative evidence to suggest that nodosaurids had more powerful bites than ankylosaurids, but the overall mechanical diversity among megaherbivores from the Dinosaur Park Formation is low, suggesting that differential jaw mechanics could have played only a subsidiary role in niche partitioning. Such mechanical conservatism may have may have been selected for, or it may simply reflect the limits imposed by evolutionary constraints. Regardless, mechanical disparity patterns remained stable throughout the ∼1.5 Ma evolution of the Dinosaur Park Formation megaherbivore chronofauna.

Spiclypeus shipporum gen. et sp. nov., a Boldly Audacious New Chasmosaurine Ceratopsid (Dinosauria: Ornithischia) from the Judith River Formation (Upper Cretaceous: Campanian) of Montana, USA

This study reports on a new ceratopsid, Spiclypeus shipporum gen et sp. nov., from the lower Coal Ridge Member of the Judith River Formation in Montana, USA, which dates to ~76 Ma (upper Campanian). The species is distinguished by rugose dorsal contacts on the premaxillae for the nasals, laterally projecting postorbital horncores, fully fused and anteriorly curled P1 and P2 epiparietals, and a posterodorsally projecting P3 epiparietal. The holotype specimen is also notable for its pathological left squamosal and humerus, which show varied signs of osteomyelitis and osteoarthritis. Although the postorbital horncores of Spiclypeus closely resemble those of the contemporaneous ‘Ceratops’, the horncores of both genera are nevertheless indistinguishable from those of some other horned dinosaurs, including Albertaceratops and Kosmoceratops; ‘Ceratops’ is therefore maintained as a nomen dubium. Cladistic analysis recovers Spiclypeus as the sister taxon to the clade Vagaceratops + Kosmoceratops, and appears transitional in the morphology of its epiparietals. The discovery of Spiclypeus adds to the poorly known dinosaur fauna of the Judith River Formation, and suggests faunal turnover within the formation.

Supplementary cranial description of the types of Edmontosaurus regalis (Ornithischia: Hadrosauridae), with comments on the phylogenetics and biogeography of Hadrosaurinae

The cranial anatomy of the flat-skulled hadrosaurine Edmontosaurus regalis (Ornithischia: Hadrosauridae) is extensively described here, based on the holotype and paratype collected from the middle part of the Horseshoe Canyon Formation in southern Alberta. Focus is given to previously undocumented features of ontogenetic and phylogenetic importance. This description facilitates overall osteological comparisons between E. regalis and other hadrosaurids (especially E. annectens), and revises the diagnosis of E. regalis, to which a new autapomorphy (the dorsal half of the jugal anterior process bearing a sharp posterolateral projection into the orbit) is added. We consider the recently named Ugrunaaluk kuukpikensis from the upper Campanian/lower Maastrichtian of Alaska a nomen dubium, and conservatively regard the Alaskan material as belonging to Edmontosaurus sp.. A phylogenetic analysis of Hadrosauroidea using maximum parsimony further corroborates the sister-taxon relationship between E. regalis and E. annectens. In the strict consensus tree, Hadrosaurus foulkii occurs firmly within the clade comprising all non-lambeosaurine hadrosaurids, supporting the taxonomic scheme that divides Hadrosauridae into Hadrosaurinae and Lambeosaurinae. Within Edmontosaurini, Kerberosaurus is posited as the sister taxon to the clade of Shantungosaurus + Edmontosaurus. The biogeographic reconstruction of Hadrosaurinae in light of the time-calibrated cladogram and probability calculation of ancestral areas for all internal nodes reveals a significantly high probability for the North American origin of the clade. However, the Laramidia–Appalachia dispersals around the Santonian–Campanian boundary, inferred from the biogeographic scenario for the North American origin of Hadrosaurinae, are in conflict with currently accepted paleogeographic models. By contrast, the Asian origin of Hadrosaurinae with its relatively low probability resulting from the biogeographic analysis is worth seriously considering, despite the lack of fossil material from the Santonian and lower Campanian of Asia. Extra fossil collecting in appropriate geographic locations and stratigraphic intervals of Asia and Europe will help to clarify the biogeographic dynamics of hadrosaurine dinosaurs in the near future.

Basilemys morrinensis, a new species of nanhsiungchelyid turtle from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada

ABSTRACT We describe a new species of nanhsiungchelyid turtle, Basilemys morrinensis, based on a nearly complete shell from the Horsethief Member (lower Maastrichtian) of the Horseshoe Canyon Formation of Alberta. The species is intermediate in age between the Campanian forms B. variolosa and B. gaffneyi and the upper Maastrichtian forms B. sinuosa and B. praeclara. It is also intermediate in its morphology, possessing a unique suite of both plesiomorphic (e.g., divided extragulars) and derived (e.g., square epiplastral beak, pygal wider than long) traits. It is further characterized by an autapomorphic square cervical scale. Phylogenetic analysis assuming parsimony recovers B. morrinensis in a polytomy with B. variolosa and B. gaffneyi, outside the clade formed by the upper Maastrichtian forms B. sinuosa and B. praeclara. The holotype of Basilemys morrinensis provides the first evidence that this genus reached large size (~1 m long) in the Horseshoe Canyon Formation and was not diminutive as previously thought based on less complete shell material. Although Basilemys is usually regarded as terrestrial in habit based on its skull and limb morphology, we note that the low profile of its shell is a derived feature usually indicative of an aquatic mode of life. This suggests that there is yet much to learn about the life habits of this interesting turtle.