Mark A. Loewen

A new centrosaurine from the Late Cretaceous of Alberta, Canada, and the evolution of parietal ornamentation in horned dinosaurs

In 1916, a centrosaurine dinosaur bonebed was excavated within the Campanian-aged deposits of what is now Dinosaur Provincial Park, Alberta, Canada. Specimens from this now-lost quarry, including two parietals, a squamosal, a skull missing the frill, and an incomplete dentary, were purchased by The Natural History Museum, London. The material was recently reprepared and identified herein as a previously unknown taxon, Spinops sternbergorum gen. et sp. nov. Based upon the available locality data and paleopalynology, the quarry lies in either the upper part of the Oldman Formation or the lower part of the Dinosaur Park Formation. The facial region of the partial skull is similar to putative mature specimens of Centrosaurus spp. and Styracosaurus albertensis, with short, rounded postorbital horncores and a large, erect nasal horncore. Parietal ornamentation is consistent on both known parietals and is unique among ceratopsids. Bilateral, procurved parietal hooks occupy the P1 (medial-most) position on the dorsal surface of the parietal and are very similar to those seen in Centrosaurus apertus. Epiparietals in the P2 or possibly P3 position (lateral to P1) manifest as extremely elongate, caudally directed spikes, unlike the condition in C. apertus, S. albertensis, or any other “derived” centrosaurine. Cladistic analysis suggests that S. sternbergorum is closely related to Centrosaurus and Styracosaurus. Historically, based upon the condition in Styracosaurus and related centrosaurines, it was assumed that the medial-most elongated spikes on centrosaurine parietals correspond to the P3 epiparietal position. The exception illustrated in the new taxon suggests that homologies of epiparietals among basal centrosaurines (e.g., Albertaceratops and Diabloceratops) and derived centrosaurines (e.g., Styracosaurus and “pachyrhinosaurs”) should be reconsidered. The medially-placed, caudally-directed “P3” process of basal centrosaurines may, in fact, be homologous with P2.

A remarkable short-snouted horned dinosaur from the Late Cretaceous (late Campanian) of southern Laramidia

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 groups 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 basal nodosaurid ankylosaur Europelta carbonensis n. gen., n. sp. from the Lower Cretaceous (lower Albian) Escucha Formation of northeastern Spain.

Nodosaurids are poorly known from the Lower Cretaceous of Europe. Two associated ankylosaur skeletons excavated from the lower Albian carbonaceous member of the Escucha Formation near Ariño in northeastern Teruel, Spain reveal nearly all the diagnostic recognized character that define nodosaurid ankylosaurs. These new specimens comprise a new genus and species of nodosaurid ankylosaur and represent the single most complete taxon of ankylosaur from the Cretaceous of Europe. These two specimens were examined and compared to all other known ankylosaurs. Comparisons of these specimens document that Europelta carbonensis n. gen., n. sp. is a nodosaur and is the sister taxon to the Late Cretaceous nodosaurids Anoplosaurus, Hungarosaurus, and Struthiosaurus, defining a monophyletic clade of European nodosaurids– the Struthiosaurinae.

Eocene Green River Formation “Oocardium tufa” reinterpreted as complex arrays of calcified caddisfly (Insecta: Trichoptera) larval cases

Abstract Fossil caddisfly (Insecta: Trichoptera) larval cases are preserved in some Eocene Green River Formation tufa, and well organized arrays of these calcified larval cases have been misidentified as “Oocardium tufa”. The arrays of caddisfly larval cases occur as superimposed layers of vertically oriented cases. The cases are parallel to each other, closely packed, equal in size, and are oriented perpendicular to the bedding planes of the tufa. In thin section, the cases are composed of detrital carbonate particles, ooids and quartz grains arranged in a single layer along the walls of the cases. The size and arrangement of the particles in the walls of the fossil caddisfly larval cases compares favorably with modern caddisfly larval cases. Correct identification of the origin of this unique tufa fabric enables us to better understand the important role of metazoans (Trichopteran larvae) in some Eocene Lake Gosiute paleoenvironments. Caddisfly larvae played direct and indirect roles in the deposition and erosion of tufa in Lake Gosiute during the deposition of the Laney Member of the Green River Formation.

A New Centrosaurine Ceratopsid, Machairoceratops cronusi gen et sp. nov., from the Upper Sand Member of the Wahweap Formation (Middle Campanian), Southern Utah

The Upper Cretaceous (middle-late Campanian) Wahweap Formation of southern Utah contains the oldest diagnostic evidence of ceratopsids (to date, all centrosaurines) in North America, with a number of specimens recovered from throughout a unit that spans between 81 and 77 Ma. Only a single specimen has been formally named, Diabloceratops eatoni, from the lower middle member of the formation. Machairoceratops cronusi gen. et sp. nov., a new centrosaurine ceratopsid from the upper member of the Wahweap Formation, is here described based on cranial material representing a single individual recovered from a calcareous mudstone. The specimen consists of two curved and elongate orbital horncores, a left jugal, a nearly complete, slightly deformed braincase, the left squamosal, and a mostly complete parietal ornamented by posteriorly projected, anterodorsally curved, elongate spikes on either side of a midline embayment. The fan-shaped, stepped-squamosal is diagnostic of Centrosaurinae, however, this element differs from the rectangular squamosal in Diabloceratops. Machairoceratops also differs in the possession of two anterodorsally (rather than laterally) curved epiparietal ornamentations on either side of a midline embayment that are distinguished by a posteromedially-oriented sulcus along the entire length of the epiparietal. Additionally, the parietosquamosal frill is lacking any other epiossifications along its periphery. Machairoceratops shares a triangular (rather than round) frill and spike-like epiparietal loci (p1) ornamentation with the stratigraphically lower Diabloceratops. Both parsimony and Bayesian phylogenetic analyses place Machairoceratops as an early-branching centrosaurine. However, the parsimony-based analysis provides little resolution for the position of the new taxon, placing it in an unresolved polytomy with Diabloceratops. The resultant Bayesian topology yielded better resolution, aligning Machairoceratops as the definitive sister taxon to a clade formed by Diabloceratops and Albertaceratops. Considered together, both phylogenetic methods unequivocally place Machairoceratops as an early-branching centrosaurine, and given the biostratigraphic position of Machairoceratops, these details increase the known ceratopsid diversity from both the Wahweap Formation and the southern portion of Laramidia. Finally, the unique morphology of the parietal ornamentation highlights the evolutionary disparity of frill ornamentation near the base of Centrosaurinae.

New material and systematic re-evaluation of Medusaceratops lokii (Dinosauria, Ceratopsidae) from the Judith River Formation (Campanian, Montana)

Abstract. Medusaceratops lokii Ryan, Russell, and Hartman, 2010 is an enigmatic taxon of ceratopsid represented by partial parietals from the Mansfield bonebed in the Campanian Judith River Formation, Montana. Originally, all ceratopsid material collected from this bonebed was referred to the centrosaurine ceratopsid Albertaceratops, but subsequently two parietals were designated the types of the chasmosaurine, M. lokii, in part, because they were interpreted to have three epiparietals bilaterally. Here we describe new material from the bonebed that allows a systematic revision of the taxon. A revised reconstruction of the frill, informed by newly discovered parietals, reveals that M. lokii had a broad midline ramus and at least five epiparietals (ep) around the margin of the frill, both traits that are characteristic of Centrosaurinae. From medial to lateral, the epiparietal ornamentation consists of a small, variably procurving epiparietal (ep 1), an anterolaterally curving pachyostotic hook (ep 2), a smaller pachyostoic process (ep 3), and two small triangular epiparietals (ep 4 and 5). A phylogenetic analysis of ceratopsids, which is the first to include Medusaceratops, indicates that M. lokii is a unique, early centrosaurine ceratopsid taxon that is more closely related to Centrosaurini and Pachyrhinosaurini than Nasutoceratopsini. No unequivocal chasmosaurine bones or diagnostic material from any other ceratopsid could be identified from the Mansfield bonebed, suggesting that it represents one of the oldest occurrences of a monodominant accumulation of a centrosaurine ceratopsid on record.

Microbial-caddisfly bioherm association from the Lower Cretaceous Shinekhudag Formation, Mongolia: Earliest record of plant armoring in fossil caddisfly cases

Caddisfly larvae construct underwater protective cases using surrounding materials, thus providing information on environmental conditions in both modern and ancient systems. Microbial bioherms associated with caddisfly cases are found in the Berriassian-Hauterivian (~140–130 Ma) Shinekhudag Formation of Mongolia, and yield new insights into aspects of lacustrine paleoecosystems and paleoenvironments. This formation contains the earliest record of plant-armored caddisfly cases and a rare occurrence of microbial-caddisfly association from the Mesozoic. The bioherms are investigated within the context of stratigraphic correlations, depositional environment interpretations, and basin-evolution models of the sedimentary fill. The bioherms form 0.5–2.0 m diameter mound-shaped bodies and are concentrated within a single, oil shale-bound stratigraphic interval. Each bioherm is composed of up to 40% caddisfly cases along with stromatolites of millimeter-scale, micritic laminations. Petrographic analyses reveal these bioherms are composed of non-systematic associations of columnar and oncoidal microbialites, constructed around colonies of caddisfly cases. The cases are straight to curved, slightly tapered, and tube-shaped, with a progressively increasing length and width trend (7–21 mm by 1.5–2.5 mm). Despite these variations, the case architectures reveal similar construction materials; the particles used for cases are dominated by plant fragments, ostracod valves, carbonate rocks, and rare mica and feldspar grains. Allochems within the bioherms include ooids, ostracods, plant fragments, rare gastropods, feldspar grains bound in micritic matrices, and are consolidated by carbonate dominated cements. The combination of microbial-caddisfly association, plant fragment case particles, and ooids/oncoids are indicative of a shallow, littoral lake setting. Stratigraphic juxtaposition of nearshore bioherms and the bounding distal oil-shale facies suggests that the bioherms developed in an underfilled lake basin, resulting from an abrupt and short-lived lake desiccation event. Lake chemistry is believed to have been relatively alkaline, saline to hypersaline, and rich in Ca, Mg, and HCO3 ions. Through analyzing bioherm characteristics, caddisfly case architecture, carbonate microfacies, and stratigraphic variability, we infer larger-scale processes that controlled basin development during their formation.