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Concept: Ceratopsidae

356

The fossil record of ceratopsid dinosaurs between the occurrence of their proximate sister taxa in the Turonian and the beginning of their well-documented radiation from the late Campanian of North America onwards (approximately 90 and 77 Ma) is poor, with only seven taxa described from this early period in their evolution. We describe a new taxon of a highly adorned basal centrosaurine, Wendiceratops pinhornensis gen. et sp. nov., from the lower part of the Oldman Formation (middle Campanian, approximately 78-79 Ma), Alberta, Canada. Over 200 bones derived from virtually all parts of the skeleton, including multiple well-preserved specimens of the diagnostic parietosquamosal frill, were collected from a medium-density monodominant bonebed, making the new taxon one of the best-represented early ceratopsids. The new taxon is apomorphic in having epiparietals at loci 2 and 3 developed as broad-based, pachyostotic processes that are strongly procurved anterodorsally to overhang the posterior and lateral parietal rami, and an ischium with a broad, rectangular distal terminus. Although the morphology of the nasal is incompletely known, Wendiceratops is inferred to have a large, upright nasal horn located close to the orbits, which represents the oldest occurrence of this feature in Ceratopsia. Given the phylogenetic position of the new taxon within Centrosaurinae, a enlarged nasal horn is hypothesized to have arisen independently at least twice in ceratopsid evolution.

Concepts: Phylogenetics, Cladistics, Fossil, Paleontology, Dinosaur, Taxon, Ceratopsidae, Albertaceratops

279

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.

Concepts: Cladistics, Clade, Cretaceous, Dinosaur, Taxon, Linnaean taxonomy, Ceratopsidae, Nomen dubium

240

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 group’s 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.

Concepts: United States, Phylogenetics, Cladistics, North America, Cretaceous, Fossil, Dinosaur, Ceratopsidae

160

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.

Concepts: Phylogenetic nomenclature, Phylogenetics, Cladistics, Ceratopsidae, Styracosaurus, Albertaceratops, Centrosaurinae, Achelousaurus

103

Ceratopsid (horned) dinosaurs are an iconic group of large-bodied, quadrupedal, herbivorous dinosaurs that evolved in the Late Cretaceous and were largely restricted to western North America [1-5]. Ceratopsids are easily recognized by their cranial ornamentation in the form of nasal and postorbital horns and frill (capped by epiossifications); these structures show high morphological disparity and also represent the largest cranial display structures known to have evolved [2, 4]. Despite their restricted occurrence in time and space, this group has one of the best fossil records within Dinosauria, showing a rapid diversification in horn and frill morphology [1]. Here a new genus and species of chasmosaurine ceratopsid is described based on a nearly complete and three-dimensionally preserved cranium recovered from the uppermost St. Mary River Formation (Maastrichtian) of southwestern Alberta. Regaliceratops peterhewsi gen. et sp. nov. exhibits many unique characters of the frill and is characterized by a large nasal horncore, small postorbital horncores, and massive parietal epiossifications. Cranial morphology, particularly the epiossifications, suggests close affinity with the late Campanian/early Maastrichian taxon Anchiceratops, as well as with the late Maastrichtian taxon Triceratops. A median epiparietal necessitates a reassessment of epiossification homology and results in a more resolved phylogeny. Most surprisingly, Regaliceratops exhibits a suite of cranial ornamentations that are superficially similar to Campanian centrosaurines, indicating both exploration of novel display morphospace in Chasmosaurinae, especially Maastrichtian forms, and convergent evolution in horn morphology with the recently extinct Centrosaurinae. This marks the first time that evolutionary convergence in horn-like display structures has been demonstrated between dinosaur clades, similar to those seen in fossil and extant mammals [6].

Concepts: Evolution, Extinction, Cretaceous, Fossil, Paleontology, Dinosaur, Triceratops, Ceratopsidae

59

Ceratopsids (“horned dinosaurs”) are known from western North America and Asia, a distribution reflecting an inferred subaerial link between the two landmasses during the Late Cretaceous. However, this clade was previously unknown from eastern North America, presumably due to limited outcrop of the appropriate age and depositional environment as well as the separation of eastern and western North America by the Western Interior Seaway during much of the Late Cretaceous. A dentary tooth from the Owl Creek Formation (late Maastrichtian) of Union County, Mississippi, represents the first reported occurrence of Ceratopsidae from eastern North America. This tooth shows a combination of features typical of Ceratopsidae, including a double root and a prominent, blade-like carina. Based on the age of the fossil, we hypothesize that it is consistent with a dispersal of ceratopsids into eastern North America during the very latest Cretaceous, presumably after the two halves of North America were reunited following the retreat of the Western Interior Seaway.

Concepts: United States, North America, Cretaceous, Dinosaur, Maastrichtian, Mosasaur, Late Cretaceous, Ceratopsidae

27

Bravoceratops polyphemus gen. et sp. nov. is a large chasmosaurine ceratopsid from the lowermost part of the Javelina Formation (early Maastrichtian) of Big Bend National Park, TX, USA. B. polyphemus has a distinctive narrow snout, a long fenestrate frill, and a fan-shaped median parietal bar with a midline epiparietal on its posterior margin, as well as a symmetrical depression on its dorsal surface at the nexus of the parietal rami. This depression is interpreted to be the attachment point for a second midline epiparietal. This parietal morphology is distinct from that exhibited by Anchiceratops or Pentaceratops. The posterior midline epiparietal in B. polyphemus and its bifurcated quadratojugal-squamosal joint are features shared with the most derived chasmosaurines, Torosaurus and Triceratops. The combination of primitive and derived traits exhibited by B. polyphemus, and its stratigraphic position, is compatible with the gradual transition from basal, to intermediate, to derived chasmosaurines observed throughout the western interior of North America, and with phylogenetic analysis, which suggests that Bravoceratops may be closely related to Coahuilaceratops.

Concepts: United States, Computational phylogenetics, Texas, Dinosaur, National Park Service, Triceratops, Ceratopsidae, Ceratopsia

18

Psittacosauridae (parrot-beaked dinosaurs) represents the first major radiation of ceratopsians (horned dinosaurs). However, psittacosaurids are divergent from the general morphology found in other ceratopsians, and this has resulted in their uncertain systematic position among ceratopsians. Here we describe a new basal neoceratopsian dinosaur, Mosaiceratops azumai gen. et sp. nov. based on a partial semi-articulated skeleton recovered from the Upper Cretaceous Xiaguan Formation of Neixiang County, Henan Province, China. Although our phylogenetic analysis supports this taxon as the most basal neoceratopsian, Mosaiceratops exhibits many features previously considered unique to the Psittacosauridae among the basal Ceratopsia. These include a relatively highly positioned external naris, a proportionally large premaxilla, the nasal extending ventral to the external naris, slender postorbital and temporal bars, a large notch between the basal tubera, and the edentulous premaxilla. Thus, the discovery of Mosaiceratops reduces the morphological disparity between the Psittacosauridae and other basal ceratopsians. Character optimization suggests that basal neoceratopsians have re-evolved premaxillary teeth; a major reversal previously unknown in any dinosaur clade. The new specimen also highlights the mosaic nature of evolution among early ceratopsians and supports the phylogenetic hypothesis that the Psittacosauridae is a relatively derived clade, rather than the most basal group of the Ceratopsia.

Concepts: Cladistics, Cretaceous, Dinosaur, Psittacosaurus, Ceratopsidae, Ceratopsia, Protoceratops, Nanyang, Henan

13

While centrosaurines and ceratopsids in general are abundant in the Late Campanian of northern Laramidia, they are much less commonly found in southern Laramidia. This has supported hypotheses of dinosaur provinciality and endemism in the Late Cretaceous with the delineation of at least two separate faunal zones, north and south Laramidia. There have been 12 genera of centrosaurines recognized from northern Laramidia while two genera, Diabloceratops and Nasutoceratops, have been named from southern Laramidia. We present an osteological description and taphonomic outline for a new centrosaurine ceratopsid from the Aguja Formation of northern Coahuila, Mexico that is not currently diagnosable to the generic level, but likely represents a new taxon. Further, we have included three-dimensional surface scans of all material attributed to this animal. Considering the large number of centrosaurines from northern Laramidia, it is likely that cladistic analyses are biased towards this faunal zone. New findings of southern centrosaurines are needed to correct this bias. This discovery expands the range of centrosaurines south to Coahuila, Mexico and adds new information to better characterize the morphology and taxonomy of centrosaurines from southern Laramidia and their evolution in comparison to their northern counterparts.

Concepts: Cladistics, Cretaceous, Dinosaur, Maastrichtian, Late Cretaceous, Triceratops, Ceratopsidae, Ceratopsia

13

Recent assessments of morphological changes in the frill during ontogeny hypothesized that the late Maastrichtian horned dinosaur Torosaurus represents the “old adult” of Triceratops, although acceptance of this finding has been disputed on several lines of evidence.

Concepts: Dinosaur, Maastrichtian, Late Cretaceous, Triceratops, Ceratopsidae, Ceratopsia, Othniel Charles Marsh, Torosaurus