Size-driven preservational and macroecological biases in the latest Maastrichtian terrestrial vertebrate assemblages of North America

The end-Cretaceous (K/Pg) mass extinction event is the most recent and well-understood of the “big five” and triggered establishment of modern terrestrial ecosystem structure. Despite the depth of research into this event, our knowledge of upper Maastrichtian terrestrial deposits globally relies primarily on assemblage-level data limited to a few well-sampled formations in North America, the Hell Creek and Lance Formations. These assemblages disproportionally affect our interpretations of this important interval. Multiple investigations have quantified diversity patterns within these assemblages, but the potential effect of formation-level size-dependent taphonomic biases and their implications on extinction dynamics remains unexplored. Here, the relationship between taphonomy and body size of the Hell Creek Formation and Lance Formation dinosaurs and mammals are quantitatively analyzed. Small-bodied dinosaur taxa (<70 kg) are consistently less complete, unlikely to be articulated, and delayed in their description relative to their large-bodied counterparts. Family-level abundance (particularly skeletons) is strongly tied to body mass, and the relative abundance of juveniles of large-bodied taxa similarly is underrepresented. Mammals show similar but nonsignificant trends. The results are remarkably similar to those from the Campanian-aged Dinosaur Park Formation, suggesting a widespread strong taphonomic bias against the preservation of small taxa, which will result in their seemingly depauperate diversity within the assemblage. This taphonomically skewed view of diversity and abundance of small-bodied taxa amid our best late Maastrichtian samples has significant implications for understanding speciation and extinction dynamics (e.g., size-dependent extinction selectivity) across the K/Pg boundary.

The oldest occurrence of brachylophosaurin hadrosaurids in Canada

Hadrosaurids are a diverse and widely distributed group of ornithischian dinosaurs that are particularly well represented in the upper Campanian Dinosaur Park Formation of the Belly River Group of Alberta. However, the origin of this hadrosaurid diversity in Alberta is poorly understood, as the lower Campanian terrestrial deposits of the underlying Oldman and Foremost formations of the group have produced comparatively few body fossils. Here we provide the first description of a partially articulated hadrosaurid and hadrosaurid material from a bonebed from the Foremost Formation and refer it to the brachylophosaurin Probrachylophosaurus sp. indet. The material represents the oldest occurrence of Brachylophosaurini in Alberta and the oldest known hadrosaurid diagnostic to the genus level from Canada. In Alberta, Hadrosaurinae display a distinct pattern of replacement with the tribes Brachylophosaurini and Kritosaurini being confined to the lower to middle Campanian strata (below the marine Bearpaw Formation) and replaced above the Bearpaw Formation by members of Saurolophini (Prosaurolophus, Saurolophus) and Edmontosaurini (Edmontosaurus), with the latter clade persisting to the end of the Maastrichtian. Although the worldwide stratigraphic distribution of the Hadrosaurinae is complex, this pattern generally holds true for northern Laramidian hadrosaurine tribes, suggesting that their pattern of evolution and replacement may be driven by some common underlying factor such as an environmental response to fluctuations in the margins of the Western Interior Seaway due to sea level change.

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.

Anatomical, morphometric, and stratigraphic analyses of theropod biodiversity in the Upper Cretaceous (Campanian) Dinosaur Park Formation

The Dinosaur Park Formation (DPF) of Alberta, Canada, has produced one of the most diverse dinosaur faunas, with the record favouring large-bodied taxa, in terms of number and completeness of skeletons. Although small theropods are well documented in the assemblage, taxonomic assessments are frequently based on isolated, fragmentary skeletal elements. Here we reassess DPF theropod biodiversity using morphological comparisons, high-resolution biostratigraphy, and morphometric analyses, with a focus on specimens/taxa originally described from isolated material. In addition to clarifying taxic diversity, we test whether DPF theropods preserve faunal zonation/turnover patterns similar to those previously documented for megaherbivores. Frontal bones referred to a therizinosaur (cf. Erlikosaurus), representing among the only skeletal record of the group from the Campanian–Maastrichtian (83–66 Ma) fossil record of North America, plot most closely to troodontids in morphospace, distinct from non-DPF therizinosaurs, a placement supported by a suite of troodontid anatomical frontal characters. Postcranial material referred to cf. Erlikosaurus in North America is also reviewed and found most similar in morphology to caenagnathids, rather than therizinosaurs. Among troodontids, we document considerable morphospace and biostratigraphic overlap between Stenonychosaurus and the recently described Latenivenatrix, as well as a variable distribution of putatively autapomorphic characters, calling the validity of the latter taxon into question. Biostratigraphically, there are no broad-scale patterns of faunal zonation similar to those previously documented in ornithischians from the DPF, with many theropods ranging throughout much of the formation and overlapping extensively, possibly reflecting a lack of sensitivity to environmental changes, or other cryptic ecological or evolutionary factors.

A JUVENILE METATARSAL OF CF. DASPLETOSAURUS TOROSUS: IMPLICATIONS FOR ONTOGENY IN TYRANNOSAURID THEROPODS

A well preserved, but isolated metatarsal III of a tyrannosaurid dinosaur, originating probably from the Dinosaur Park Formation of Alberta, Canada, is tentatively referred to Daspletosaurus torosus. The size of the specimen suggests that it likely comes from a large juvenile, since the width of the distal end is about 63 % of that of a much larger individual. The morphology of the specimen supports the recently suggested hypotheses that apomorphies of tyrannosaurid taxa may have developed at young growth stages, and that juveniles of albertosaurines and tyrannosaurines may be easier to distinguish from one another than previously thought. Additionally, the specimen reported here is important in that it provides an addition to the very poor juvenile fossil record of Daspletosaurus.

A new elasmosaurid (Sauropterygia: Plesiosauria) from the non-marine to paralic Dinosaur Park Formation of southern Alberta, Canada

Elasmosaurid plesiosaurian remains have been documented from non-marine to paralic (fluvial to estuarine) sediments of the upper Campanian Dinosaur Park Formation (DPF) of southern Alberta since 1898. Despite this long collection history, this material has received relatively little research attention, largely due to the highly fragmentary nature of most recovered specimens. However, this assemblage is significant, as it constitutes a rare occurrence of plesiosaurian remains in a non-marine depositional environment. This study reports on a recently collected and prepared specimen, which represents the most complete elasmosaurid yet collected from the DPF. This specimen preserves the trunk region, the base of the neck and tail, a partial fore and hind limb, and tooth, and is sufficiently complete to be assigned as the holotype of a new genus and species. This new taxon is diagnosed by a distinctive character state combination including a boomerang-shaped clavicular arch with acute anterior process, convex anterolateral margin, deeply embayed posterior margin, and pronounced ventral keel, together with the presence of 22 dorsal vertebrae, and the anterior dorsal centra bearing a ventral notch. The DPF plesiosaurian fossils were recovered from both estuarine/bay and fluvial palaeochannel sediments. The holotype skeleton represents an osteologically mature individual with an estimated body length of around 5 m, although the largest referred DPF elasmosaurid might have been closer to 7 m, which is considerably larger than other plesiosaurians reported from non-marine deposits. This suggests small-body lengths relative to typical elasmosaurids from marine settings, but is consistent with other plesiosaurians recovered from non-marine sediments. The identification of a distinct elasmosaurid taxon in the DPF might be evidence of niche-partitioning among the predominantly oceanic members of the ubiquitous plesiosaurian clade.

A large pterosaur limb bone from the Kaiparowits Formation (late Campanian) of Grand Staircase-Escalante National Monument, Utah, USA

Pterosaurs were widespread during the Late Cretaceous, but their fossils are comparatively rare in terrestrial depositional environments. A large pterosaur bone from the Kaiparowits Formation (late Campanian, ~76–74 Ma) of southern Utah, USA, is tentatively identified as an ulna, although its phylogenetic placement cannot be precisely constrained beyond Pterosauria. The element measures over 36 cm in preserved maximum length, indicating a comparatively large individual with an estimated wingspan between 4.3 and 5.9 m, the largest pterosaur yet reported from the Kaiparowits Formation. This size estimate places the individual at approximately the same wingspan as the holotype for Cryodrakon boreas from the penecontemporaneous Dinosaur Park Formation of Alberta. Thus, relatively large pterosaurs occurred in terrestrial ecosystems in both the northern and southern parts of Laramidia (western North America) during the late Campanian.

Testing size–frequency distributions as a method of ontogenetic aging: a life-history assessment of hadrosaurid dinosaurs from the Dinosaur Park Formation of Alberta, Canada, with implications for hadrosaurid paleoecology

Hadrosaurid dinosaurs, the dominant large-bodied terrestrial herbivores in most Laurasian Late Cretaceous ecosystems, have an exceptional fossil record consisting of many species known from partial ontogenetic series, making them an ideal clade with which to conduct life-history studies. Previous research considered the Dinosaur Park Formation (DPF) of Alberta as an attritional, or time-averaged, sample and interpreted size–frequency distribution of long bones collected from the DPF with three size classes to suggest that hadrosaurids from the DPF attained near-asymptotic body size in under 3 years. This conflicted with previously published osteohistological estimates of 6+ years for penecontemporaneous hadrosaurids from the Two Medicine Formation (TMF) of Montana, suggesting either extreme variation in hadrosaurid growth rates or that size–frequency distributions and/or osteohistology and growth modeling inaccurately estimate ontogenetic age.We tested the validity of the previously proposed size–age relationship of hadrosaurids from the DPF by significantly increasing sample size and combining data from size–frequency distributions and osteohistology across multiple long-bone elements. The newly constructed size–frequency distributions typically reveal four relatively distinct size–frequency peaks that, when integrated with the osteohistological data, aligned with growth marks. The yearling size class was heavily underrepresented in the size–frequency distribution. If not due to preservation, this suggests that either juvenile (<2 years of age) hadrosaurids from the DPF had increased survivorship following an initially high nestling mortality rate or that yearlings were segregated from adults. A growth-curve analysis revealed asymptotic body size was attained in approximately 7 years, which is consistent with hadrosaurids from the TMF. The data suggest size–frequency distributions of attritional samples underestimate age and overestimate growth rates, but when paired with osteohistology can provide unique life-history insights.

Caenagnathids of the Dinosaur Park Formation (Campanian) of Alberta, Canada: anatomy, osteohistology, taxonomy, and evolution

Our understanding of caenagnathid anatomy, diversity, and ecology has improved considerably in the past twenty years, but numerous issues still remain. Among these, the diversity and taxonomy of caenagnathids from the Dinosaur Park Formation of Alberta, Canada, have remained problematic. Whereas some authors recognize three genera, others suggest only two were present, and there is considerable disagreement about which specimens are referable to which genus. This study aims to resolve this issue by reviewing the known specimens and using osteohistology, to establish a testable taxonomic framework of Dinosaur Park Formation caenagnathids. Numerous new specimens from all regions of the skeleton provide insight into morphological variation in caenagnathids, and three morphotypes are recognized based on a combination of morphological features and body size. Osteohistology shows that representatives in each body size class are at skeletal maturity, and therefore supports the delineation of three taxa: the smaller Citipes elegans gen. nov., the intermediate Chirostenotes pergracilis, and the larger Caenagnathus collinsi, new material of which shows it rivalled Anzu wyliei in size. However, these analyses also raise concerns about the referral of isolated material to each taxon in the absence of skeletal overlap between specimens or osteohistological analysis. Caenagnathids are consistently recovered throughout the Dinosaur Park Formation interval, and two geographic clusters of increased abundance probably reflect collection and taphonomic biases. The coexistence of three taxa was apparently facilitated by differences in both adult body size and functional morphology of the dentary and pes, which suggests that caenagnathids minimized niche overlap rather than subdividing niche space. Regardless, little is known of the exact roles caenagnathids played in Late Cretaceous ecosystems. Incorporation of the new material and taxonomic framework into a phylogenetic analysis drastically improves our understanding of the relationships between caenagnathines, and sheds light on the evolution of body size in caenagnathids and its role in their diversification.

An ornithurine bird coracoid from the Late Cretaceous of Alberta, Canada

The Cretaceous birds of Alberta are poorly known, as skeletal elements are rare and typically consist of fragmentary postcranial remains. A partial avian coracoid from the upper Campanian Dinosaur Park Formation of Alberta, Canada, can be referred to the Ornithurae, and is referred to here as Ornithurine G (cf. Cimolopteryx). Its structure is similar to previously described ornithurine coracoids from Alberta and other localities in North America, particularly those belonging to the genus Cimolopteryx. A comparison of these elements indicates that the new coracoid is distinct; however, its preservation prevents complete diagnosis. As other Cimolopteryx are Maastrichtian in age, Ornithurine G (cf. Cimolopteryx) also represents the earliest occurrence of a Cimolopteryx-like anatomy. A pneumatized coracoid is a diagnostic trait of Neornithes, identified by the presence of a pneumatic foramen. Ornithurine G (cf. Cimolopteryx) does not preserve this feature. CT and micro-CT scans of both pneumatic and apneumatic coracoids of modern birds show similar internal structures to Ornithurine G (cf. Cimolopteryx), indicating that pneumaticity of the coracoid cannot be determined in the absence of an external pneumatic foramen. A comparison between members of Cimolopterygidae, including Cimolopteryx and Lamarqueavis, raises questions about the assignment of Lamarqueavis to the Cimolopterygidae, and the validity of this family as a whole.