Randall B. Irmis

Shuvosauridae is an unusual clade of Triassic poposauroid pseudosuchians from western North America and Argentina. Gracile, bipedal, and edentulous, these pseudosuchian archosaurs appear generally ...

On September 18, 1996, Grand Staircase-Escalante National Monument (GSENM) became the first national monument managed by the US Bureau of Land Management (BLM) and one of the first to protect a landscape based partly on its opportunity for scientific discovery. Its creation was a watershed moment in public land management, because to meet the mandates for its first monument, BLM opted to implement unprecedented support of resource investigations for numerous natural and cultural sciences, including establishing its first ever in-house paleontological field program. The rationale for this was taken directly from the establishing presidential proclamation (6920) which called out GSENM’s untapped paleontological treasure trove as “world-class.” The proclamation also singled out the Late Cretaceous vertebrate fossil record of the Kaiparowits Plateau, largely known at the time through the pioneering work of Drs. Jeff Eaton and Rich Cifelli, who had spent years teasing out the mammalian evolutionary story preserved within. Their work on Mesozoic mammals, alongside sporadic work by other institutions (mainly the University of Utah and Brigham Young University) in the 1970s and 1980s, demonstrated that the Kaiparowits Plateau also held a substantial macrovertebrate record that included beautifully preserved dinosaur skeletons. However, a lack of coordinated effort and the difficult nature of fieldwork in the rugged badlands led to what can only be described as desultory results. The leverage that came with monument status, including logistical and financial support provided by BLM, made this resource more accessible to the paleontological community, stimulating a sudden burst of new field research and discovery. Initial, coordinated, and collaborative fossil inventories started in 2000 by joint BLM, Utah Museum of Natural History, Museum of Northern Arizona, and Utah Geological Survey teams led to a cascade of discoveries, including sites preserving plants, invertebrates, trace fossils, microvertebrates, and macrovertebrates, contextualized by new geological insights. Many of these new fossil finds represent species entirely new to science, with some sites preserving intact snapshots of Late Cretaceous ecosystems that are unmatched globally. Unique geologic conditions resulted in spectacular preservation, sometimes even including soft tissue traces. This renaissance in North American Late Cretaceous paleontology would not have been possible without the focused resources and effort facilitated by the creation of GSENM and the subsequent prioritization of inventory and basic research in its mission. In addition to the science, the public benefits of these efforts have been immense, providing opportunities for direct involvement in the scientific process through volunteer programs, training for several generations of future paleontologists and geologists, innumerable educational programs, and exposure in national and international media outlets through articles, television, and interviews. The collaborative and far-reaching paleontological effort at GSENM has highlighted an often overlooked aspect of public lands management: the importance of US public lands for scientific discovery and education.

Monstersauria (Squamata, Anguimorpha) fossils are present in most Upper Cretaceous sedimentary basins in western North America, but despite almost a century of collection, their record remains extremely fragmentary. Here, we describe new material belonging to large-bodied monstersaurs, including a new taxon, Bolg amondol gen. et sp. nov., based on a fragmentary associated skeleton and co-occurring specimens from the middle unit of the upper Campanian Kaiparowits Formation of Grand Staircase–Escalante National Monument in southern Utah, USA. Phylogenetic analyses recover B. amondol within Monstersauria, with two unique anatomical features: fused osteoderms on the jugal and the presence of autotomy septa on the distal caudal vertebrae. Critically, B. amondol is morphologically distinct from the problematic Late Cretaceous North American monstersaur Palaeosaniwa canadensis , whereas co-occurring monstersaur vertebrae and parietals from the Kaiparowits Formation (cf. P. canadensis ) highlight a pressing need for a reassessment of this important, widespread taxon. These results offer new evidence that at least three lineages of distinct, large-bodied monstersaurian lizard were present on the palaeolandmass of Laramidia during the Campanian Stage. Importantly, B. amondol represents the most complete squamate recovered from late Campanian southern Laramidia and reveals key anatomical characteristics for future identification of isolated lizard fossil elements.

Acaenasuchus geoffreyi is a diminutive armored archosaur from the Upper Triassic Chinle Formation of northern Arizona, U.S.A., with uncertain evolutionary relationships and skeletal maturity. Known only from osteoderms, the taxon has been considered a valid taxon of aetosaur, juvenile specimens synonymous with the aetosaur Desmatosuchus spurensis, or a non-aetosaurian pseudosuchian archosaur. Here, we describe new fossils of Acaenasuchus geoffreyi that represent cranial, vertebral, and appendicular elements as well as previously unknown variations in the dorsal carapace and ventral shield. The skull bones are ornamented with the same anastomosing complex of ridges and grooves found on the paramedian and lateral osteoderms, and the appendicular skeleton resembles that of Revueltosaurus callenderi, Euscolosuchus olseni, aetosaurs, and other armored archosaurs such as erpetosuchids. Histology of osteoderms from the hypodigm of Acaenasuchus geoffreyi shows multiple growth lines, laminar tissue, and low vascularity, evidence that the individuals were close to skeletal maturity and not young juveniles. A revised phylogenetic analysis of early archosaurs recovers Acaenasuchus geoffreyi and Euscolosuchus olseni as sister taxa and members of a new clade that is the sister taxon of Aetosauria. This new phylogeny depicts a broader distribution of osteoderm character states previously thought to only occur in aetosaurs, demonstrating the danger of using only armor character states in aetosaur taxonomy and phylogeny. Acaenasuchus geoffreyi is also a good example of how new fossils can stabilize ‘wild card’ taxa in phylogenetic analyses and contributes to our understanding of the evolution of the aetosaur carapace.

Skeletal remains of a small reptile with a distinctive dentition from the Lower Keuper (Erfurt Formation; Middle Triassic, Ladinian) of the Schumann quarry near Eschenau, in the municipality of Vellberg in Baden-Württemberg, Germany, represent a new taxon of non-archosaurian archosauriforms, Polymorphodon adorfi. It is diagnosed by various craniodental autapomorphies, including mesial and distal carinae of labiolingually flattened maxillary and dentary tooth crowns with large, somewhat hook-shaped denticles aligned at distinct angle to apicobasal axis of tooth crown; premaxilla with long, leaf-shaped posterodorsal process that is slightly longer than body of element; presence of prominent lateral fossa on premaxilla anteroventral to external narial fenestra; premaxilla with five gently recurved, conical teeth; medial surface of maxilla with distinct ledge above the interdental plates; and maxilla and dentary with distinctly heterodont dentition. Phylogenetic analysis recovered Polymorphodon adorfi in a position crownward of Erythrosuchus africanus but in an unresolved polytomy with derived non-archosaurian archosauriforms such as Proterochampsidae and Euparkeria capensis and with Archosauria. The maxillary and dentary teeth of Polymorphodon adorfi differ from those of other non-archosaurian archosauriforms and indicate a different, possibly omnivorous diet, suggesting that these reptiles were more diverse in terms of feeding habits than previously assumed.

The sternum is a functionally important but poorly understood component of the pectoral girdle in both living and extinct dinosaurs. When ossified, the dinosaur sternum consists of paired plates situated just posterior to the clavicles and scapulocoracoid. In Avialae, the sternal plates are fused at the midline to form a single element, aiding in powered flight. Our understanding of the early evolutionary history of the theropod sternum is complicated by a lack of fossil material, especially outside Maniraptoriformes, both because of taphonomic biases and incomplete ossification. Here, we report the oldest-known dinosaur sternal plates from the Late Triassic taxon Tawa hallae recovered from the Hayden Quarry at Ghost Ranch, New Mexico, as well as an isolated sternal plate from just south of Petrified Forest National Park, Arizona. These specimens expand the fossil record and allow for reconstruction of early theropod sternal musculature. They bear morphological features that are surprisingly similar to sternal traits in avialans, including a sternocoracoidal process, costal processes, and possible homologs to the ‘coracoid facet’ and pila coracoidea. The presence of these features in T. hallae pushes their appearance back to the Late Triassic, well before the development of the modern avian sternum. The variable distribution of sternal character states across avemetatarsalians was previously recognized in later Cretaceous avialans and can now be shown to also characterize the early evolution of the theropod sternum.

The Upper Triassic Dockum Group of Garza County, Texas (lower, middle, and upper Cooper Canyon Formation) captures the radiation of Triassic non-marine tetrapods by preserving a variety of Late Triassic taxa from the southwestern United States. Our understanding of the vertebrate assemblage from these strata largely comes from a single site, the Post Quarry (lower Cooper Canyon Formation), with previous research documenting a variety of temnospondyls, sphenodontians, non-archosauriform archosauromorphs, and archosauriforms including a phytosaur, three species of aetosaurs, a poposauroid, a rauisuchid, a crocodylomorph, and several dinosauromorphs. To more completely reconstruct the vertebrate assemblage of the Dockum Group of Garza County we use an apomorphy-based approach to identify morphologically similar disarticulated and fragmentary elements from a variety of localities that span the entire Cooper Canyon Formation (Norian-Rhaetian), allowing assignments from the large clade level to the species level. Many skeletal elements are incomplete yet diagnostic and are assigned to the least inclusive clade if discrete character states do not allow for an unambiguous species-level identification. We identify new specimens referable to numerous clades including Tanystropheidae, Allokotosauria + Prolacerta + Archosauriformes, Vancleavea + Litorosuchus, Phytosauria, Paracrocodylomorpha, Dinosauriformes, and Saurischia, in addition to additional species identifications of the aetosaur Scutarx deltatlyus, and the dinosauromorph Dromomeron gregorii. Our study of this material demonstrates the utility of an apomorphy-based approach in making testable and repeatable observations for identifying small, isolated fragmentary fossil tetrapod material to reconstruct a more accurate faunal hypothesis for a portion of the Late Triassic of Texas. Previous claims of the earliest dinosaurs from near the base of the Dockum Group do not pass the apomorphy-based identification test, and the question of whether the oldest known North American dinosaurs are present in the Chinle Formation or Dockum Group can be resolved by utilizing vertebrate biostratigraphic correlation. Our revision of these fossil assemblages supports the hypothesis that early diapsids, early archosauromorphs, and non-dinosaurian dinosauromorphs were more common, diverse, and widespread in low latitudes during this time than previously thought.

Trialestes romeri (Reig) is an early crocodylomorph from the Ischigualasto Formation (late Carnian – early Norian; Ischigualasto – Villa Unión Basin, Argentina) and one of the oldest‐known members of this clade. Two specimens of this species are known, the holotype (PVL 2561) and a referred specimen (PVL 3889), both consisting of associated cranial and postcranial remains. These specimens are incomplete and poorly preserved thus leading previous authors to propose different phylogenetic hypotheses for the species. Trialestes romeri was originally interpreted as a basal dinosaur, subsequently considered to be a crocodylomorph with some dinosaurian characters (e.g. mesotarsal ankle joint, functionally tridactyl pes), and even proposed as a chimera representing two different genera. Some recent workers have mentioned and discussed Trialestes, but none have described it in detail or included it in a quantitative phylogenetic analysis. Here, we describe in detail all the material assignable to the species and test its phylogenetic relationships using a comprehensive data matrix focused on early archosaurs. We support the referral of PVL 3889 to Trialestes and reject the presence of a mesotarsal ankle joint in this specimen. We recovered Trialestes within Crocodylomorpha, closer to Crocodyliformes than Pseudhesperosuchus, Hesperosuchus, Dromicosuchus and Sphenosuchus. Several stratigraphical units worldwide approximately coeval with the Ischigualasto Formation, have also yielded early crocodylomorph remains, such as the upper Santa María (Brazil), Pekin (USA), and lower Maleri (India) formations. However, the crocodylomorph specimens recovered from those units are more incomplete, possess uncertain phylogenetic affinities, or are part of chimaerae. Therefore, Trialestes represents the most completely known of the earliest non‐crocodyliform crocodylomorph taxa known to date.

Significance: Many hypotheses have been put forth to explain the origin and early radiation of dinosaurs, but poor age constraints for constituent fossil assemblages make these scenarios difficult to test. Using precise radioisotopic ages, we demonstrate that the temporal gap between assemblages containing only dinosaur precursors and those with the first dinosaurs was 5–10 million years shorter than previously thought. Thus, these data suggest that the origin of dinosaurs was a relatively rapid evolutionary event. Combined with our synthesis of paleoecological data, we demonstrate there was little compositional difference between the dinosaur precursor assemblages and the earliest dinosaur assemblages, and thus, the initial appearance of dinosaurs was not associated with a fundamental shift in ecosystem composition, as classically stated.