The Cretaceous-Paleogene (KPg) boundary is demarked by two critical and intimately linked events: a global-scale impact caused by a massive extraterrestrial body, and a resultant catastrophic planet-wide mass-extinction, with pivotal long-term consequences for life on Earth. The site of impact has been identified as the ~180 km wide Chicxulub crater complex [Yucatan peninsula, Mexico] based on strong geochemical, sedimentological, and temporal evidence. However, the impactor, which was obliterated on contact during the cratering process, has eluded identification. Previous studies struggled to identify the projectile based on scant geochemical and isotopic traces admixed with the ejecta and melt-rock, plus a single heavily altered microscopic fragment possibly from the impactor. While those efforts helped to rule out some potential sources and narrowed down the best potential candidates to a CM, CR, or CO carbonaceous chondrite, the data was insufficient to provide an incontrovertible diagnosis, and the identity of the Chicxulub projectile remained a critical missing detail. Here we describe new, exceptionally-preserved fragments of cosmic origin that were directly associated with Chicxulub impact ejecta from a temporally constrained KPg boundary site in North Dakota (U.S.A.). The fragments occur as inclusions within unaltered glassy ejecta spherules, which likely protected them against chemical and physical degradation, leading to their marked preservation. The spherules themselves were also uniquely preserved in amber, which inhibited their usual breakdown to smectitic clay. Geochemical examination of the inclusions were undertaken using electron microprobe, laser-ablation inductively-coupled-plasma-mass-spectroscopy, and synchrotron X-ray techniques. Results provide multiple lines of evidence that support a cosmic origin and chemistry indicative of a CM subtype carbonaceous chondrite, offering new support of prior hypotheses. The data is also in agreement with projectile residues from the Chicxulub impact fallout. In addition to its utility in reconstructing the dynamics of the Chicxulub event, this data helps provide extra dimension to our understanding of major impact processes and their effects on Earth.

<p>Dinosaurs and pterosaurs, critical components of late Mesozoic terrestrial biomes, have thus far never been reported in-situ from deposits immediately below the Cretaceous-Paleogene (KPg) boundary. Strata from the KPg boundary preserve worldwide evidence of a massive cosmic impact that resulted in grave consequences for Earth’s biota, triggering a rapid global mass extinction that ultimately claimed ~75% of species. While trends in long-term biotic effects are well-documented, little is known about the fate of biota coeval with the impact because well-preserved fossil evidence is lacking for that brief time interval. Here we report the first-known occurrences of in-situ dinosaur and pterosaur remains coincident with the KPg boundary, providing a vantage point that is closer in temporal proximity to the Chicxulub impact than any prior known records for these clades. The fossils, preserved in a Chicxulub impact-triggered surge deposit and mass-death assemblage in the Hell Creek Formation, U.S.A., consist of a well-developed semi-articulated prenatal pterosaur in ovum and partially articulated remains of a subadult ornithischian dinosaur (Family Thescelosauridae). Examination via synchrotron rapid scanning X-ray fluorescence (SRS-XRF), scanning electron microscopy (SEM), and traditional light microscopy revealed extensive preservation of soft tissue consisting of distinct signatures of organic residues as well as three-dimensional structure. The pterosaur embryo, which comprises the first known from the Late Cretaceous and the only example thus far from North America, contributes information pivotal to understanding the prenatal development and early ontogeny of these animals from a time interval for which no such data exists. The ornithischian (c.f. <em>Thescelosaurus</em> sp.) dinosaur, which includes a remarkably complete articulated hind limb enclosed within three-dimensional lithified tubercular skin envelope, provides important data regarding the osseous and soft-tissue anatomy of the Thescelosauridae. In addition to contributing invaluable new anatomical and physiological insights, these specimens constitute the first conclusive evidence of the direct participation of pterosaurs and dinosaurs in the terminal-Cretaceous extinction event.</p>

Ichthyornis has long been recognized as a pivotally important fossil taxon for understanding the latest stages of the dinosaur-bird transition, but little significant new postcranial material has been brought to light since initial descriptions of partial skeletons in the 19th Century. Here, we present new information on the postcranial morphology of Ichthyornis from 40 previously undescribed specimens, providing the most complete morphological assessment of the postcranial skeleton of Ichthyornis to date. The new material includes four partially complete skeletons and numerous well-preserved isolated elements, enabling new anatomical observations such as muscle attachments previously undescribed for Mesozoic euornitheans. Among the elements that were previously unknown or poorly represented for Ichthyornis, the new specimens include an almost-complete axial series, a hypocleideum-bearing furcula, radial carpal bones, fibulae, a complete tarsometatarsus bearing a rudimentary hypotarsus, and one of the first-known nearly complete three-dimensional sterna from a Mesozoic avialan. Several pedal phalanges are preserved, revealing a remarkably enlarged pes presumably related to foot-propelled swimming. Although diagnosable as Ichthyornis, the new specimens exhibit a substantial degree of morphological variation, some of which may relate to ontogenetic changes. Phylogenetic analyses incorporating our new data and employing alternative morphological datasets recover Ichthyornis stemward of Hesperornithes and Iaceornis, in line with some recent hypotheses regarding the topology of the crownward-most portion of the avian stem group, and we establish phylogenetically-defined clade names for relevant avialan subclades to help facilitate consistent discourse in future work. The new information provided by these specimens improves our understanding of morphological evolution among the crownward-most non-neornithine avialans immediately preceding the origin of crown group birds.

The end-Cretaceous Chicxulub impact triggered Earth's last mass-extinction, extinguishing ~ 75% of species diversity and facilitating a global ecological shift to mammal-dominated biomes. Temporal details of the impact event on a fine scale (hour-to-day), important to understanding the early trajectory of mass-extinction, have largely eluded previous studies. This study employs histological and histo-isotopic analyses of fossil fish that were coeval with a unique impact-triggered mass-death assemblage from the Cretaceous-Paleogene (KPg) boundary in North Dakota (USA). Patterns of growth history, including periodicity of ẟ18O and ẟ13C and growth band morphology, plus corroborating data from fish ontogeny and seasonal insect behavior, reveal that the impact occurred during boreal Spring/Summer, shortly after the spawning season for fish and most continental taxa. The severity and taxonomic symmetry of response to global natural hazards are influenced by the season during which they occur, suggesting that post-impact perturbations could have exerted a selective force that was exacerbated by seasonal timing. Data from this study can also provide vital hindsight into patterns of extant biotic response to global-scale hazards that are relevant to both current and future biomes.

The flat-end surfaces of dinosaur vertebral centra led to the presumption that intervertebral discs occupied the space between their vertebrae. A set of fused hadrosaur vertebrae allowed that hypothesis to be tested. The Tyrannosaurus rex responsible for this pathology did not escape unscathed. It left behind a tooth crown that had fractured. Fragments of that tooth were scattered through the intervertebral space, evidencing that there was no solid structure to impede its movement. That eliminates the possibility of an intervertebral disc and instead proves the presence of an articular space, similar to that in modern reptiles, but at variance to what is noted in birds. While avian cervical vertebral centra appear to be separated by diarthrodial joints, the preponderance of their thoracic vertebral centra is not separated by synovial joints.

SignificanceThe Chicxulub impact played a crucial role in the Cretaceous–Paleogene extinction. However the earliest postimpact effects, critical to fully decode the profound influence on Earth’s biota, are poorly understood due to a lack of high-temporal-resolution contemporaneous deposits. The Tanis site, which preserves a rapidly deposited, ejecta-bearing bed in the Hell Creek Formation, helps to resolve that long-standing deficit. Emplaced immediately (minutes to hours) after impact, Tanis provides a postimpact “snapshot,” including ejecta accretion and faunal mass death, advancing our understanding of the immediate effects of the Chicxulub impact. Moreover, we demonstrate that the depositional event, calculated to have coincided with the arrival of seismic waves from Chicxulub, likely resulted from a seismically coupled local seiche.

A set of associated left pedal elements of a sauropod dinosaur from the Upper Jurassic Morrison Formation in Weston County, Wyoming, is described here. Several camarasaurids, a nearly complete small brachiosaur, and a small diplodocid have been found at this locality, but none match the exceptionally large size of the pedal elements. Next to the associated pedal elements, an isolated astragalus, phalanx and ungual were found, which match the large metatarsals in size. The elements cannot be ascribed to diplodocids due to the lack of a ventral process of metatarsal I. Moreover, the morphology of metatarsal V has a broad proximal end, with a long and narrow distal shaft, which differs from Camarasaurus . The size of the material and a medially beveled distal articular surface of metatarsal IV imply an identification as a brachiosaurid. This is the largest pes ever reported from a sauropod dinosaur and represents the first confirmed pedal brachiosaur elements from the Late Jurassic of North America. Furthermore, this brachiosaur material (the pes and the small nearly complete specimen) is the northernmost occurrence of brachiosaurids in the Morrison Formation.

The skull of living birds is greatly modified from the condition found in their dinosaurian antecedents. Bird skulls have an enlarged, toothless premaxillary beak and an intricate kinetic system that includes a mobile palate and jaw suspensorium. The expanded avian neurocranium protects an enlarged brain and is flanked by reduced jaw adductor muscles. However, the order of appearance of these features and the nature of their earliest manifestations remain unknown. The Late Cretaceous toothed bird Ichthyornis dispar sits in a pivotal phylogenetic position outside living groups: it is close to the extant avian radiation but retains numerous ancestral characters1–3. Although its evolutionary importance continues to be affirmed3–8, no substantial new cranial material of I. dispar has been described beyond incomplete remains recovered in the 1870s. Jurassic and Cretaceous Lagerstätten have yielded important avialan fossils, but their skulls are typically crushed and distorted9. Here we report four three-dimensionally preserved specimens of I. dispar—including an unusually complete skull—as well as two previously overlooked elements from the Yale Peabody Museum holotype, YPM 1450. We used these specimens to generate a nearly complete three-dimensional reconstruction of the I. dispar skull using high-resolution computed tomography. Our study reveals that I. dispar had a transitional beak—small, lacking a palatal shelf and restricted to the tips of the jaws—coupled with a kinetic system similar to that of living birds. The feeding apparatus of extant birds therefore evolved earlier than previously thought and its components were functionally and developmentally coordinated. The brain was relatively modern, but the temporal region was unexpectedly dinosaurian: it retained a large adductor chamber bounded dorsally by substantial bony remnants of the ancestral reptilian upper temporal fenestra. This combination of features documents that important attributes of the avian brain and palate evolved before the reduction of jaw musculature and the full transformation of the beak. High-resolution computed tomography of three-dimensionally preserved specimens of Ichthyornis dispar clarifies the mosaic evolution of the avian head, revealing a kinetic feeding apparatus reminiscent of modern birds, a transitional beak and a dinosaurian temporal region.