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>

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.

Hagfish depart so much from other fishes anatomically that they were sometimes considered not fully vertebrate. They may represent: (i) an anatomically primitive outgroup of vertebrates (the morphology-based craniate hypothesis); or (ii) an anatomically degenerate vertebrate lineage sister to lampreys (the molecular-based cyclostome hypothesis). This systematic conundrum has become a prominent case of conflict between morphology- and molecular-based phylogenies. To date, the fossil record has offered few insights to this long-branch problem or the evolutionary history of hagfish in general, because unequivocal fossil members of the group are unknown. Here, we report an unequivocal fossil hagfish from the early Late Cretaceous of Lebanon. The soft tissue anatomy includes key attributes of living hagfish: cartilages of barbels, postcranial position of branchial apparatus, and chemical traces of slime glands. This indicates that the suite of characters unique to living hagfish appeared well before Cretaceous times. This new hagfish prompted a reevaluation of morphological characters for interrelationships among jawless vertebrates. By addressing nonindependence of characters, our phylogenetic analyses recovered hagfish and lampreys in a clade of cyclostomes (congruent with the cyclostome hypothesis) using only morphological data. This new phylogeny places the fossil taxon within the hagfish crown group, and resolved other putative fossil cyclostomes to the stem of either hagfish or lamprey crown groups. These results potentially resolve the morphological-molecular conflict at the base of the Vertebrata. Thus, assessment of character nonindependence may help reconcile morphological and molecular inferences for other major discords in animal phylogeny.

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