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We provide a reassessment of the hypothesis of ammonite survival across the Cretaceous–Paleogene (Maastrichtian–Danian) boundary, based on new data from the lower Danian Cerithium Limestone Member at Stevns Klint, eastern Denmark. The occurrence of this unit in disjunct basins between eroded crests of uppermost Maastrichtian bryozoan mounds prompts a reconsideration of ammonite redeposition as an alternative to the survival hypothesis. We describe new ammonite specimens from the Cerithium Limestone, representing the genera Hoploscaphites , Baculites and Fresvillia . In order to elucidate the nature of these fossils, we study their local depositional settings, based on detailed, chiefly taphonomic and sedimentological (microfacies) observations. Results for the main part of the Cerithium Limestone point to the autochthonous nature of the enclosed ammonites, which implies that they are Danian survivors. Only a single individual from the lowermost part of the Cerithium Limestone is considered a reworked Maastrichtian fossil. In summary, our results confirm ammonite survival into the Danian for the bulk of the Cerithium Limestone fauna, stimulating questions for further research of what actually killed the last ammonites that lived on Earth.

We examine temporal and spatial variation in morphology of the ammonoid cephalopod Discoscaphites iris using a large dataset from multiple localities in the Late Cretaceous (Maastrichtian) of the U.S. Gulf and Atlantic Coastal Plains, spanning a distance of 2000 km along the paleoshoreline. Our results suggest that the fossil record of D. iris is consistent with no within-species net accumulation of phyletic evolutionary change across morphological traits or the lifetime of this species. Correlations between some traits and paleoenvironmental conditions as well as changes in the coefficient of variation may support limited population-scale ecophenotypic plasticity; however, where stratigraphic data are available, no directional changes in morphology occur before the Cretaceous/Paleogene (K/Pg) boundary. This is consistent with models of “dynamic” evolutionary stasis. Combined with knowledge of life-history traits and paleoecology of scaphitid ammonoids, specifically a short planktonic phase after hatching followed by transition to a nektobenthic adult stage, these data suggest that scaphitids had significant potential for rapid morphological change in conjunction with limited dispersal capacity. It is therefore likely that evolutionary mode in the Scaphitidae (and potentially across the broader ammonoid clade) follows a model of cladogenesis wherein a dynamic morphological stasis is periodically interrupted by more substantial evolutionary change at speciation events. Finally, the lack of temporal changes in our data suggest that global environmental changes had a limited effect on the morphology of ammonoid faunas during the latest Cretaceous.

Ammonites are among the best-known fossils of the Phanerozoic, yet their habitat is poorly understood. Three common ammonite families (Baculitidae, Scaphitidae, and Sphenodiscidae) co-occur with well-preserved planktonic and benthic organisms at the type locality of the upper Maastrichtian Owl Creek Formation, offering an excellent opportunity to constrain their depth habitats through isotopic comparisons among taxa. Based on sedimentary evidence and the micro- and macrofauna at this site, we infer that the 9-m-thick sequence was deposited at a paleodepth of 70–150 m. Taxa present throughout the sequence include a diverse assemblage of ammonites, bivalves, and gastropods, abundant benthic foraminifera, and rare planktonic foraminifera. No stratigraphic trends are observed in the isotopic data of any taxon, and thus all of the data from each taxon are considered as replicates. Oxygen isotope-based temperature estimates from the baculites and scaphites overlap with those of the benthos and are distinct from those of the plankton. In contrast, sphenodiscid temperature estimates span a range that includes estimates of the planktonic foraminifera and of the warmer half of the benthic values. These results suggest baculites and scaphites lived close to the seafloor, whereas sphenodiscids sometimes inhabited the upper water column and/or lived closer to shore. In fact, the rarity and poorer preservation of the sphenodiscids relative to the baculites and scaphites suggests that the sphenodiscid shells may have only reached the Owl Creek locality by drifting seaward after death.

Numerous geochemical anomalies exist at the K-Pg boundary that indicate the addition of extraterrestrial materials; however, none fingerprint volatilization, a key process that occurs during large bolide impacts. Stable Zn isotopes are an exceptional indicator of volatility-related processes, where partial vaporization of Zn leaves the residuum enriched in its heavy isotopes. Here, we present Zn isotope data for sedimentary rock layers of the K-Pg boundary, which display heavier Zn isotope compositions and lower Zn concentrations relative to surrounding sedimentary rocks, the carbonate platform at the impact site, and most carbonaceous chondrites. Neither volcanic events nor secondary alteration during weathering and diagenesis can explain the Zn concentration and isotope signatures present. The systematically higher Zn isotope values within the boundary layer sediments provide an isotopic fingerprint of partially evaporated material within the K-Pg boundary layer, thus earmarking Zn volatilization during impact and subsequent ejecta transport associated with an impact at the K-Pg. Elevated Zn isotope compositions occur in K-Pg sedimentary layers of three different depositional environments across North America and the Caribbean. The data indicate a volatilization event, and act as a robust mechanistic indicator of the meteorite impact at the end of the Cretaceous.

Sedimentary deposits in Stoddard County, southeastern Missouri, reveal a K-Pg transition sequence represented by the uppermost Maastrichtian Owl Creek Formation and the Paleocene Clayton Formation. The Clayton Formation is characterized by a basal fossiliferous coquinite that contains reworked late Maastrichtian macrofossils. Dinoflagellate biostratigraphy is used to determine the age of the coquinite layer and specifically whether or not it is an end-K tsunamite deposit resulting from the Chicxulub impact event. Results indicate a mixed assemblage of late Maastrichtian and early Danian dinocysts within the basal coquinite of the Clayton Formation. Maastrichtian dinocyst taxa identified are Riculacysta amplexa, Pierceites pentagonus, Phelodinium tricuspe and Dinogymnium sp. and dinocysts utilized as global indicators of the basal Danian, also present in the coquinite, consist of Senoniasphaera inornata, Carpatella cornuta, Damassadinium californicum, and Lanternosphaeridium reinhardtii. A gray mud occurring above the coquinite in the middle of the Clayton Formation contains the mid-Danian dinoflagellate Senegalinium iterlaaense. Collectively, these data suggest that the coquinite was deposited well after the K-Pg event but before the middle Danian. The mixed assemblage of Late Cretaceous and Paleocene dinocysts preserved in the coquinite weakens the hypothesis that it is an end-K tsunamite deposit and suggests instead that it may result from a long-term transgressive lag. We also extend the stratigraphic range of the Paleocene Senegalinium simplex downward into the uppermost Maastrichtian.

Spider dragline (major ampullate) silk outperforms virtually all other natural and manmade materials in terms of tensile strength and toughness. For this reason, the mass-production of artificial spider silks through transgenic technologies has been a major goal of biomimetics research. Although all known arthropod silk proteins are extremely large (>200 kiloDaltons), recombinant spider silks have been designed from short and incomplete cDNAs, the only available sequences. Here we describe the first full-length spider silk gene sequences and their flanking regions. These genes encode the MaSp1 and MaSp2 proteins that compose the black widow's high-performance dragline silk. Each gene includes a single enormous exon (>9000 base pairs) that translates into a highly repetitive polypeptide. Patterns of variation among sequence repeats at the amino acid and nucleotide levels indicate that the interaction of selection, intergenic recombination, and intragenic recombination governs the evolution of these highly unusual, modular proteins. Phylogenetic footprinting revealed putative regulatory elements in non-coding flanking sequences. Conservation of both upstream and downstream flanking sequences was especially striking between the two paralogous black widow major ampullate silk genes. Because these genes are co-expressed within the same silk gland, there may have been selection for similarity in regulatory regions. Our new data provide complete templates for synthesis of recombinant silk proteins that significantly improve the degree to which artificial silks mimic natural spider dragline fibers.

Background Orb-web weaving spiders and their relatives use multiple types of task-specific silks. The majority of spider silk studies have focused on the ultra-tough dragline silk synthesized in major ampullate glands, but other silk types have impressive material properties. For instance, minor ampullate silks of orb-web weaving spiders are as tough as draglines, due to their higher extensibility despite lower strength. Differences in material properties between silk types result from differences in their component proteins, particularly members of the spidroin (spider fibroin) gene family. However, the extent to which variation in material properties within a single silk type can be explained by variation in spidroin sequences is unknown. Here, we compare the minor ampullate spidroins (MiSp) of orb-weavers and cobweb weavers. Orb-web weavers use minor ampullate silk to form the auxiliary spiral of the orb-web while cobweb weavers use it to wrap prey, suggesting that selection pressures on minor ampullate spidroins (MiSp) may differ between the two groups. Results We report complete or nearly complete MiSp sequences from five cobweb weaving spider species and measure material properties of minor ampullate silks in a subset of these species. We also compare MiSp sequences and silk properties of our cobweb weavers to published data for orb-web weavers. We demonstrate that all our cobweb weavers possess multiple MiSp loci and that one locus is more highly expressed in at least two species. We also find that the proportion of β-spiral-forming amino acid motifs in MiSp positively correlates with minor ampullate silk extensibility across orb-web and cobweb weavers. Conclusions MiSp sequences vary dramatically within and among spider species, and have likely been subject to multiple rounds of gene duplication and concerted evolution, which have contributed to the diverse material properties of minor ampullate silks. Our sequences also provide templates for recombinant silk proteins with tailored properties.

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