Integrated biostratigraphy based on planktonic foraminifera and dinoflagellates across the Cretaceous/Paleogene (K/Pg) transition at the Izeh section (SW Iran)

<p>After 42 years, the debate over the end-Cretaceous mass extinction still rages with arguments made for Chicxulub impact and Deccan volcanism as the real cause of this catastrophe. We briefly review the evidence for the pre-KPB age of the Chicxulub impact based on the primary impact spherule layer, which we link to Deccan volcanism based on the global mercury (Hg) fallout from Deccan eruptions. Mercury from volcanic eruptions is distributed around the world during its atmospheric residence time of 6 months to one year, after which it rains out over land and oceans. Major pulsed volcanic eruptions yield high Hg concentrations during fallout, which we termed Extreme Events (EE). We identified 20 of these Hg extreme events during the last 550 ky of the late Maastrichtian in sequences from Tunisia, Israel, Egypt and Mexico. At Elles, Tunisia, we dated these events (EE1 to EE20) based on orbital cyclicity and biostratigraphy with precision of one cycle (20 ky) with an error margin of 10-20 ky (Keller et al., 2020). We linked these dates to U-Pb zircon ages of the Deccan Traps with similarly high precision (Schoene et al., 2019). The resulting mercury stratigraphy yielded excellent age control linking Deccan eruption pulses across the globe. Results from two localities in NE Mexico revealed the Chicxulub impact crashed into Yucatan above the base of the <em>Plummerita hantkeninoides</em> zone CF1 and EE6 at about 200 ky prior to the KPB mass extinction. This deposit is unlike any other of the over 100 reworked spherule layers mixed with abundant shallow water debris. This oldest and primary impact spherule layer consists of compressed pure melt rock glass and glass spherules that settled rapidly to the deep seafloor. The environmental effects of this large impact were short-lived and caused no species extinctions. The effects of this 10 km-sized bolide impact had been vastly overrated.</p><p>The KPB mass extinction was identified between the longest lava flows across India to the Krishna-Godavari Basin and into the Bay of Bengal. Based on peak Hg fallout, we identified these volcanic eruptions as the largest most rapid sequence of pulsed events in Tunisia, Israel, Egypt and Mexico, all coinciding with the rapid mass extinction observed in India. The mass extinction began with the onset and ramp-up of pulsed Deccan eruptions resulting in toxic and acidic waters that caused 50% species extinctions. Extremely rapid large pulsed eruptions followed and resulted in the longest lave flows and hyperthermal warming that caused the rapid demise of all but one species, the disaster opportunist <em>Guembelitria cretacea</em>. Deccan eruptions quickly diminished after the mass extinction and climate cooled rapidly giving rise to the first new species. Volcanic eruptions remained low and cool temperatures persisted through the early Paleocene interrupted by a smaller eruption phase about 100 ky after the mass extinction.  These data reveal that Deccan volcanism caused the KPB mass extinction without any extraterrestrial aid.</p><p>Keywords: Chicxulub, Deccan Volcanism, Mass Extinction, Mercury Stratigraphy, Age control</p><p> </p>

The human race faces many global to local challenges in the near future. Among these are massive biodiversity losses. The 2012 IUCN/SSC Red List reported evaluations of ~56 % of all vertebrates. This included 97 % of amphibians, mammals, birds, cartilaginous fishes, and hagfishes. It also contained evaluations of ~50 % of lampreys, ~38 % of reptiles, and ~29 % of bony fishes. A cursory examination of extinction magnitudes does not immediately reveal the severity of current biodiversity losses because the extinctions we see today have happened in such a short time compared to earlier events in the fossil record. So, we still must ask how current losses of species compare to losses in mass extinctions from the geological past. The most recent and best understood mass extinction is the Cretaceous terminal extinction which ends at the Cretaceous–Paleogene (K–Pg) border, 65 MYA. This event had massive losses of biodiversity (~17 % of families, >50 % of genera, and >70 % of species) and exterminated the dinosaurs. Extinction estimates for non-dinosaurian vertebrates at the K–Pg boundary range from 36 to 43 %. However, there remains much uncertainty regarding the completeness, preservation rates, and extinction magnitudes of the different classes of vertebrates. Fuzzy arithmetic was used to compare recent vertebrate extinction reported in the 2012 IUCN/SSC Red List with biodiversity losses at the end of K–Pg. Comparisons followed 16 different approaches to data compilation and 288 separate calculations. I tabulated the number of extant and extinct species (extinct + extinct in the wild), extant island endemics, data deficient species, and so-called impaired species [species with IUCN/SSC Red List designations from vulnerable (VU) to critically endangered (CR)]. Species that went extinct since 1500 and since 1980 were tabulated. Vertebrate extinction moved forward 24–85 times faster since 1500 than during the Cretaceous mass extinction. The magnitude of extinction has exploded since 1980, with losses about 71–297 times larger than during the K–Pg event. If species identified by the IUCN/SSC as critically endangered through vulnerable, and those that are data deficient are assumed extinct by geological standards, then vertebrate extinction approaches 8900–18,500 times the magnitude during that mass extinction. These extreme values and the great speed with which vertebrate biodiversity is being decimated are comparable to the devastation of previous extinction events. If recent levels of extinction were to continue, the magnitude is sufficient to drive these groups extinct in less than a century.

<p>Recent multi-disciplinary efforts demonstrate a correlation between continental flood basalt (CFB) volcanism and major environmental catastrophes associated with four out of the five largest Phanerozoic mass extinctions. Unique among these is the end-Cretaceous mass extinction, which is potentially coincident with both the Chicxulub bolide impact and the Deccan volcanism. Among these two drivers, the role of the Deccan volcanism is crucial in order to decipher if there is a causal relationship between volcanism and environmental stress, and if so, how stressed the environment was during the latest Maastrichtian. To assess the cause-and-effect relationship between Deccan volcanism and climate change and mass extinctions, high-resolution biostratigraphy, quantitative species analysis coupled with geochemical measurements have been performed on complete sections of Mudurnu-Göynük and Haymana basins (Turkey).</p><p>In both basins Maastrichtian sedimentation is characterized by monotonous mudstones, which sharply in turn to marl-calcareous mudstone alternations in the earliest Danian. Detailed quantitative study on planktonic foraminifera of the Haymana Basin revealed that planktonic foraminiferal community in the latest Maastrichtian is dominated by ecological generalists with small, simple morphologies (e.g., Heterohelix, Globigerinelloides, Guembelitria). Among them low oxygen tolerant Heterohelix globulosa is the most dominant taxa and their abundance changing with the presence of stress marker Guembelitria cretacea. In all sections, the K/Pg boundary itself is characterized by 2-3 mm thick reddish oxidized layer which corresponds to sudden annihilation of large, ornamented ecological specialists (e.g., Globotruncana, Rugoglobigerina, Racemiguembelina). Right after the boundary, there is an acme of calcareous dinoflagellate cysts (Thoracosphaera) and a surge of Guembelitria cretacea indicate ecosystem collapse in post-K/Pg environment.</p><p>On the other hand, detailed quantitative analysis shows a systematic reduction in the species richness throughout the Plummerita hantkeninoides Zone corresponding to the final 150 kyr of the Cretaceous. Proliferations of the Guembelitria cretacea through late Maastrichtian is known as an indicator of high terrigenous influx; therefore, enhanced food resources. The high sedimentation rates observed in all the studied sections might be linked to increased greenhouse conditions due to Deccan volcanism leading to enhanced weathering. Overall, our multiproxy approach including quantitative biostratigraphy and geochemical analyses highlights the influence of the Deccan volcanism by releasing high amounts of atmospheric CO<sub>2</sub> and SO<sub>2</sub>, leading to the climatic changes and associated biotic stress, which predisposed faunas to eventual extinction at the K/Pg boundary.</p>

Deccan volcanism induced high-stress environment during the Cretaceous–Paleogene transition at Zumaia, Spain: Evidence from magnetic, mineralogical and biostratigraphic records

Paleoecology of the Cretaceous–Tertiary mass extinction in planktonic foraminifera

Dinoflagellate cyst evidence for the age, palaeoenvironment and paleoclimate of a new Cretaceous–Paleogene (K/Pg) boundary section at the Bou Angueur syncline, Middle Atlas, Morocco

<p indent="0mm">The Cretaceous-Paleogene (K-Pg) mass extinction that ended the age of the dinosaurs occurred 66 million years ago, with competing hypotheses including the Deccan Traps volcanic eruption, climate change, and sea level change. However, it is the Chicxulub asteroid impact that has been widely accepted as the most plausible cause. Whatever the reason, the Earth’s ecological environment changed dramatically, leading to a mass extinction of both terrestrial and marine life. Primary producers lie at the base of the food chain and support higher trophic levels, so play an important role in the marine ecosystem. The predominant primary producers in the ocean, coccolithophores (phytoplankton with small CaCO₃ scales covering their cell surface), were among those most severely devastated by the K-Pg mass extinction. So, studying fossil coccolithophores across the K-Pg boundary can provide a better understanding of the destruction and post-event recovery processes of marine primary production (PP) and export production (EP), and the changes and controlling mechanisms of the marine biological carbon pump in the current global climate change. The four hypotheses previously proposed to interpret the post-event primary production, namely strangelove ocean (collapsed PP), living ocean (collapsed EP), resilient ocean (continued PP and EP), and heterogeneous ocean (geography-dependent responses of PP and EP), are illustrated and compared, and their pros and cons reviewed. A dataset is compiled from multiple locations in the global ocean basin including Deep Sea Drilling Program Site 465, Ocean Drilling Program Sites 690, 1210, and 1262, and the outcrop at El Kef (Tunisia), with each location having a high-precision cyclostratigraphic time control and an ocean surface-to-deep (bulk carbonate-benthic foraminifera) <italic>δ</italic>¹³C vertical gradient. This global dataset is analyzed to explore the changes in the efficiency of global marine biological carbon pump after the K-Pg boundary event. Three important patterns are observed: (1) Prior to the K-Pg event, the vertical <italic>δ</italic>¹³C gradient in all ocean basins falls between 0.8‰–1.5‰ and is consistent with that in the modern ocean, and this gradient seems to gradually increase from the shelf to the deep ocean possibly due to higher efficiency of the biological carbon pump in the open ocean commonly characterized by oligotrophy; (2) the global ocean vertical <italic>δ</italic>¹³C gradient collapses after the K-Pg event, suggesting drastic changes in primary and/or export production in the euphotic zone; (3) the vertical <italic>δ</italic>¹³C gradient collapse results from the abrupt negative shift of the surface ocean <italic>δ</italic>¹³C, and the magnitude of this negative shift gradually decreases from the continental shelf to the deep sea, which may be related to the selective extinction of shallow-water grazers, swimming organisms and plankton, as well as the limited buffering capacity of shelf environments. After the K-Pg event, the main primary producers change from larger coccolithophores to smaller picoplankton, and the more recalcitrant dissolved organic carbon is able to sink to the seafloor due to the operation of the microbial carbon pump in the photic zone, leading to sustained efficiency of the biological carbon pump. Although the picoplankton-dominated biological carbon pump is not as efficient as that by coccolithophores, it is still capable of removing enough nutrients and harmful substances from the photic zone via long-term operation, not only providing food for higher trophic level organisms, but improving the marine physicochemical environment that deteriorated after the K-Pg asteroid impact. This process paves the way for the restoration and development of the entire marine ecosystem. The switch of key marine primary producers and the controlling factors involved are critical scientific issues that deserve more focus in future studies of the deep-time marine carbon cycle. The regional or global variation patterns in the biological carbon pump after the K-Pg event reveal how different marine ecosystems operate under extreme environmental stress, and provide a reference for better understanding and predicting how they respond to the ongoing rapid global climate change.

A Cretaceous–Palaeogene boundary geological site, revealed by planktic foraminifera and dinoflagellate cysts, at Ouled Haddou, eastern external Rif Chain, Morocco

Recent advances in Deccan volcanic studies indicate three volcanic phases with the phase-1 at 67.5 Ma followed by a 2 m.y. period of quiescence. Phase-2 marks the main Deccan volcanic eruptions in Chron 29r near the end of the Maastrichtian and accounts for approximately 80% of the entire 3500 m thick Deccan lava pile. At least four of the world's longest lava flows spanning 1000 km across India and out into the Gulf of Bengal mark phase-2. The final phase-3 was smaller, coincided with the early Danian Chron 29n and also witnessed several of the longest lava flows. The KT boundary and mass extinction was first discovered based on planktic foraminifera from shallow marine intertrappean sediments exposed in Rajahmundry quarries between the longest lava flows of the main volcanic phase-2 and smaller phase-3. At this locality early Danian (zone P1a) planktic foraminiferal assemblages directly overlie the top of phase-2 eruptions and indicate that the masse extinction coincided with the end of this volcanic phase. Planktic foraminiferal assemblages also mark the KT boundary in intertrappean sediments at Jhilmili, Chhindwara, where freshwater to estuarine conditions prevailed during the early Danian and indicate the presence of a marine seaway across India at KT time. Dinosaur bones, nesting sites with complete eggs and abundant eggshells are known from central India surrounding the hypothesized seaway through the Narmada-Tapti rift zone. A Maastrichtian age is generally assigned to these dinosaur remains. Age control may now be improved based on marine microfossils from sequences deposited in the seaway and correlating these strata to nearby terrestrial sequences with dinosaur remains.

Biotic effects of impacts and volcanism

Mass extinction events (MEEs), defined as significant losses of species diversity in significantly short time periods, have attracted the attention of biologists because of their link to major environmental change. MEEs have traditionally been studied through the fossil record, but the development of birth‐death models has made it possible to detect their signature based on extant‐taxa phylogenies. Most birth‐death models consider MEEs as instantaneous events where a high proportion of species are simultaneously removed from the tree (“single pulse” approach), in contrast to the paleontological record, where MEEs have a time duration. Here, we explore the power of a Bayesian Birth‐Death Skyline (BDSKY) model to detect the signature of MEEs through changes in extinction rates under a “time‐slice” approach. In this approach, MEEs are time intervals where the extinction rate is greater than the speciation rate. Results showed BDSKY can detect and locate MEEs but that precision and accuracy depend on the phylogeny's size and MEE intensity. Comparisons of BDSKY with the single‐pulse Bayesian model, CoMET, showed a similar frequency of Type II error and neither model exhibited Type I error. However, while CoMET performed better in detecting and locating MEEs for smaller phylogenies, BDSKY showed higher accuracy in estimating extinction and speciation rates.

The Cretaceous–Paleogene (K/Pg) boundary in the Haymana Basin, Central Anatolia, Turkey, was delineated using planktonic foraminiferal biostratigraphy, microfacies analysis, and sequence stratigraphy. An ∼ 29 m outcrop consisting of limestone and marl was measured, and four planktonic foraminiferal biozones were identified spanning the boundary. Planktonic foraminiferal extinction across the K/Pg boundary was catastrophic and abrupt. The extinction level is overlain by a unit (Zone P0) showing an increase in echinoid fecal pellets and authigenic clay minerals such as glauconite, suggesting low sedimentation rates in the early Danian. Ten microfacies types were identified indicating inner-ramp to basinal paleoenvironments based on the sedimentological characteristics and microfossil and macrofossil assemblages. Maastrichtian carbonates contain large benthic foraminifera, calcareous red algae, bryozoans, fragments of echinoderms and mollusks, and planktonic foraminifera. Overlying Maastrichtian–Danian silty marls and silty limestones have common planktonic and benthic foraminifera. Progradation of carbonates into the basin took place during the highstand systems tract, and deposition of a silty marl succession occurred during the transgressive systems tract. The K/Pg boundary is in the upper part of the transgressive systems tract, below a maximum flooding surface. Sequence stratigraphic analysis of a second section, Campo Pit, New Jersey, USA, showed that the K/Pg boundary occurs within a transgressive systems tract in New Jersey as well, suggesting a global sea-level rise across the K/Pg boundary.

Approximate periodicity for peak rates of global extinction during the past 250 m.y. may have resulted from delayed recovery following major extinction events. Two components can be envisioned for such delays: persistence of inimical environmental conditions for some time after the onset of the crisis, and slow restoration of vulnerable taxa. This general hypothesis is consistent with statistical evidence of linkage between measured rates of extinction of marine invertebrate genera for contiguous stages and substages of the geologic column. The nine broad valleys between the “periodic” peak rates for the past 250 m.y. exhibit only three trivial secondary peaks, indicating that, if the pattern is not artifactual, trends in global rates of extinction have not readily been abruptly reversed. Moreover, the smooth observed trends reflect the fact that regional crises tend to remove many species but few genera. To some degree, high rates of extinction that precede peak rates must represent bias of the incomplete fossil record (the Signor-Lipps effect). High rates that immediately follow peak rates also may, to a degree, reflect biological legacy: (1) final extinction of weakened genera or (2) extinction of new genera that contain few species or represent failed evolutionary “experiments.” Nonetheless, there is evidence that protracted intervals of stressful environmental conditions contributed to high rates of extinction preceding or following certain peak intervals, including the Scythian, Cenomanian, Early Paleocene, and Early Oligocene. The reef-building rudists, for example, suffered heavy extinction during both Cenomanian and Turonian time and then failed to recover quickly. The late Neogene record of bivalve molluscs in the Western Atlantic offers a more detailed picture of delayed recovery. Here early intervals of glacial expansion caused heavy extinction, leaving an impoverished, eurythermal fauna that was virtually unaffected by late Pleistocene glacial episodes. The episode of heavy extinction in Late Eocene time exhibits a similar phenomenon on a worldwide scale. Among the planktonic foraminifera, warm-adapted stenothermal species died out, and eurythermal forms predominated throughout Oligocene time; restoration of vulnerable, stenothermal species proceeded gradually during the Miocene Epoch. This example of delayed recovery and others like it following earlier global crises may have prevented such crises from following one another in rapid succession, yielding an appearance of periodicity.

The Cretaceous-Paleogene (K-Pg) boundary is marked by a major mass extinction, yet this event is thought to have had little effect on the diversity of lizards and snakes (Squamata). A revision of fossil squamates from the Maastrichtian and Paleocene of North America shows that lizards and snakes suffered a devastating mass extinction coinciding with the Chicxulub asteroid impact. Species-level extinction was 83%, and the K-Pg event resulted in the elimination of many lizard groups and a dramatic decrease in morphological disparity. Survival was associated with small body size and perhaps large geographic range. The recovery was prolonged; diversity did not approach Cretaceous levels until 10 My after the extinction, and resulted in a dramatic change in faunal composition. The squamate fossil record shows that the end-Cretaceous mass extinction was far more severe than previously believed, and underscores the role played by mass extinctions in driving diversification.

Paleo-ecological niche modeling (paleoENM) estimates the niches and distributions of extinct species using fossil paleo-coordinates and local environmental data. While general circulation models (GCMs) have been used to estimate climate conditions in deep time, primarily for terrestrial vertebrates, variations in paleo-elevation models used in GCM construction can influence paleoENM outcomes. This study (1) examines the impact of the Cretaceous–Paleogene (K-Pg) mass extinction on the niche dimensions of the marine invertebrate group Turritellinae (Cerithoidea: Turritellidae) and (2) compares two paleo-elevation models’ effects on GCM-based species’ distribution predictions. Fossil occurrence data from the Maastrichtian and Danian periods were collected from the Paleobiology Database (PBDB), museum collections, and published literature. Environmental data were extracted from HadCM3L GCM simulations using Scotese- and Getech-based paleogeographic and pCO2 boundary conditions. We estimated the niche dimensions of turritellines using maximum entropy (MaxEnt) and performed ordination analysis using kernel density estimation. MaxEnt model metrics showed that the Getech-based GCM outperformed the Scotese-based GCM. Geographic projections revealed minor differences in suitable habitat between the Maastrichtian and Danian in the Getech-based GCM, but overinflated predictions in the Scotese-based GCM. Niche overlap between the Maastrichtian and Danian was high, with both GCMs supporting niche similarity and equivalency. Our results suggest that differences in elevation model boundary conditions affected predicted distribution and niche patterns. This study offers a novel approach to understanding ecological persistence in invertebrates after mass extinction events, examines the robustness of GCM boundary conditions in paleoENM studies, and provides a framework for future paleoecological research on fossil invertebrates.