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Pelagic ecosystem function is integral to global biogeochemical cycling, and plays a major role in modulating atmospheric CO2 concentrations (pCO2). Uncertainty as to the effects of human activities on marine ecosystem function hinders projection of future atmospheric pCO2 To this end, events in the geological past can provide informative case studies in the response of ecosystem function to environmental and ecological changes. Around the Cretaceous-Palaeogene (K-Pg) boundary, two such events occurred: Deccan large igneous province (LIP) eruptions and massive bolide impact at the Yucatan Peninsula. Both perturbed the environment, but only the impact coincided with marine mass extinction. As such, we use these events to directly contrast the response of marine biogeochemical cycling to environmental perturbation with and without changes in global species richness. We measure this biogeochemical response using records of deep-sea carbonate preservation. We find that Late Cretaceous Deccan volcanism prompted transient deep-sea carbonate dissolution of a larger magnitude and timescale than predicted by geochemical models. Even so, the effect of volcanism on carbonate preservation was slight compared with bolide impact. Empirical records and geochemical models support a pronounced increase in carbonate saturation state for more than 500 000 years following the mass extinction of pelagic carbonate producers at the K-Pg boundary. These examples highlight the importance of pelagic ecosystems in moderating climate and ocean chemistry., (© 2016 The Author(s).)

Mass extinction at the Cretaceous-Paleogene (K-Pg) boundary coincides with the Chicxulub bolide impact and also falls within the broader time frame of Deccan trap emplacement. Critically, though, empirical evidence as to how either of these factors could have driven observed extinction patterns and carbon cycle perturbations is still lacking. Here, using boron isotopes in foraminifera, we document a geologically rapid surface-ocean pH drop following the Chicxulub impact, supporting impact-induced ocean acidification as a mechanism for ecological collapse in the marine realm. Subsequently, surface water pH rebounded sharply with the extinction of marine calcifiers and the associated imbalance in the global carbon cycle. Our reconstructed water-column pH gradients, combined with Earth system modeling, indicate that a partial ∼50% reduction in global marine primary productivity is sufficient to explain observed marine carbon isotope patterns at the K-Pg, due to the underlying action of the solubility pump. While primary productivity recovered within a few tens of thousands of years, inefficiency in carbon export to the deep sea lasted much longer. This phased recovery scenario reconciles competing hypotheses previously put forward to explain the K-Pg carbon isotope records, and explains both spatially variable patterns of change in marine productivity across the event and a lack of extinction at the deep sea floor. In sum, we provide insights into the drivers of the last mass extinction, the recovery of marine carbon cycling in a postextinction world, and the way in which marine life imprints its isotopic signal onto the geological record.

Plain Language Summary: The Early Eocene Climatic Optimum (EECO) is an interval of prolonged warmth that occurred ∼53 to 49 million years ago. Planktic foraminifera are important (alongside coccolithophores) for understanding the carbon cycle and determining export production in the ocean. To understand how foraminifera can be impacted by extreme heat, we analyzed samples from the Pacific Ocean through measuring changes in (a) the relative abundance, that is, the count of different taxa in a sample, and (b) body (shell) size of planktic foraminifera. At the start of the EECO, the abundance of the genera Morozovella and Chiloguembelina decreased. Despite this decline, the number of foraminifera being buried and the size of the largest shells in a sample does not change. We attribute this to the increased abundance of the genus Acarinina. In general, the accumulation of foraminifera remains stable while the relative abundance of foraminifera to coccolithophores decreases. Together, this implies that coccolithophores are increasing in abundance, and/or mass. During the EECO, one group of foraminifera was able to counterbalance the decrease in abundance of other genera. This highlights the resilience of open‐ocean carbonate production and the base of the marine food web. The Early Eocene Climatic Optimum (EECO; ∼53 and 49 Ma) records the warmest long‐term global average temperature and highest CO2 levels of the Cenozoic. Multiple transient global warming events occur within the EECO, offering an opportunity to investigate the impact of extreme heat on planktic foraminifera. Pacific Ocean Drilling Program (ODP) Sites 1209–1210 (Shatsky Rise) provide an excellent age model and stable isotope records to link biotic data with the carbon cycle across the interval here analyzed (55.6 and 49.93 Ma). We combine carbonate production proxies with changes in planktic foraminiferal assemblages and test‐size. Our data show that during the EECO planktic foraminiferal assemblages were permanently modified, besides transient changes. At the EECO onset, abundance of the genera Morozovella and Chiloguembelina decreased at 53.28 and 52.85 Ma, respectively, confirming published Atlantic Ocean data and thus the global decline of these genera. Given the dominance and large size of Morozovella in early Eocene tropical assemblages, we postulated that this change would have reduced foraminiferal production and the assemblage test‐size. In contrast, we record a slight increase in test‐size within assemblages, controlled by the now dominant genus, Acarinina. The decrease in coarse fraction weight, partially controlled by dissolution, during times of stable foraminferal mass accumulation rate indicates enhanced calcareous nannofossil productivity reducing the foraminiferal contribution to the sediment. During the EECO, despite the decrease in abundance of some genera, species replacement within communities highlights the resilience of pelagic carbonate production, emphasizing the critical role of planktic foraminifera in regulating the marine food web and global carbon cycling. Key Points: At the EECO onset, the abundance of the planktic foraminiferal genera Morozovella and Chiloguembelina decreased and never recoveredSpecies replacement and test‐size increase within assemblages sustained production of pelagic carbonate by foraminiferaGreater adaptability and flexibility of the genus Acarinina led to its dominance over the more specialized morozovellids [ABSTRACT FROM AUTHOR]

Carbonate‐forming organisms play an integral role in the marine inorganic carbon cycle, yet the links between carbonate production and the environment are insufficiently understood. Carbonate production is driven by the abundance of calcifiers and the amount of calcite produced by each individual (their size and weight). Here we investigate how foraminiferal carbonate production changes in the Atlantic, Pacific and Southern Ocean in response to a 4–5°C warming and a 0.3 surface ocean pH reduction during the Palaeocene‐Eocene Thermal Maximum (PETM). To put these local data into a global context, we apply a trait‐based plankton model (ForamEcoGEnIE) to the geologic record for the first time. Our data illustrates negligible change in the assemblage test size and abundance of foraminifers. ForamEcoGEnIE resolves small reductions in size and biomass, but these are short‐lived. The response of foraminifers shows spatial variability linked to a warming‐induced poleward migration and suggested differences in nutrient availability between open‐ocean and shelf locations. Despite low calcite saturation at high latitudes, we reconstruct stable foraminiferal size‐normalized weight. Based on these findings, we postulate that sea surface warming had a greater impact on foraminiferal carbonate production during the PETM than ocean acidification. Changes in the composition of bulk carbonate suggest a higher sensitivity of coccolithophores to environmental change during the PETM than foraminifers. Key Points: No substantial change in planktic foraminiferal carbonate production (size‐normalized weight, test size, or abundance) during the Palaeocene‐Eocene Thermal MaximumSea surface warming and the resulting migration of taxa increased carbonate production in the Southern OceanPelagic carbonate composition changes due to differing impacts on foraminifers and coccolithophores which we link to nutrient availability [ABSTRACT FROM AUTHOR]

Planktic foraminifera are major marine calcifiers in the modern ocean, regulating the marine inorganic carbon pump, and generating marine fossil archives of past climate change. The foraminifera contain ecogroups with and without spines and algal symbionts, creating functional trait diversity which expands their ecological niches. Here, we incorporate symbiosis and spine traits into the symbiont-barren non-spinose foraminifer functional type in EcoGEnIE to represent all the extant foraminifera species. We calibrated the modelled new traits using Latin hypercube sampling (LHS) and identified the optimal model parameters from an ensemble of 1200 runs compared with global observations from core-top sediment samples, sediment traps, and plankton nets. The model successfully describes the global distribution and seasonal abundance variation of the four major foraminiferal functional groups. The model reproduces the dominance of the symbiont-obligate group in subtropical gyres and of the symbiont-barren types in the productive subpolar oceans. Global annual mean biomass and foraminifer-derived carbon export rate are correctly predicted compared to data, with biomass ranging from 0.001 to 0.010 mmol C m -3 and organic carbon export 0.002–0.031 mmol C m -2 d -1. The model captures the seasonal peak time of biomass and organic carbon export but struggles to reproduce the amplitude of both in productive areas. The sparseness and uneven distribution of observations and the model's limitation in upwelling regions likely contribute to this discrepancy. Our model overcomes the lack of major groups in the previous ForamEcoGEnIE 1.0 version and offers the potential to explore foraminiferal ecology dynamics and its impact on biogeochemistry in modern, future, and paleogeographic environments. [ABSTRACT FROM AUTHOR]

Planktic foraminifera are major marine calcifiers in the modern ocean regulating the marine inorganic carbon pump and generating marine fossil archives of past climate change. Some planktic foraminifera evolved spine and symbiosis, increasing functional trait diversity and expanded their ecological niches. Here we incorporate symbiosis and spine traits into the ForamEcoGENIE model, a trait-based model focusing on functional trait rather than individual species, to enable us to study the importance of foraminifera biodiversity in the palaeoceanographic environment. We calibrated the modelled new traits using Latin Hypercube Sampling. We identified the best model run from an ensemble of 1200 runs compared with observations from global core-top, sediment trap, and plankton nets. The model successfully captures the global distribution and seasonal variation of the 4 major functional groups including dominance of the symbiont-obligate type in subtropical gyres and the symbiont-barren type in the productive subpolar oceans. The carbon export rate is correctly predicted for spinose foraminifera, but the model overestimates the global mean biomass of each group by 8 times and global export rate of nonspinose foraminifera by 4 times. Both the observational bias and the model's limitation in linking biomass to export production likely contributes to the discrepancy. Our model approximates a 3.05 g m-2 yr-1 global mean foraminifer-derived calcite flux and 1.1 Gt yr-1 total calcite export, account for 19% of the global pelagic marine calcite budget within the lower range of modern calcite estimates. The calcite export is mostly derived from the symbiont-barren non-spinose group (39%) and the symbiont-obligate spinose group (13%). Our model overcomes the lack of biodiversity in previous version and offers the potential to explore foraminifera ecology dynamics and its impact on biogeochemistry in modern, future and paleogeographic environments. [ABSTRACT FROM AUTHOR]

Planktonic foraminifera are one of the primary calcifiers in the modern ocean, contributing 23%–56% of total global pelagic carbonate production. However, a mechanistic understanding of how physiology and environmental conditions control their abundance and distribution is lacking, hindering the projection of the impact of future climate change. This understanding is important, not only for ecosystem dynamics, but also for marine carbon cycling because of foraminifera's key role in carbonate production. Here we present and apply a global trait‐based ecosystem model of non‐spinose planktonic foraminifera ('ForamEcoGEnIE') to assess their ecology and global distribution under future climate change. ForamEcoGEnIE considers the traits of calcium carbonate production, shell size, and foraging. It captures the main characteristic of biogeographical patterns of non‐spinose species – with maximum biomass concentrations found in mid‐ to high‐latitude waters and upwelling areas. The model also reproduces the magnitude of global carbonate production relatively well, although the foraminifera standing stock is systematically overestimated. In response to future scenarios of rising atmospheric CO2 (RCP6 and RCP8.5), on a regional scale, the modelled foraminifera biomass and export flux increases in the subpolar regions of the North Atlantic and the Southern Ocean while it decreases everywhere else. In the absence of adaptation, the biomass decline in the low‐latitude South Pacific suggests extirpation. The model projects a global average loss in non‐spinose foraminifera biomass between 8% (RCP6) and 11% (RCP8.5) by 2050 and between 14% and 18% by 2100 as a response to ocean warming and associated changes in primary production and ecological dynamics. Global calcium carbonate flux associated with non‐spinose foraminifera declines by 13%–18% by 2100. That decline can slow down the ocean carbonate pump and create short‐term positive feedback on rising atmospheric pCO2. [ABSTRACT FROM AUTHOR]

Silicate weathering is a primary control on the carbon cycle and therefore long-term climate. Tracing silicate weathering in the geological record has been a challenge for decades, with a number of proxies proposed and their limits determined. Recently lithium isotopes in marine carbonates have emerged as a potential tracer. Bulk carbonates are increasingly being used as a Li isotope archive, though with limited tests thus far of the robustness of this approach in the modern ocean. As the bulk composition of marine pelagic carbonates has changed through time and geographically, assessing the fidelity of bulk carbonate as proxy carrier is fundamental. To address the impact of compositional variability in bulk carbonate on Li isotopes, we examine 27 Bahamian aragonitic bulk carbonates and 16 Atlantic largely calcitic core-top sediment samples. Two core-tops only have trace (<10 %) carbonate, and are analysed to test whether carbonates in such sections are still a viable archive. We selectively extract the exchangeable and carbonate fractions from the core-top samples. The exchangeable fraction contains ∼2 % of the total Li and has a fairly constant offset from seawater of 16.5 ± 0.8‰. When leaching silicate-containing carbonates, acetic acid buffered with sodium acetate appears a more robust method of solely attacking carbonates compared to dilute HCl, which may also liberate some silicate-bound Li. Carbonates from samples that do not contain aragonite have the isotopic fractionation of seawater of Δ7Li seawater-calcite = 6.1 ± 1.3‰ (2sd), which is not affected by latitude or the water depth the sample was deposited at. The pure aragonite bulk carbonates from the Bahamas have a fractionation of Δ7Li seawater-aragonite = 9.6 ± 0.6‰. A sediment sample from the Galician coast that mostly consists of quartz is highly offset from seawater by ∼20‰ and also has relatively high Li/Ca ratios. These high values are not due to leaching of silicate material directly (Al/Ca ratios are low). We interpret this addition via cation exchange of Li from silicate during recrystallisation. Overall bulk carbonates from the open ocean are a reliable archive of seawater δ 7Li, but care must be taken with carbonate mineralogy and low-carbonate samples. Overall, therefore, any examination of the palaeo-seawater δ 7Li record must be reproduced in different global settings (e.g. multiple global cores) before it can be considered robust. [ABSTRACT FROM AUTHOR]

The Paleogene was punctuated by perturbations of the global carbon cycle, many associated with transient global warming events (hyperthermals). The Dan-C2 event (~160 kyr after Cretaceous/Paleogene boundary; K/Pg) was the oldest of these eccentricity-linked carbon cycle disturbances (ELCD). In contrast to other hyperthermals, the Dan-C2 event was not characterised by bottom water warming, and surface water warming probably was not global. Benthic foraminiferal assemblages across Dan-C2 at SE Atlantic Ocean Drilling Program (ODP) Site 1262 are diverse and strongly dominated by calcareous species. Epifaunal and infaunal morphogroups are equally abundant, suggesting meso-oligotrophic seafloor conditions. Assemblages decreased in diversity gradually before Dan-C2, and Nuttallides truempyi decreased in relative abundance while Stensioeina beccariiformis and the agglutinant Spiroplectammina spectabilis increased, suggesting enhanced food supply to the seafloor. Benthic foraminifera were not highly affected by the Dan-C2 event. An increase in relative abundance of the opportunistic species Bulimina kugleri and Seabrookia cretacea after Dan-C2 points to a change in the type of organic matter arriving at the seafloor. These changes may have been caused by ongoing environmental and/or evolutionary instability following K/Pg mass extinction of oceanic plankton. Variability in composition of pelagic ecosystems, thus the type and/or amount of food arriving at the seafloor, may have been caused by the gradual recovery of pelagic ecosystems after that extinction, possibly affected by warming and pH changes due to Deccan volcanism. • Benthic foraminifera did not respond to the Dan-C2 event. • Recovery of primary producers after K/Pg cause unstable food-supply to the seafloor. • Deccan volcanism probably influenced in the recovery of primary producers. • Dan-C2 is an eccentricity-linked carbon cycle disturbance, not a hyperthermal. [ABSTRACT FROM AUTHOR]