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Past responses to environmental change provide vital baseline data for estimating the potential resilience of extant taxa to future change. Here, we investigate the latitudinal range contraction that terrestrial and freshwater turtles (Testudinata) experienced from the Late Cretaceous to the Paleogene (100.5-23.03 mya) in response to major climatic changes. We apply ecological niche modeling (ENM) to reconstruct turtle niches, using ancient and modern distribution data, paleogeographic reconstructions, and the HadCM3L climate model to quantify their range shifts in the Cretaceous and late Eocene. We then use the insights provided by these models to infer their probable ecological responses to future climate scenarios at different representative concentration pathways (RCPs 4.5 and 8.5 for 2100), which project globally increased temperatures and spreading arid biomes at lower to mid-latitudes. We show that turtle ranges are predicted to expand poleward in the Northern Hemisphere, with decreased habitat suitability at lower latitudes, inverting a trend of latitudinal range contraction that has been prevalent since the Eocene. Trionychids and freshwater turtles can more easily track their niches than Testudinidae and other terrestrial groups. However, habitat destruction and fragmentation at higher latitudes will probably reduce the capability of turtles and tortoises to cope with future climate changes.

The ocean biological pump is the mechanism by which carbon and nutrients are transported to depth. As such, the biological pump is critical in the partitioning of carbon dioxide between the ocean and atmosphere, and the rate at which that carbon can be sequestered through burial in marine sediments. How the structure and function of planktic ecosystems in the ocean govern the strength and efficiency of the biological pump and its resilience to disruption are poorly understood. The aftermath of the impact at the Cretaceous/Palaeogene (K/Pg) boundary provides an ideal opportunity to address these questions as both the biological pump and marine plankton size and diversity were fundamentally disrupted. The excellent fossil record of planktic foraminifera as indicators of pelagic-biotic recovery combined with carbon isotope records tracing biological pump behaviour, show that the recovery of ecological traits (diversity, size and photosymbiosis) occurred much later (approx. 4.3 Ma) than biological pump recovery (approx. 1.8 Ma). We interpret this decoupling of diversity and the biological pump as an indication that ecosystem function had sufficiently recovered to drive an effective biological pump, at least regionally in the South Atlantic.

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.

Despite the important role of planktonic foraminifera in regulating the ocean carbonate production and their unrivalled value in reconstructing paleoenvironments, our knowledge on their ecology is limited. A variety of observational techniques such as plankton tows, sediment traps and experiments have contributed to our understanding of foraminifera ecology. But, fundamental questions around costs and benefits of calcification and the effect of nutrients, temperature and ecosystem structure on these organisms remain unanswered. To tackle these questions, we take a novel mechanistic approach to study planktonic foraminifera ecology based on trait theory. We develop a zero-dimensional (0-D) trait-based model to account for the biomass of prolocular (20 m) and adult (160 m) stages of non-spinose foraminifera species and investigate their potential interactions with phytoplankton and other zooplankton under different temperature and nutrient regimes. Building on the costs and benefits of calcification, we model two ecosystem structures to explore the effect of resource competition and temperature on planktonic foraminifera biomass. By constraining the model results with ocean biomass estimations of planktonic foraminifera, we estimate that the energetic cost of calcification could be about 10 50% and 10 40% for prolocular and adult stages respectively. Our result suggest that the shell provides protection against processes other than predation (e.g. pathogen invasion). We propose that the low standing biomass of planktonic foraminifera plays a key role in their survival from predation, along with their shell protection. Our model suggests a shift from temperature as a main regulator of foraminifera biomass in the early developmental stage to resource competition for adult biomass.

Climate change is predicted to alter temperature, carbonate chemistry and oxygen availability in the oceans, which will affect individuals, populations and ecosystems. We use the fossil record of benthic foraminifers to assess developmental impacts in response to environmental changes during the Palaeocene-Eocene Thermal Maximum (PETM). Using an unprecedented number of µ-computed tomography scans, we determine the size of the proloculus (first chamber), the number of chambers and the final size of two benthic foraminiferal species which survived the extinction at sites 690 (Atlantic sector, Southern Ocean, palaeodepth 1900 m), 1210 (central equatorial Pacific, palaeodepth 2100 m) and 1135 (Indian Ocean sector, Southern Ocean, palaeodepth 600-1000 m). The population at the shallowest site, 1135, does not show a clear response to the PETM, whereas those at the other sites record reductions in diameter or proloculus size. Temperature was similar at all sites, thus it is not likely to be the reason for differences between sites. At site 1210, small size coincided with higher chamber numbers during the peak event, and may have been caused by a combination of low carbonate ion concentrations and low food supply. Dwarfing at site 690 occurred at lower chamber numbers, and may have been caused by decreasing carbonate saturation at sufficient food levels to reproduce. Proloculus size varied strongly between sites and through time, suggesting a large influence of environment on both microspheric and megalospheric forms without clear bimodality. The effect of the environmental changes during the PETM was more pronounced at deeper sites, possibly implicating carbonate saturation.This article is part of a discussion meeting issue 'Hyperthermals: rapid and extreme global warming in our geological past'.

Climate change is predicted to alter temperature, carbonate chemistry and oxygen availability in the oceans, which will affect individuals, populations and ecosystems. We use the fossil record of benthic foraminifers to assess developmental impacts in response to environmental changes during the Palaeocene-Eocene Thermal Maximum (PETM). Using an unprecedented number of µ-computed tomography scans, we determine the size of the proloculus (first chamber), the number of chambers and the final size of two benthic foraminiferal species which survived the extinction at sites 690 (Atlantic sector, Southern Ocean, palaeodepth 1900 m), 1210 (central equatorial Pacific, palaeodepth 2100 m) and 1135 (Indian Ocean sector, Southern Ocean, palaeodepth 600-1000 m). The population at the shallowest site, 1135, does not show a clear response to the PETM, whereas those at the other sites record reductions in diameter or proloculus size. Temperature was similar at all sites, thus it is not likely to be the reason for differences between sites. At site 1210, small size coincided with higher chamber numbers during the peak event, and may have been caused by a combination of low carbonate ion concentrations and low food supply. Dwarfing at site 690 occurred at lower chamber numbers, and may have been caused by decreasing carbonate saturation at sufficient food levels to reproduce. Proloculus size varied strongly between sites and through time, suggesting a large influence of environment on both microspheric and megalospheric forms without clear bimodality. The effect of the environmental changes during the PETM was more pronounced at deeper sites, possibly implicating carbonate saturation.This article is part of a discussion meeting issue 'Hyperthermals: rapid and extreme global warming in our geological past'.

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]