Planktonic Foraminifera are important marine calcifiers, and the ongoing change in the oceanic carbon system makes it essential to understand the influence of environmental factors on the biomineralization of their shells. The amount of calcite deposited by planktonic Foraminifera during calcification has been hypothesized to reflect a range of environmental factors. However, it has never been assessed whether their calcification only passively responds to the conditions of the ambient seawater or whether it reflects changes in resource allocation due to physiological stress. To disentangle these two end-member scenarios, an experiment is required where the two processes are separated. A natural analogue to such an experiment occurred during the deposition of the Mediterranean sapropels, where large changes in surface water composition and stratification at the onset of the sapropel deposition were decoupled from local extinctions of planktonic Foraminifera species. We took advantage of this natural experiment and investigated the reaction of calcification intensity, expressed as mean area density (MAD), of four species of planktonic Foraminifera to changing conditions during the onset of Sapropel S5 (126–121 ka) in a sediment core from the Levantine Basin. We observed a significant relationship between MAD and surface water properties, as reflected by stable isotopes in the calcite of Foraminifera shells, but we failed to observe any reaction of calcification intensity on ecological stress during times of decreasing abundance culminating in local extinction. The reaction of calcification intensity to surface water perturbation at the onset of the sapropel was observed only in surface-dwelling species, but all species calcified more strongly prior to the sapropel deposition and less strongly within the sapropel than at similar conditions during the present-day. These results indicate that the high-salinity environment of the glacial Mediterranean Sea prior to sapropel deposition induced a more intense calcification, whereas the freshwater injection to the surface waters associated with sapropel deposition inhibited calcification. The results are robust to changes in carbonate preservation and collectively imply that changes in normalized shell weight in planktonic Foraminifera should reflect mainly abiotic forcing

[1] The Pacific Ocean is the largest water body on Earth, and circulation in the Pacific contributed significantly to climate evolution in the latest Cretaceous, the culmination of a period of long-term cooling. Here, we present new high-resolution late Campanian to Maastrichtian benthic and planktic foraminiferal stable isotope data and a neodymium (Nd) isotope record obtained from sedimentary ferromanganese oxide coatings of Ocean Drilling Program Hole 1210B from the tropical Pacific Ocean (Shatsky Rise). These new records resolve 13 million years in the latest Cretaceous, providing insights into changes in surface and bottom water temperatures and source regions of deep to intermediate waters covering the carbon isotope excursions of the Campanian-Maastrichtian Boundary Event (CMBE) and the Mid-Maastrichtian event (MME). Our new benthic foraminiferal δ 18O and Nd isotope records together with published Nd isotope data show markedly parallel trends across the studied interval over a broad range of bathyal to abyssal water depths interpreted to reflect changes in the intensity of deep-ocean circulation in the tropical Pacific. In particular, we observe a three-million-year-long period of cooler conditions in the early Maastrichtian (72.5 to 69.5 Ma) when a concomitant change toward less radiogenic seawater Nd isotope signatures probably marks a period of enhanced admixture and northward flow of deep waters with Southern Ocean provenance. We suggest this change to have been triggered by intensified formation and convection of deep waters in the high southern latitudes, a process that weakened during the MME (69.5 to 68.5 Ma). The early Maastrichtian cold interval is closely related to the negative and positive carbon isotope trends of the CMBE and MME. The millions-of-years long duration of these carbon cycle perturbations suggests a tectonic forcing of climatic cooling, possibly related to changes in ocean basin geometry and bathymetry.

Extinction is a remarkably difficult phenomenon to study under natural conditions. This is because the outcome of stress exposure and associated fitness reduction is not known until the extinction occurs and it remains unclear whether there is any phenotypic reaction of the exposed population that can be used to predict its fate. Here we take advantage of the fossil record, where the ecological outcome of stress exposure is known. Specifically, we analyze shell morphology of planktonic Foraminifera in sediment samples from the Mediterranean, during an interval preceding local extinctions. In two species representing different plankton habitats, we observe shifts in trait state and decrease in variance in association with non-terminal stress, indicating stabilizing selection. At terminal stress levels, immediately before extinction, we observe increased growth asymmetry and trait variance, indicating disruptive selection and bet-hedging. The pre-extinction populations of both species show a combination of trait states and trait variance distinct from all populations exposed to non-terminal levels of stress. This finding indicates that the phenotypic history of a population may allow the detection of threshold levels of stress, likely to lead to extinction. It is thus an alternative to population dynamics in studying and monitoring natural population ecology.

The North Atlantic Current (NAC) transports warm salty water to high northern latitudes, with important repercussions for ocean circulation and global climate. A southward displacement of the NAC and Subarctic Front, which separate subpolar and subtropical water masses, is widely suggested for the Last Glacial Maximum (LGM) and may have acted as a positive feedback in glacial expansion at this time. However, the role of the NAC during the intensification of Northern Hemisphere glaciation (iNHG) at ~3.5 to 2.5 Ma is less clear. Here we present new records from Integrated Ocean Drilling Program Site U1313 (41°N) spanning ~2.8–2.4 Ma to trace the influence of Subarctic Front waters above this mid‐latitude site. We reconstruct surface and permanent pycnocline temperatures and seawater δ18O using paired Mg/Ca‐δ18O measurements on the planktic foraminifers Globigerinoides ruber and Globorotalia crassaformis and determine abundances of the subpolar foraminifer Neogloboquadrina atlantica. We find that the first significant glacial incursions of Subarctic Front surface waters above Site U1313 did not occur until ~2.6 Ma. At no time during our study interval was (sub)surface reorganization in the midlatitude North Atlantic analogous to the LGM. Our findings suggest that LGM‐like processes sensu stricto cannot be invoked to explain interglacial‐glacial cycle amplification during iNHG. They also imply that increased glacial productivity at Site U1313 during iNHG was not only driven by southward deflections of the Subarctic Front. We suggest that nutrient injection from cold‐core eddies and enhanced glacial dust delivery may have played additional roles in increasing export productivity in the midlatitude North Atlantic from 2.7 Ma.

18 O transition reflects a change in intermediate-to deep-water circulation from low-latitude to high-latitude water masses, then this change would result in cooler temperatures, higher oxygen concentration, and possibly lower organic-matter flux at the seafloor, resulting in a major benthic foraminiferal assemblage change. If, however, the d 18 O transition was mainly triggered by ice formation, no considerable compositional difference in benthic foraminiferal assemblages would be expected. Our data show a separation of the studied succession into two parts with distinctly different benthic foraminiferal assemblages. Species dominating the older part (73.0-70.5 Ma) tolerate less bottom water oxygenation and are typical components of low-latitude assemblages. In contrast, the younger part (70.0-68.0 Ma) is characterized by species that indicate well-oxygenated bottom waters and species common in high-latitude assemblages. We interpret the observed change in benthic foraminiferal assemblages toward a well-oxygenated environment to reflect the onset of a shift from low-latitude toward high-latitude dominated intermediate-to deep-water sources. This implies that a change in oceanic circulation was at least a major component of the CMBE.