Description: The composition of planktonic foraminiferal (PF) calcite is routinely used to reconstruct climate variability. However, PF ecology leaves a large imprint on the proxy signal: seasonal and vertical habitats of PF species vary spatially, causing variable offsets from annual mean surface conditions recorded by sedimentary assemblages. PF seasonality changes with temperature in a way that minimises the environmental change that individual species experience and it is not unlikely that changes in depth habitat also result from such habitat tracking. While this behaviour could lead to an underestimation of spatial or temporal trends as well as of variability in proxy records, most palaeoceanographic studies are (implicitly) based on the assumption of a constant habitat. Up to now, the effect of habitat tracking on foraminifera proxy records has not yet been formally quantified on a global scale. Here we attempt to characterise this effect on the amplitude of environmental change recorded in sedimentary PF using core top δ18O data from six species. We find that the offset from mean annual near-surface δ18O values varies with temperature, with PF δ18O indicating warmer than mean conditions in colder waters (on average by −0.1 ‰ (equivalent to 0.4 °C) per °C), thus providing a first-order quantification of the degree of underestimation due to habitat tracking. We use an empirical model to estimate the contribution of seasonality to the observed difference between PF and annual mean δ18O and use the residual Δδ18O to assess trends in calcification depth. Our analysis indicates that given an observation-based model parametrisation calcification depth increases with temperature in all species and sensitivity analysis suggests that a temperature-related seasonal habitat adjustment is essential to explain the observed isotope signal. Habitat tracking can thus lead to a significant reduction in the amplitude of recorded environmental change. However, we show that this behaviour is predictable. This allows accounting for habitat tracking, enabling more meaningful reconstructions and improved data–model comparison.

Description: Cruise M140 combined sampling of plankton, mineral dust and other particles in the water column with recovery of data and samples from long-term observational platforms (sediment traps and dust-collecting buoys). The aim of the cruise was to provide new observations to improve our understanding of the ecology of planktonic foraminifera as important carriers of paleoceanographic proxies and to investigate how mineral dust deposition and the production of marine snow and biogenic particle ballast vary in space and time and how they affect the marine biological pump. To this end, the cruise followed a transect in the central western Atlantic between oligotrophic waters of the subtropical gyre and the productive coastal waters off Mauretania affected by coastal upwelling. To characterise population dynamics, ecology and physiology of planktonic foraminifera, we obtained a series of fourteen vertically resolved plankton net profiles along the cruise track, together with profiles of physical and chemical properties of the ambient water masses. Live foraminifera extracted from these profiles were used to quantify photosynthetic activity of selected species and determine their photoadaptation. High-resolution spatial and temporal sampling of the upper 300 m over 24 hours was carried out at two locations (recovering 41 and 46 vertical profiles), allowing the characterisation of patchiness and daily vertical migration of planktonic foraminifera. Moorings with sediment traps monitoring the seasonal and short-term variability of particle fluxes and buoys monitoring atmospheric dust deposition in the region were successfully recovered in the central Atlantic (M3), south of Cabo Verde (M1) and off Mauretania (CB and CBi) and redeployed in the latter two regions to continue the monitoring. Short-term variability of sizes and types of sinking particles in the water column were characterised in each of the monitoring regions with drifting sediment traps and in the Cape Blanc region off Mauretania also with continuous vertical particle camera profile. All aims of the cruise have been met – the plankton sampling and particle characterization studies were carried out successfully and all moorings and buoys could be recovered and/or redeployed as planned.

Description: Diurnal vertical migration (DVM) is a widespread phenomenon in the upper ocean, but it remains unclear to what degree it also involves passively transported micro- and meso-zooplankton. These organisms are difficult to monitor by in situ sensing and observations from discrete samples are often inconclusive. Prime examples of such ambiguity are planktonic foraminifera, where contradictory evidence for DVM continues to cast doubt on the stability of species vertical habitats, which introduces uncertainties in geochemical proxy interpretation. To provide a robust answer, we carried out highly replicated randomised sampling with 41 vertically resolved plankton net hauls taken within 26 hours in a confined area of 400 km2 in the tropical North Atlantic, where DVM in larger plankton occurs. Manual enumeration of planktonic foraminifera cell density consistently reveals the highest total cell concentrations in the surface mixed layer (top 50 m) and analysis of cell density in seven individual species representing different shell sizes, life strategies and presumed depth habitats reveals consistent vertical habitats not changing over the 26 hours sampling period. These observations robustly reject the existence of DVM in planktonic foraminifera in a setting where DVM occurs in other organisms.

Description: The climate of the last two millennia was characterised by decadal to multi‐centennial variations which were recorded in terrestrial records and had important societal impacts. The cause of these climatic events is still under debate but changes in the North Atlantic circulation have often been proposed to play an important role. In this review we compile available high‐resolution paleoceanographic datasets from the northern North Atlantic and Nordic Seas. The records are grouped into regions related to modern ocean conditions and their variability is discussed. We additionally discuss our current knowledge from modelling studies, with a specific focus on the dynamical changes that are not well inferred from the proxy records. An illustration is provided through the analysis of two climate model ensembles and an individual simulation of the last millennium. This review thereby provides an up‐to‐date paleo‐perspective on the North Atlantic multidecadal to multi‐centennial ocean variability across the last two millennia.

Description: The species composition of many groups of marine plankton appears well predicted by sea surface temperature (SST). Consequently, fossil plankton assemblages have been widely used to reconstruct past SST. Most applications of this approach make use of the highest possible taxonomic resolution. However, not all species are sensitive to temperature, and their distribution may be governed by other parameters. There are thus reasons to question the merit of including information about all species, both for transfer function performance and for its effect on reconstructions. Here we investigate the effect of species selection on planktonic foraminifera transfer functions. We assess species importance for transfer function models using a random forest technique and evaluate the performance of models with an increasing number of species. Irrespective of using models that use the entire training set (weighted averaging) or models that use only a subset of the training set (modern analogue technique), we find that the majority of foraminifera species does not carry useful information for temperature reconstruction. Less than one-third of the species in the training set is required to provide a temperature estimate with a prediction error comparable to a transfer function that uses all species in the training set. However, species selection matters for paleotemperature estimates. We find that transfer function models with a different number of species but with the same error may yield different reconstructions of sea surface temperature when applied to the same fossil assemblages. This ambiguity in the reconstructions implies that fossil assemblage change reflects a combination of temperature and other environmental factors. The contribution of the additional factors is site and time specific, indicating ecological and geological complexity in the formation of the sedimentary assemblages. The possibility of obtaining multiple different reconstructions from a single sediment record presents a previously unrecognized source of uncertainty for sea surface temperature estimates based on planktonic foraminifera assemblages. This uncertainty can be evaluated by determining the sensitivity of the reconstructions to species pruning.

Description: The ocean—the Earth’s largest ecosystem—is increasingly affected by anthropogenic climate change1,2. Large and globally consistent shifts have been detected in species phenology, range extension and community composition in marine ecosystems3,4,5. However, despite evidence for ongoing change, it remains unknown whether marine ecosystems have entered an Anthropocene6 state beyond the natural decadal to centennial variability. This is because most observational time series lack a long-term baseline, and the few time series that extend back into the pre-industrial era have limited spatial coverage7,8. Here we use the unique potential of the sedimentary record of planktonic foraminifera—ubiquitous marine zooplankton—to provide a global pre-industrial baseline for the composition of modern species communities. We use a global compilation of 3,774 seafloor-derived planktonic foraminifera communities of pre-industrial age9 and compare these with communities from sediment-trap time series that have sampled plankton flux since ad 1978 (33 sites, 87 observation years). We find that the Anthropocene assemblages differ from their pre-industrial counterparts in proportion to the historical change in temperature. We observe community changes towards warmer or cooler compositions that are consistent with historical changes in temperature in 85% of the cases. These observations not only confirm the existing evidence for changes in marine zooplankton communities in historical times, but also demonstrate that Anthropocene communities of a globally distributed zooplankton group systematically differ from their unperturbed pre-industrial state.

Description: Species of planktonic foraminifera exhibit specific seasonal production patterns and different preferred vertical habitats. The seasonality and vertical habitats are not constant throughout the range of the species and changes therein must be considered when interpreting paleoceanographic reconstructions based on fossil foraminifera. Accounting for the effect of vertical and seasonal habitat tracking on foraminifera proxies at times of climate change is difficult because it requires independent fossil evidence. An alternative that could reduce the bias in paleoceanographic reconstructions is to predict species-specific habitat shifts under climate change using an ecosystem modeling approach. To this end, we present a new version of a planktonic foraminifera model, PLAFOM2.0, embedded into the ocean component of the Community Earth System Model, version 1.2.2. This model predicts monthly global concentrations of the planktonic foraminiferal species: Neogloboquadrina pachyderma, N. incompta, Globigerina bulloides, Globigerinoides ruber (white), and Trilobatus sacculifer throughout the world ocean, resolved in 24 vertical layers to 250m depth. The resolution along the vertical dimension has been implemented by applying the previously used spatial parameterization of biomass as a function of temperature, light, nutrition, and competition on depth-resolved parameter fields. This approach alone results in the emergence of species-specific vertical habitats, which are spatially and temporally variable. Although an explicit parameterization of the vertical dimension has not been carried out, the seasonal and vertical distribution patterns predicted by the model are in good agreement with sediment trap data and plankton tow observations. In the simulation, the colder-water species N. pachyderma, N. incompta, and G. bulloides show a pronounced seasonal cycle in their depth habitat in the polar and subpolar regions, which appears to be controlled by food availability. During the warm season, these species preferably occur in the subsurface, while towards the cold season they ascend through the water column and are found closer to the sea surface. The warm-water species G. ruber (white) and T. sacculifer exhibit a less variable shallow depth habitat with highest biomass concentrations within the top 40m of the water column. Nevertheless, even these species show vertical habitat variability and their seasonal occurrence outside the tropics is limited to the warm surface layer that develops at the end of the warm season. The emergence in PLAFOM2.0 of species-specific vertical habitats that are consistent with observations indicates that the population dynamics of planktonic foraminifera species may be driven by the same factors in time, space, and with depth, in which case the model can provide a reliable and robust tool to aid the interpretation of proxy records.

Description: Shell fluxes of planktonic Foraminifera species vary intra-annually in a pattern that appears to follow the seasonal cycle. However, the variation in the timing and prominence of seasonal flux maxima in space and among species remains poorly constrained. Thus, although changing seasonality may result in a flux-weighted temperature offset of more than 5° C within a species, this effect is often ignored in the interpretation of Foraminifera-based paleoceanographic records. To address this issue we present an analysis of the intra-annual pattern of shell flux variability in 37 globally distributed time series. The existence of a seasonal component in flux variability was objectively characterised using periodic regression. This analysis yielded estimates of the number, timing and prominence of seasonal flux maxima. Over 80% of the flux series across all species showed a statistically significant periodic component, indicating that a considerable part of the intra-annual flux variability is predictable. Temperature appears to be a powerful predictor of flux seasonality, but its effect differs among species. Three different modes of seasonality are distinguishable. Tropical and subtropical species (Globigerinoides ruber (white and pink varieties), Neogloboquadrina dutertrei, Globigerinoides sacculifer, Orbulina universa, Globigerinella siphonifera, Pulleniatina obliquiloculata, Globorotalia menardii, Globoturborotalita rubescens, Globoturborotalita tenella and Globigerinoides conglobatus) appear to have a less predictable flux pattern, with random peak timing in warm waters. In colder waters, seasonality is more prevalent: peak fluxes occur shortly after summer temperature maxima and peak prominence increases. This tendency is stronger in species with a narrower temperature range, implying that warm-adapted species find it increasingly difficult to reproduce outside their optimum temperature range and that, with decreasing mean temperature, their flux is progressively more focussed in the warm season. The second group includes the temperate to cold-water species Globigerina bulloides, Globigerinita glutinata, Turborotalita quinqueloba, Neogloboquadrina incompta, Neogloboquadrina pachyderma, Globorotalia scitula, Globigerinella calida, Globigerina falconensis, Globorotalia theyeri and Globigerinita uvula. These species show a highly predictable seasonal pattern, with one to two peaks a year, which occur earlier in warmer waters. Peak prominence in this group is independent of temperature. The earlier-when-warmer pattern in this group is related to the timing of productivity maxima. Finally, the deep-dwelling Globorotalia truncatulinoides and Globorotalia inflata show a regular and pronounced peak in winter and spring. The remarkably low flux outside the main pulse may indicate a long reproductive cycle of these species. Overall, our analysis indicates that the seasonality of planktonic Foraminifera shell flux is predictable and reveals the existence of distinct modes of phenology among species. We evaluate the effect of changing seasonality on paleoceanographic reconstructions and find that, irrespective of the seasonality mode, the actual magnitude of environmental change will be underestimated. The observed constraints on flux seasonality can serve as the basis for predictive modelling of flux pattern. As long as the diversity of species seasonality is accounted for in such models, the results can be used to improve reconstructions of the magnitude of environmental change in paleoceanographic records.

Description: Previous work showed that South Atlantic sediments have lower glacial than Holocene 231Pa/230Th, which was attributed to a switch in the flow direction of Atlantic deep-water. Debate exists, however as to the degree to which two processes - circulation and scavenging - determine sedimentary 231Pa/230Th, making this interpretation contentious. Here we address this issue using 145-kyr records of paleocirculation proxies. Benthic foraminiferal d13C, neodymium isotopes (ENd) and sedimentary 231Pa/230Th were all measured in a single sediment core from the South Atlantic subtropical gyre. This site largely excludes the influence of local productivity changes on 231Pa/230Th records. Measured 231Pa/230Th ranges between ~0.041 during glacials to ~0.055 during interglacial periods and are consistently lower than the production ratio, indicating export of 231Pa from the central South Atlantic for the entire duration of the record. The lower glacial 231Pa/230Th is regionally consistent, suggesting that basin-scale oceanographic processes cause the decrease. In turn, less radiogenic ENd and lower benthic d13C confirm the classical picture of an increase in Southern Component Water (SCW) influence in the Atlantic during glacial periods and point to a circulation control on the observed 231Pa/230Th decrease rather than a local productivity change. We suggest that associated with this change in water mass distribution the dominant sink for 231Pa shifted from the margins of the South Atlantic and/or the Southern Ocean during interglacials, to the North Atlantic during glacial periods. Indeed, elevated 231Pa/230Th in the deep North Atlantic during glacials supports this mechanism of northward transport of 231Pa by SCW.