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Coccolithophores are among the most important calcifying pelagic organisms. To assess how coccolithophore species with different coccolith-carbonate mass and distinct ecological resilience to ocean warming will influence the “rain ratio” and the “biological carbon pump”, 1 yr of species-specific coccolith-carbonate export fluxes were quantified using sediment traps moored at four sites between NW Africa and the Caribbean (i.e., CB-20°N/21°W, at 1214 m; M1-12°N/23°W, at 1150 m; M2-14°N/37°W, at 1235 m; M4-12°N/49°W, at 1130 m). Highest coccolith-CaCO₃ fluxes at the westernmost site M4, where the nutricline is deepest along the tropical North Atlantic, were dominated by deep-dwelling small-sized coccolith species Florisphaera profunda and Gladiolithus flabellatus. Total coccolith-CaCO₃ fluxes of 371 mg m−2 yr−1 at M4 were followed by 165 mg m−2 yr−1 at the north-easternmost CB, 130 mg m−2 yr−1 at M1, and 114 mg m−2 yr−1 at M2 in between. Coccoliths accounted for nearly half of the total carbonate flux at M4 (45%), much higher compared to 23% at M2 and 15% at M1 and CB. At site M4, highest ratios of coccolith-CaCO₃ to particulate organic carbon fluxes and weak correlations between the carbonate of deep-dwelling species and particulate organic carbon suggest that increasing productivity in the lower photic zone in response to ocean warming might enhance the rain ratio and reduce the coccolithballasting efficiency. The resulting weakened biological carbon pump could, however, be counterbalanced by increasing frequency of Saharan dust outbreaks across the tropical Atlantic, providing mineral ballast as well as nutrients to fuel fast-blooming and ballast-efficient coccolithophore species.

A natural pH gradient caused by marine CO2 seeps off the Methana peninsula (Saronikos Gulf, eastern Peloponnese peninsula) was used as a natural laboratory to assess potential effects of ocean acidification on coccolithophores. Coccolithophore communities were therefore investigated in plankton samples collected during September 2011, September 2016 and March 2017. The recorded cell concentrations were up to ~50 x103 cells/l, with a high Shannon index of up to 2.8, along a pH gradient from 7.61 to 8.18, with values being occasionally <7. Numerous holococcolithophore species represented 60-90% of the surface water assemblages in most samples during September samplings. Emiliania huxleyi was present only in low relative abundances in September samples, but it dominated in March assemblages. Neither malformed nor corroded coccolithophores were documented. Changes in the community structure can possibly be related to increased temperatures, while the overall trend associates low pH values with high cell densities. Our preliminary results indicate that in long-termed acidified, warm and stratified conditions, the study of the total coccolithophore assemblage may prove useful to recognize the intercommunity variability, which favors the increment of lightly calcified species such as holococcolithophores.

We present a new high-resolution reconstruction of annual sea-surface temperatures (SSTa) and net primary productivity (NPP) using novel coccolithophore-based models developed for the Eastern Equatorial Pacific (EEP). We combined published coccolithophore census counts from core-tops in the Eastern Pacific with 32 new samples from the Equatorial region, to derive a new statistical model to reconstruct SSTa. Results show that the addition of the new EEP samples improves existing coccolithophore-based SST-calibrations, and allow reconstructing SSTa in the EEP with higher confidence. We also merged the relative abundance of deep-photic species Florisphaera profunda in the same surface sediment samples with existing calibration datasets for tropical regions, to reconstruct annual NPP. Both temperature and productivity calibrations were successfully applied to fossil coccolith data from Ocean Drilling Project Site 1240, in the EEP. The coccolith-based SSTa estimates show a cooling during the Last Glacial Maximum (LGM) and the Younger Dryas, and warming at the start of the Holocene. This pattern differs in the timing and magnitude of the temperature changes from other available SST-reconstructions based on biogeochemical and faunal proxies. We discuss these discrepancies to be the result of different proxy sensitivities to insolation forcing, seasonal bias, and/or preservation artifacts. Reconstructed annual NPP shows a general decreasing trend from the late last glacial period to recent times, which we relate to the weakening of wind-driven equatorial upwelling towards the Holocene. We also calculated carbon export using our SSTa and NPP reconstructions, and compared to other geochemical-based reconstructions for the same location. Our coupled SSTa-NPP reconstruction provides key data to more fully assess the evolution of primary and export productivity as well as organic carbon burial in the EEP, with implications for its role in global biogeochemical cycles across glacial terminations.

Throughout the Cenozoic, calcareous nannoplankton and planktonic foraminifera have been the main producers of pelagic carbonate preserved on the seafloor. While past variability in pelagic carbonate production has been previously studied, relatively little is known about the variability in the relative contribution of the two components. This is important because the responses of the two groups to environmental forcing could be different such that they could amplify or reduce the magnitude of fluctuations in total carbonate production. Here we present new data from the tropical Atlantic that allow us to quantify changes in the relative contribution of the two groups to the total pelagic carbonate burial flux on orbital scales and between different climate states since the Miocene. We find that the composition of the deposited pelagic carbonate remained similar on long time scales, with foraminifera making up about 30 % of the deposited carbonate, but varied by up to a factor of two on orbital time scales. We show that the relative contribution of planktonic foraminifera and coccoliths did not correlate with the total pelagic carbonate production, either in the Pliocene, when its dominant cyclicity was in the precession band and in phase with orbital parameters modulations, or in the Miocene, when its predominant cyclicity was in the eccentricity band and in antiphase with orbital parameters modulations. The observed variability in tropical pelagic carbonate productivity between foraminifera and coccolithophores suggests that the two main groups of pelagic calcifiers responded fundamentally differently to orbital forcing and associated oceanographic changes in the tropical ocean, but the resulting changes in their proportions did not drive changes in overall pelagic carbonate deposition on either geological or orbital time scales.

Coccolithophores are the main type of calcifying phytoplankton in the Southern Ocean (SO) and are key organisms in the production of particulate inorganic carbon (PIC). However, in situ studies of coccolithophores and in particular of their importance for the input of PIC in the SO are sparse in space and time due to the harsh weather conditions in the subantarctic realm. An alternative tool for monitoring PIC is the use of optical remote sensing, as coccolithophores account for most of the optical PIC backscattering in the sea. The aim of the present study is to provide coccolithophore-based estimates of PIC derived from Scanning Electron Microscope coccolith morphometric analyses and MODIS Aqua Level-2 and Level-3 PIC concentration values along two latitudinal transects from New Zealand to Antarctica and across the Drake Passage. In general, the coccolith-estimated PIC and satellite-derived PIC datasets show comparable trends in the Subantarctic and Polar Frontal zones of both transects, with coccolith-derived PIC values being generally lower than satellite PIC values. According to the coccolithophorid data, Emiliania huxleyi type A, over-calcified type A, and other taxa (e.g., Calcidiscus leptoporus) only contribute to coccolithophore PIC in the northernmost sampling locations, whereas E. huxleyi morphogroup B contributes substantially to the PIC content south of the Subantarctic Front in both transects. High satellite-derived PIC concentrations south of the Polar Front are not apparent in the coccolith-based PIC data. We suggest that the high reflectance signal in the Antarctic Zone may instead relate to the presence of small biogenic opal particles (e.g., diatoms, silicoflagellates, and/or small siliceous plankton) or other unknown highly reflective particles (such as Phaeocystis aggregations). Our results highlight the challenges presented by the lack of reliable satellite data in some parts of the SO as well as the importance of in situ measurements and methodological accuracy when estimating PIC values.

The present study was conducted to provide information about the upper water column distribution of living coccolithophores in the subtropical Gulf of Mexico. In total, 52 plankton samples from 6 stations collected in March 2006 were analysed. Coccolithophore standing crops range from zero in deep-water samples (200 m) to about 23 000 to 46 500 coccospheres per litre at intermediate water depths. From 39 identified taxa, only nine species contribute significant cell numbers of more than 1000 cells 1-1 and comprise more than 5% of the communities in at least one sample. Emiliania huxleyi was the most abundant species throughout the stations with concentrations of up to 22 700 cells l-1. At all stations, a vertical succession of coccolithophore species was found. Umbellosphaera tenuis (type IV), Discosphaera tubifera, Rhabdosphaera clavigera and, as a minor component, species of the genus Syracosphaera, were mainly restricted to surface waters (≤50 m), all adapted to lower nutrient levels. The assemblage in the deep photic zone (50-150 m water depth) was composed mainly of abundant Florisphaera profunda, Gladiolithus flabellatus and Algirosphaera robusta indicating a tolerance of lower light availability. Thus, the vertical distribution of all coccolithophorid taxa, except the placolith-bearing species, in particular E. huxleyi, was probably controlled by upper photic-zone water temperature and stratification of the water column.

Marine plankton is an important component of the global carbon cycle. Whereas the production and seafloor export of organic carbon produced by the plankton, the biological pump, has received much attention, the long-term variability in plankton calcification, controlling the carbonate counter pump, remains less well understood. However, it has been shown that changes in pelagic calcification (biological compensation) could affect the ocean's buffering capacity and thus regulate global carbon budget on geological timescales. Here we use Neogene pelagic sediments deposited on the Ceara Rise in the tropical Atlantic to characterize the variability in pelagic carbonate production with a focus on warm climates. A re-evaluation of published records of carbonate accumulation at the Ceara Rise reveals a systematic increase in sedimentation rates since the late Miocene, but the carbonate accumulation rate does not show a clear trend. Instead, we observe substantial orbital timescale variability in carbonate accumulation, combined with a trend towards less carbonate on average at sites located below 4 km, likely due to the effect of carbonate dissolution. To evaluate long-term changes against possible orbital-scale variability, we generated new high-resolution records of carbonate accumulation rate at Ocean Drilling Program (ODP) Site 927 across two Quaternary interglacials (MIS 5 and MIS 9), the Pliocene warm period (MIS KM5) and the Miocene Climatic Optimum (MCO). We observe that the highest carbonate accumulation rates occurred during the Pliocene but that each of the studied intervals was characterized by large-magnitude orbital variability. Prominent variations in carbonate accumulation prior to the Quaternary preservation cycles appear to follow Earth obliquity and eccentricity. These results imply that pelagic carbonate accumulation in the tropical ocean, buffered from large temperature changes, varied on orbital timescales. The magnitude of the orbital-scale variability was similar or even higher than the long-term mean differences among the studied intervals. Since preservation can be excluded as a driver of these changes prior to the Quaternary, the observed variations must reflect changes in the export flux of pelagic biogenic carbonate. We conclude that the overall carbonate production by pelagic calcifiers responded to local changes in light, temperature, and nutrients delivered by upwelling, which followed long orbital cycles, as well as to long-term shifts in climate and/or ocean chemistry. The inferred changes on both timescales were sufficiently large such that when extrapolated on a global scale, they could have played a role in the regulation of the carbon cycle and global climate evolution during the transition from the Miocene warm climates into the Quaternary icehouse.

The Southern Ocean is experiencing rapid and profound changes in its physical and biogeochemical properties that may influence the distribution and composition of pelagic plankton communities. Coccolithophores are the most prolific carbonate-producing phytoplankton group, playing an important role in Southern Ocean biogeochemical cycles. However, knowledge is scarce about the record of (sub-)fossil coccolith assemblages in the Southern Ocean, which constitute invaluable indicators for palaeoenvironmental reconstructions. This study investigates coccolith assemblages preserved in surface sediments of southernmost Chile and across the Drake Passage that were retrieved during R/V Polarstern expedition PS97. We focused on the coccolith response to steep environmental gradients across the frontal system of the Antarctic Circumpolar Current and to hydrodynamic and post-depositional processes occurring in this region. We used statistical analyses to explore which environmental parameters influenced the coccolith assemblages by means of cluster and redundancy analyses. We specifically assessed the morphological diversity of the dominant taxa, i.e. Emiliania huxleyi, emphasizing biogeographical variability of morphotypes, coccolith sizes and calcite carbonate mass estimations. High coccolith abundances and species diversity compared to studies in the same area and in other sectors of the Southern Ocean occur, with a high species richness especially south of the Polar Front. While the surface sediments offshore Chile and north of the Polar Front provide suitable material to reconstruct overlying surface ocean conditions, further factors such as temporary thriving coccolithophore communities in the surface waters or transport of settling coccoliths via surface and bottom currents and eddies influence the (sub-)fossil coccolith assemblages south of the Polar Front. Additionally, deeper samples in the southern part of the study area are particularly affected by selective carbonate dissolution. We identified five E. huxleyi morphotypes (A, A overcalcified, R, B/C and O) and estimated coccolith carbonate masses on the basis of scanning electron microscope images. E. huxleyi morphologies reflect diverging biogeographical distributions, trending towards smaller and lighter coccoliths to the south and emphasizing the importance of documenting those morphologies in relation to changing environmental conditions to assess their response to projected environmental change in the Southern Ocean.

Continuous multiyear records of sediment-trap-gained microorganism fluxes are scarce. Such studies are important to identify and to understand the main forcings behind seasonal and multiannual evolution of microorganism flux dynamics. Here, we assess the long-term flux variations and population dynamics of diatoms, coccolithophores, calcareous and organic dinoflagellate cysts, foraminifera and pteropods in the eastern boundary upwelling ecosystem of the Canary Current. A multiannual, continuous sediment trap experiment was conducted at the mooring site CBeu (Cap Blanc eutrophic; ∼20∘ N, 18∘ W; trap depth is ca. 1300 m) off Mauritania (northwest Africa), between June 2003 and March 2008. Throughout the study, the reasonably consistent good match of fluxes of microorganisms and bulk mass reflects the seasonal occurrence of the main upwelling season and relaxation and the contribution of microorganisms to mass flux off Mauritania. A clear successional pattern of microorganisms, i.e., primary producers followed by secondary producers, is not observed. High fluxes of diatoms, coccolithophores, organic dinoflagellate cysts, and planktonic foraminifera occur simultaneously. Peaks of calcareous dinoflagellate cysts and pteropods mostly occurred during intervals of upwelling relaxation. A striking feature of the temporal variability of population occurrences is the persistent pattern of seasonal groups contributions. Species of planktonic foraminifera, diatoms, and organic dinoflagellate cysts typical of coastal upwelling, as well as cooler-water planktonic foraminifera and the coccolithophore Gephyrocapsa oceanica, are abundant at times of intense upwelling (late winter through early summer). Planktonic foraminifera and calcareous dinoflagellate cysts are dominant in warm pelagic surface waters, and all pteropod taxa are more abundant in fall and winter when the water column stratifies. Similarly, coccolithophores of the upper and lower photic zones, together with Emiliania huxleyi, and organic dinoflagellate cysts dominate the assemblage during phases of upwelling relaxation and deeper layer mixing. A significant shift in the “regular” seasonal pattern of taxa relative contribution is observed between 2004 and 2006. Benthic diatoms strongly increased after fall 2005 and dominated the diatom assemblage during the main upwelling season. Additional evidence for a change in population dynamics is the short dominance of the coccolithophore Umbilicosphaera annulus, the occurrence of the pteropod Limacina bulimoides and the strong increase in the flux of calcareous dinoflagellate cysts, abundant in warm tropical oligotrophic waters south of the study area after fall 2005. Altogether, this suggests that pulses of southern waters were transported to the sampling site via the northward Mauritania Current. Our multiannual trap experiment provides a unique opportunity to characterize temporal patterns of variability that can be extrapolated to other eastern boundary upwelling ecosystems (EBUEs), which are experiencing or might experience similar future changes in their plankton community.

Mesoscale eddies are abundant in the eastern tropical North Atlantic and act as oases for phytoplankton growth due to local enrichment of nutrients in otherwise oligotrophic waters. It is not clear whether these eddies can efficiently transfer organic carbon and other flux components to depth and if they are important for the marine carbon budget. Due to their transient and regionally restricted nature, measurements of eddies' contribution to bathypelagic particle flux are difficult to obtain. Rare observations of export flux associated with low-oxygen eddies have suggested efficient export from the surface to the deep ocean, indicating that organic carbon flux attenuation might be low. Here we report on particle flux dynamics north of the Cabo Verde islands at the oligotrophic Cape Verde Ocean Observatory (CVOO; approx. 17∘35′ N, 24∘15′ W). The CVOO site is located in the preferred pathways of highly productive eddies that ultimately originate from the Mauritanian upwelling region. Between 2009 and 2016, we collected biogenic and lithogenic particle fluxes with sediment traps moored at ca. 1 and 3 km water depths at the CVOO site. From concurrent hydrography and oxygen observations, we confirm earlier findings that highly productive eddies are characterized by colder and less saline waters and a low-oxygen signal as well. Overall, we observed quite consistent seasonal flux patterns during the passage of highly productive eddies in the winters of 2010, 2012 and 2016. We found flux increases at 3 km depth during October–November when the eddies approached CVOO and distinct flux peaks during February–March, clearly exceeding low oligotrophic background fluxes during winter 2011 and showing an enhanced particle flux seasonality. During spring, we observed a stepwise flux decrease leading to summer flux minima. The flux pattern of biogenic silicate (BSi) showed a stronger seasonality compared to organic carbon. Additionally, the deep fluxes of total mass showed an unusually higher seasonality compared to the 1 km traps. We assume that BSi and organic carbon/lithogenic material had different sources within the eddies. BSi-rich particles may originate at the eddy boundaries where large diatom aggregates are formed due to strong shear and turbulence, resulting in gravitational settling and, additionally, in an active local downward transport. Organic carbon associated with lithogenic material is assumed to originate from the interior of eddies or from mixed sources, both constituting smaller, dust-ballasted particles. Our findings suggest that the regularly passing highly productive eddies at CVOO repeatedly release characteristic flux signals to the bathypelagic zone during winter–spring seasons that are far above the oligotrophic background fluxes and sequester higher organic carbon than during oligotrophic settings. However, the reasons for a lower carbon flux attenuation below eddies remain elusive.