Bermuda Atlantic Time-series Study Research Articles

Recurring seasonality exposes dominant species and niche partitioning strategies of open ocean picoeukaryotic algae

Ocean spring phytoplankton blooms are dynamic periods important to global primary production. We document vertical patterns of a diverse suite of eukaryotic algae, the prasinophytes, in the North Atlantic Subtropical Gyre with monthly sampling over four years at the Bermuda Atlantic Time-series Study site. Water column structure was used to delineate seasonal stability periods more ecologically relevant than seasons defined by calendar dates. During winter mixing, tiny prasinophytes dominated by Class II comprise 46 ± 24% of eukaryotic algal (plastid-derived) 16S rRNA V1-V2 amplicons, specifically Ostreococcus Clade OII, Micromonas commoda, and Bathycoccus calidus. In contrast, Class VII are rare and Classes I and VI peak during warm stratified periods when surface eukaryotic phytoplankton abundances are low. Seasonality underpins a reservoir of genetic diversity from multiple prasinophyte classes during warm periods that harbor ephemeral taxa. Persistent Class II sub-species dominating the winter/spring bloom period retreat to the deep chlorophyll maximum in summer, poised to seed the mixed layer upon winter convection, exposing a mechanism for initiating high abundances at bloom onset. Comparisons to tropical oceans reveal broad distributions of the dominant sub-species herein. This unparalleled window into temporal and spatial niche partitioning of picoeukaryotic primary producers demonstrates how key prasinophytes prevail in warm oceans.

Evaluation of three contrasting models in estimating primary production from ocean color remote sensing using long-term time-series data at oceanic and coastal sites

Accurate estimates of depth-integrated Net Primary Production (NPP, mg C m−2 d−1) and the creation of a robust climate data record of NPP for the global oceans are essential goals of the ocean color remote sensing community. Here, we take advantage of in situ NPP measurements from three long-term time-series sites, the HOT (Hawaii Ocean Time-series), BATS (Bermuda Atlantic Time-series Study) and CARIACO (Ocean Time-Series Program from the Cariaco basin), spanning over 30 years to evaluate three contrasting models in estimating NPP from ocean color remote sensing. These models for NPP estimation include the Absorption-based Model (AbPM), which relies on phytoplankton absorption coefficient, the Vertically Generalized Production Model (VGPM), which centers on chlorophyll-a concentration, and the Carbon-based Productivity Model (CbPM) centering on phytoplankton carbon. In addition to the accuracy of NPP estimation from these models, we laid great emphasis on evaluating their skills in capturing the monthly to seasonal variations and interannual trends in NPP at the three sites. Comparison with in situ NPP at all three long-term sites (∼20 years) showed that AbPM yielded the highest coefficient of determination (R2 = 0.67) and the lowest uncertainties (Bias = 0.03 and unbiased root mean square difference = 0.17). Seasonal and interannual variations apparent in the in situ NPP time-series records were best captured by AbPM. These results showcase the robust capabilities of AbPM and its superiority for global carbon cycling and climate change studies, largely because it takes into account optical and photosynthetic parameters of local phytoplankton.

Forty years of ocean acidification observations (1983–2023) in the Sargasso Sea at the Bermuda Atlantic Time-series Study site

Ocean physical and biogeochemical conditions are rapidly changing over time. Forty years of observations from 1983 to 2023 collected at the Bermuda Atlantic Time-series Study (BATS) site near Bermuda in the North Atlantic Ocean shows continuing trends of surface warming, increase in salinity, loss of dissolved oxygen (DO), increase in carbon dioxide (CO2), and ocean acidification (OA) effects. Over this period, the ocean has warmed by about +1°C, increased in salinity by +0.136, and lost DO by 12.5 µmol kg−1 or ~6%. Since the 1980s, ocean dissolved inorganic carbon (DIC), total alkalinity (TA), a tracer of anthropogenic CO2 (CTrOCA), and fugacities/partial pressures of CO2 (i.e., fCO2 and pCO2) have continued to increase substantially, with no evidence of a reduction in the rates of change over time. Contemporaneously, ocean pH has decreased by ~0.1 pH units [with ocean acidity (i.e., H+) increasing by >30%], and the saturation states of calcium carbonate minerals (Ωcalcite and Ωaragonite) have decreased. These OA indicators show that the chemical conditions for calcification have become less favorable over the past 40 years. Updating of data and trends at the BATS site show how ocean chemistry of the 2020s is now outside the range observed in the 1980s, and how essential these data are for predicting the response of ocean chemistry and marine ecosystems to future shifting earth and ocean conditions.

Enhanced Turbulent Diapycnal Mixing in the Northern Sargasso Sea Inferred From a Finescale Parameterization

AbstractThe Sargasso Sea, accommodating marine species and mode water, is highly productive relative to other oligotrophic subtropical regions with intense air‐sea carbon and heat exchanges. Turbulent diapycnal mixing regulates these processes and is generally thought to be weakened in a more stratified ocean under anthropogenic forcing. However, by applying a finescale parameterization to long‐term hydrographic profiles collected during the Bermuda Atlantic Time‐Series Study (BATS), we show that its intensity has become stronger from 1994 to 2019 in the permanent thermocline. The enhanced turbulent diapycnal mixing is mainly attributed to the increase of wind power on internal waves along the Gulf Stream extension. Numerical simulations suggest that stronger internal waves excited along the Gulf Stream extension radiate downward and equatorward, causing enhanced turbulent diapycnal mixing in the northern Sargasso Sea including the BATS station. The findings have important implications for understanding responses of mode water and ecosystem in the Sargasso Sea to anthropogenic forcing.

Morphological and taxonomic diversity of mesozooplankton is an important driver of carbon export fluxes in the ocean.

Mesozooplankton is a very diverse group of small animals ranging in size from 0.2 to 20 mm not able to swim against ocean currents. It is a key component of pelagic ecosystems through its roles in the trophic networks and the biological carbon pump. Traditionally studied through microscopes, recent methods have been however developed to rapidly acquire large amounts of data (morphological, molecular) at the individual scale, making it possible to study mesozooplankton using a trait-based approach. Here, combining quantitative imaging with metabarcoding time-series data obtained in the Sargasso Sea at the Bermuda Atlantic Time-series Study (BATS) site, we showed that organisms' transparency might be an important trait to also consider regarding mesozooplankton impact on carbon export, contrary to the common assumption that just size is the master trait directing most mesozooplankton-linked processes. Three distinct communities were defined based on taxonomic composition, and succeeded one another throughout the study period, with changing levels of transparency among the community. A co-occurrences' network was built from metabarcoding data revealing six groups of taxa. These were related to changes in the functioning of the ecosystem and/or in the community's morphology. The importance of Diel Vertical Migration at BATS was confirmed by the existence of a group made of taxa known to be strong migrators. Finally, we assessed if metabarcoding can provide a quantitative approach to biomass and/or abundance of certain taxa. Knowing more about mesozooplankton diversity and its impact on ecosystem functioning would allow to better represent them in biogeochemical models.

Anthropogenic carbon estimation in the surface ocean from atmospheric CO2 fugacity at the BATS (Bermuda Atlantic Time-series Study) station

In surface seawater, it is usually very difficult to quantify anthropogenic carbon concentrations. Many processes (such as air-sea exchanges of gases and heat, biological activity, and mixing of water masses), are at play and often on different timescales. Thus, various hypotheses are used to estimate the anthropogenic concentrations in surface waters. Here, using the relatively long (1980s to present) time series data sets from the Bermuda Atlantic Time-series Study site (BATS; 31°40′N, 64°10′W) in the North Atlantic Ocean, we evaluate results based upon two different hypotheses. The results clearly confirm that it is very difficult to assess anthropogenic carbon concentrations in surface waters from sole oceanic properties. However, this study further indicates that at this ocean site, they can be appropriately determined from low-frequency variations of atmospheric CO2 concentrations. Consequently, the impact of anthropogenic carbon penetration in surface waters on their acidification could be predicted.

Authigenic mineral phases as a driver of the upper-ocean iron cycle.

Iron is important in regulating the ocean carbon cycle1. Although several dissolved and particulate species participate in oceanic iron cycling, current understanding emphasizes the importance of complexation by organic ligands in stabilizing oceanic dissolved iron concentrations2-6. However, it is difficult to reconcile this view of ligands as a primary control on dissolved iron cycling with the observed size partitioning of dissolved iron species, inefficient dissolved iron regeneration at depth or the potential importance of authigenic iron phases in particulate iron observational datasets7-12. Here we present a new dissolved iron, ligand and particulate iron seasonal dataset from the Bermuda Atlantic Time-series Study (BATS) region. We find that upper-ocean dissolved iron dynamics were decoupled from those of ligands, which necessitates a process by which dissolved iron escapes ligand stabilization to generate a reservoir of authigenic iron particles that settle to depth. When this 'colloidal shunt' mechanism was implemented in a global-scale biogeochemical model, it reproduced both seasonal iron-cycledynamics observations and independent global datasets when previous models failed13-15. Overall, we argue that the turnover of authigenic particulate iron phases must be considered alongside biological activity and ligands in controlling ocean-dissolved iron distributions and the coupling between dissolved and particulate iron pools.

Subtropical Gyre Nutrient Cycling in the Upper Ocean: Insights From a Nutrient‐Ratio Budget Method

AbstractWe use a nutrient‐ratio budget method to investigate the relative importance of different nutrient source and sink terms at time‐series Station ALOHA and Bermuda Atlantic Time‐series Study (BATS) in the North Pacific and North Atlantic subtropical gyres, respectively. At mean state conditions over annual and multi‐year time scales, vertical phosphate () supply from the subsurface accounts for ∼60% of the total phosphorus supply at both sites. Dissolved organic matter transport and zooplankton excretion are more important phosphorous export pathways than sinking particles at Station ALOHA and BATS. The nutrient‐ratio budget approach provides quantitative, observation‐based constraints on nutrient sources and sinks in the surface ocean, which helps improve our understanding of the biological carbon pump in oligotrophic oceans.

The photic-aphotic divide is a strong ecological and evolutionary force determining the distribution of ciliates (Alveolata, Ciliophora) in the ocean.

The bulk of knowledge on marine ciliates is from shallow and/or sunlit waters. We studied ciliate diversity and distribution across epi- and mesopelagic oceanic waters, using DNA metabarcoding and phylogeny-based metrics. We analyzed sequences of the 18S rRNA gene (V4 region) from 369 samples collected at 12 depths (0-1,000 m) at the Bermuda Atlantic Time-series Study site of the Sargasso Sea (North Atlantic) monthly for 3 years. The comprehensive depth and temporal resolutions analyzed led to three main findings. First, there was a gradual but significant decrease in alpha-diversity (based on Faith's phylogenetic diversity index) from surface to 1,000-m waters. Second, multivariate analyses of beta-diversity (based on UniFrac distances) indicate that ciliate assemblages change significantly from photic to aphotic waters, with a switch from Oligotrichea to Oligohymenophorea prevalence. Third, phylogenetic placement of sequence variants and clade-level correlations (EPA-ng and GAPPA algorithms) show Oligotrichea, Litostomatea, Prostomatea, and Phyllopharyngea as anti-correlated with depth, while Oligohymenophorea (especially Apostomatia) have a direct relationship with depth. Two enigmatic environmental clades include either prevalent variants widely distributed in aphotic layers (the Oligohymenophorea OLIGO5) or subclades differentially distributed in photic versus aphotic waters (the Discotrichidae NASSO1). These results settle contradictory relationships between ciliate alpha-diversity and depth reported before, suggest functional changes in ciliate assemblages from photic to aphotic waters (with prevalence of algivory and mixotrophy versus omnivory and parasitism, respectively), and indicate that contemporary taxon distributions in the vertical profile have been strongly influenced by evolutionary processes. Integration of DNA sequences with organismal data (microscopy, functional experiments) and development of databases that link these sources of information remain as major tasks to better understand ciliate diversity, ecological roles, and evolution in the ocean.

Ecophysiological basis of spatiotemporal patterns in picophytoplankton pigments in the global ocean

Information on the intracellular content and functional diversity of phytoplankton pigments can provide valuable insight on the ecophysiological state of primary producers and the flow of energy within aquatic ecosystems. Combined global datasets of analytical flow cytometry (AFC) cell counts and High-Performance Liquid Chromatography (HPLC) pigment concentrations were used to examine vertical and seasonal variability in the ratios of phytoplankton pigments in relation to indices of cellular photoacclimation. Across all open ocean datasets, the weight-to-weight ratio of photoprotective to photosynthetic pigments showed a strong depth dependence that tracked the vertical decline in the relative availability of light. The Bermuda Atlantic Time-series Study (BATS) dataset revealed a general increase in surface values of the relative concentrations of photoprotective carotenoids from the winter-spring phytoplankton communities dominated by low-light acclimated eukaryotic microalgae to the summer and early autumn communities dominated by high-light acclimated picocyanobacteria. In Prochlorococcus-dominated waters, the vertical decline in the relative contribution of photoprotective pigments to total pigment concentration could be attributed in large part to changes in the cellular content of photosynthetic pigments (PSP) rather than photoprotective pigments (PPP), as evidenced by a depth-dependent increase of the intracellular concentration of the divinyl chlorophyll-a (DVChl-a) whilst the intracellular concentration of the PPP zeaxanthin remained relatively uniform with depth. The ability of Prochlorococcus cells to adjust their DVChl-a cell-1 over a large gradient in light intensity was reflected in more highly variable estimates of carbon-to-Chl-a ratio compared to those reported for other phytoplankton groups. This cellular property is likely the combined result of photoacclimatory changes at the cellular level and a shift in dominant ecotypes. Developing a mechanistic understanding of sources of variability in pigmentation of picocyanobacteria is critical if the pigment markers and bio-optical properties of these cells are to be used to map their biogeography and serve as indicators of photoacclimatory state of subtropical phytoplankton communities more broadly. It would also allow better assessment of effects on, and adaptability of phytoplankton communities in the tropical/subtropical ocean due to climate change.

Nitrous Oxide Distributions in the Oxygenated Water Column of the Sargasso Sea

ABSTRACT This study presents dissolved nitrous oxide (N2O) concentrations in the water column at the Bermuda Atlantic Time-series Study (BATS) station and uses a subset of these measurements to estimate air-to-sea flux for four specific time points between September 2018 and June 2019. N2O concentrations at BATS were in the range of 4.0 nmol L−1–16.9 nmol L−1, with vertical profiles which were the mirror inverse of dissolved oxygen. Regardless of season, N2O concentration maxima were found within the oxygen minimum zone (OMZ). The highest maximum N2O values were observed in November and lowest in October. As the water column at BATS remains consistently at dissolved oxygen concentrations greater than 140 µmol L−1, and therefore aerobic, we assume that the bulk of N2O production occurs through nitrification. A nitrification source is supported by a correlation between excess N2O (ΔN2O) below the mixed layer, apparent oxygen utilization (AOU) and nitrate concentrations. We estimate a pooled average yield of 0.027% to 0.038% N2O from nitrification at BATS. Finally, estimates of air–sea exchange of N2O using regional average monthly wind speeds indicated that this region acts as a weak source or a sink of atmospheric N2O, and varies between months.

Uncertainties in ocean biogeochemical simulations: Application of ensemble data assimilation to a one-dimensional model

Marine biogeochemical (BGC) models are highly uncertain in their parameterization. The value of the BGC parameters are poorly known and lead to large uncertainties in the model outputs. This study focuses on the uncertainty quantification of model fields and parameters within a one-dimensional (1-D) ocean BGC model applying ensemble data assimilation. We applied an ensemble Kalman filter provided by the Parallel Data Assimilation Framework (PDAF) into a 1-D vertical configuration of the BGC model Regulated Ecosystem Model 2 (REcoM2) at two BGC time-series stations: the Bermuda Atlantic Time-series Study (BATS) and the Dynamique des Flux Atmosphériques en Méditerranée (DYFAMED). We assimilated 5-day satellite chlorophyll-a (chl-a) concentration and monthly in situ net primary production (NPP) data for 3 years to jointly estimate 10 preselected key BGC parameters and the model state. The estimated set of parameters resulted in improvements in the model prediction up to 66% for the surface chl-a and 56% for NPP. Results show that assimilating satellite chl-a concentration data alone degraded the prediction of NPP. Simultaneous assimilation of the satellite chl-a data and in situ NPP data improved both surface chl-a and NPP simulations. We found that correlations between parameters preclude estimating parameters independently. Co-dependencies between parameters also indicate that there is not a unique set of optimal parameters. Incorporation of proper uncertainty estimation in BGC predictions, therefore, requires ensemble simulations with varying parameter values.

Fragility of marine photosynthesis

Ecosystem fragility is an often used term in oceanography yet to this day it lacks a precise and widely accepted definition. Defining and subsequently quantifying fragility would be of great value, for such measures could be used to objectively ascertain the level of risk marine ecosystems face. Risk assessments could further be used to define the level of protection a given ocean region requires from economic activity, such as fisheries. With this aim we introduce to the oceanographic literature the concepts of marginal production and fragility, which we define for marine photosynthesis, the base of the oceanic food web. We demonstrate that marine photosynthesis is always fragile with respect to light, implying variability in surface irradiance acts unfavourably on biomass. We also demonstrate that marine photosynthesis can be both fragile and antifragile with respect to the mixed-layer depth, implying variability in mixed-layer depth can act both favourably and unfavourably on biomass. Quantification of marginal production and fragility is presented on data from two open ocean stations: Hawaii Ocean Time Series and Bermuda Atlantic Time-Series Study. Seasonal cycle of biomass is modelled and the effects of primary production fragility are analysed. A new tipping point for marine phytoplankton is identified in the form of a depth horizon. Using the new definitions presented here a rich archive of data can be used straightforwardly to quantify primary production fragility. The definitions can also be used to predict when primary production enters the fragile state during the seasonal cycle.

Linkages Among Dissolved Organic Matter Export, Dissolved Metabolites, and Associated Microbial Community Structure Response in the Northwestern Sargasso Sea on a Seasonal Scale.

Deep convective mixing of dissolved and suspended organic matter from the surface to depth can represent an important export pathway of the biological carbon pump. The seasonally oligotrophic Sargasso Sea experiences annual winter convective mixing to as deep as 300 m, providing a unique model system to examine dissolved organic matter (DOM) export and its subsequent compositional transformation by microbial oxidation. We analyzed biogeochemical and microbial parameters collected from the northwestern Sargasso Sea, including bulk dissolved organic carbon (DOC), total dissolved amino acids (TDAA), dissolved metabolites, bacterial abundance and production, and bacterial community structure, to assess the fate and compositional transformation of DOM by microbes on a seasonal time-scale in 2016–2017. DOM dynamics at the Bermuda Atlantic Time-series Study site followed a general annual trend of DOC accumulation in the surface during stratified periods followed by downward flux during winter convective mixing. Changes in the amino acid concentrations and compositions provide useful indices of diagenetic alteration of DOM. TDAA concentrations and degradation indices increased in the mesopelagic zone during mixing, indicating the export of a relatively less diagenetically altered (i.e., more labile) DOM. During periods of deep mixing, a unique subset of dissolved metabolites, such as amino acids, vitamins, and benzoic acids, was produced or lost. DOM export and compositional change were accompanied by mesopelagic bacterial growth and response of specific bacterial lineages in the SAR11, SAR202, and SAR86 clades, Acidimicrobiales, and Flavobacteria, during and shortly following deep mixing. Complementary DOM biogeochemistry and microbial measurements revealed seasonal changes in DOM composition and diagenetic state, highlighting microbial alteration of the quantity and quality of DOM in the ocean.

Seasonal changes in seawater calcium and alkalinity in the Sargasso Sea and across the Bermuda carbonate platform

Ocean acidification may shift coral reefs from a state of net ecosystem calcification (+NEC) to net ecosystem dissolution (–NEC). Changes in NEC are typically inferred from either measured or calculated total alkalinity (TA) or the dissolved calcium (Ca) to salinity ratio relative to a reference value. The alkalinity anomaly technique has historically been the primary method to estimate NEC due to the greater analytical challenges and uncertainty associated with dissolved Ca measurements in seawater. However, this method assumes that changes in salinity-normalized TA are exclusively the result of calcification and dissolution processes. In many cases, this assumption is valid, but in some environments additional processes can significantly influence seawater TA (e.g., nutrient fluxes and redox processes). Seawater Ca is unaffected or less sensitive to these processes, and therefore, Ca and TA anomalies can be used to estimate absolute or relative changes in NEC with greater confidence. Here, we present a two-year time series of monthly seawater Ca and TA measurements across the Bermuda carbonate platform and the nearby Bermuda Atlantic Time-series Study (BATS) location offshore. High precision Ca measurements (±6 μmol kg−1) were conducted using an improved spectrophotometric titration system and showed mostly good agreement with changes in TA over the same spatial and temporal scales. Ca and TA measurements across the Bermuda platform showed seasonal fluctuations relative to offshore waters, with +NEC during summer months and near-zero or possible –NEC (net dissolution) during winter months. These seasonal patterns were most pronounced at the inshore locations with the longest residence times (10+ days), which allow stronger biogeochemical signals to develop relative to the offshore source water. Although obtaining high accuracy and precision Ca measurements remains challenging, parallel measurements of Ca and TA from both inshore and offshore waters over a multi-annual timescale could strengthen the validity of predictions for when and where a reef system, such as the Bermuda platform, may shift from +NEC to –NEC.

Microscopy and DNA‐based characterization of sinking particles at the Bermuda Atlantic Time‐series Study station point to zooplankton mediation of particle flux

AbstractPlankton‐derived, microscopic, and macroscopic sinking aggregates constitute most of the particulate organic carbon (POC) flux in the oceans. While the flux of particulate organic matter and associated elements has been quantified at the Bermuda Atlantic Time‐series Study (BATS) station for several decades, we lack an understanding of the source and composition of sinking particles, as well as the fate of predominant phytoplankton taxa. We determined the composition of individual sinking particles and their microbial communities in the upper 300 m depth at the BATS station in fall 2017 and spring 2018 by image analysis and V4 amplicon sequencing of the 16S and 18S rRNA genes. The sinking particles were primarily composed of phytodetrital aggregates, fecal aggregates, and fecal pellets. In the fall, phytodetrital aggregates were numerically dominant and drove the majority of the POC flux; however, in the spring, particle flux of all particle categories declined below 150 m, and the POC flux at 200 m shifted to one driven by fecal aggregates. The relative composition of the microbial communities associated with phytodetrital and fecal aggregates were statistically indistinguishable in both seasons, and prokaryotic taxa known to be associated with the gut microbiomes of zooplankton were indicators of the sinking particles. Our results point to the utilization and modification of sinking particles by resident midwater zooplankton populations, and to fecal pellets as the predominant mechanism transporting picophytoplankton to depth.

Reconciling Between Optical and Biological Determinants of the Euphotic Zone Depth

AbstractThe conventional use of optically determined 1% of surface photosynthetically available radiation (PAR) depth (Z1%PAR, λ = 400–700 nm) as a metric for the euphotic zone depth (Zeu) has been a matter of debate for several decades because of frequent inconsistencies with the base of euphotic zone determined biologically, that is, the compensation depth (Zc). In this study, we attempt to reconcile between optical and biological determinants of the euphotic zone through the use of a large data set of coincidental profiles of downwelling irradiance and primary production. These measurements cover open ocean waters in the tropics, subtropics, temperate regions, and from two time‐series stations, the Hawaii Ocean Time‐series (HOT) and the Bermuda Atlantic Time‐series Study (BATS). We report that, at least for these measurements, Z1%PAR is too shallow (by 14.1%) compared to Zc, while Z0.1%PAR is too deep (by 32.7%). Further, the irradiance at Zc (i.e., the compensation irradiance, Ic) varies by a factor of more than five, but its ratio to surface irradiance is relatively stable. In general, Ic corresponds to 0.48 ± 0.23% of surface PAR, or 0.87 ± 0.40% of surface usable solar radiation (USR, for λ = 400–560 nm), or 1.50 ± 0.67% of surface downwelling irradiance at 490 nm. These results suggest that Z0.5%PAR, or Z0.9%USR, or Z1.5%490 could be promising alternatives to bridge the optical and biological determinants of the euphotic zone depths for waters in the open ocean at low to middle latitudes.

Phytoplankton community composition and biomass in the oligotrophic Gulf of Mexico

Abstract Biomass and composition of the phytoplankton community were investigated in the deep-water Gulf of Mexico (GoM) at the edges of Loop Current anticyclonic eddies during May 2017 and May 2018. Using flow cytometry, high-performance liquid chromatography pigments and microscopy, we found euphotic zone integrated chlorophyll a of ~10 mg m−2 and autotrophic carbon ranging from 463 to 1268 mg m−2, dominated by picoplankton (<2 μm cells). Phytoplankton assemblages were similar to the mean composition at the Bermuda Atlantic Time-series Study site, but differed from the Hawaii Ocean Times-series site. GoM phytoplankton biomass was ~2-fold higher at the deep chlorophyll maximum (DCM) relative to the mixed layer (ML). Prochlorococcus and prymnesiophytes were the dominant taxa throughout the euphotic zone; however, other eukaryotic taxa had significant biomass in the DCM. Shallower DCMs were correlated with more prymnesiophytes and prasinophytes (Type 3) and reduced Prochlorococcus. These trends in ML and DCM taxonomic composition likely reflect relative nutrient supply—with ML populations relying on remineralized ammonium as a nitrogen source, and the taxonomically diverse DCM populations using more nitrate. These spatially separated phytoplankton communities represent different pathways for primary production, with a dominance of picoplankton in the ML and more nano- and microplankton at the DCM.

Acceleration of ocean warming, salinification, deoxygenation and acidification in the surface subtropical North Atlantic Ocean

Ocean chemical and physical conditions are changing. Here we show decadal variability and recent acceleration of surface warming, salinification, deoxygenation, carbon dioxide (CO2) and acidification in the subtropical North Atlantic Ocean (Bermuda Atlantic Time-series Study site; 1980s to present). Surface temperatures and salinity exhibited interdecadal variability, increased by ~0.85 °C (with recent warming of 1.2 °C) and 0.12, respectively, while dissolved oxygen levels decreased by ~8% (~2% per decade). Concurrently, seawater DIC, fCO2 (fugacity of CO2) and anthropogenic CO2 increased by ~8%, 22%, and 72% respectively. The winter versus summer fCO2 difference increased by 4 to 8 µatm decade−1 due to seasonally divergent thermal and alkalinity changes. Ocean pH declined by 0.07 (~17% increase in acidity) and other acidification indicators by ~10%. Over the past nearly forty years, the highest increase in ocean CO2 and ocean acidification occurred during decades of weakest atmospheric CO2 growth and vice versa.

Controls on carbon export in the subtropical North Atlantic

We present a lower trophic level pelagic ecosystem model that allows the investigation of the role of hydrography, bacterial remineralization and detritus consumption on the strength of carbon export and its attenuation. We apply the model to investigate the controls on the carbon export at the Bermuda Atlantic Time-Series Study site (BATS) and the European Station for Time series in the Ocean, Canary Islands (ESTOC) in the subtropical North Atlantic. In previous field studies, export ratios at 200 m (the ratio between particulate carbon export below the euphotic zone and primary production) were found to be 300–400% smaller at ESTOC compared to BATS. Our model results show that the magnitude and temporal variability of primary productivity and modeled carbon export are modulated by the intensity and duration of vertical mixing events. BATS, with a more dynamic physical environment, has winter mixed-layer depths on average 80 m deeper than ESTOC and is characterized by pulses of enhanced productivity and export. Our model demonstrates the influence of hydrography on export attenuation through (i) more stable water column dynamics at ESTOC that increase particle remineralization time scales and weaken export strength at ESTOC compared to BATS; and (ii) higher water temperatures at BATS in the mesopelagic between 200 and 500 m that increase remineralization rates compared to ESTOC. This results in reduced differences in export ratios within the mesopelagic between both stations that is confirmed by observations. Strengthening remineralization through (i) adding zooplankton feeding on detritus and (ii) increasing bacterial remineralization rates decrease export at all depths at both stations, and increase the modeled difference in 200 m export ratios between BATS and ESTOC from 7% to 17%, a difference that is still much smaller than observed. While we could demonstrate the skill of our model in testing the mechanisms that could lead to differences in regional carbon export, our results indicate that hydrography-driven residence times, zooplankton feeding on detritus and enhanced bacterial remineralization rates alone are insufficient in driving export ratio differences between BATS and ESTOC. On the other hand, modeled export proved to be highly sensitive to prescribed particle sinking speeds. Thus, community and site dependent processes that lead to variations in particle sinking speed and remineralization, together with potential differences in vertical migration by zooplankton and horizontal transport, may be additional processes explaining the observed differences in export ratios at BATS and ESTOC.