Bermuda Atlantic Time-series Study Site 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.

Seasonal and daily patterns in known dissolved metabolites in the northwestern Sargasso Sea

AbstractOrganic carbon in seawater plays a significant role in the global carbon cycle. The concentration and composition of dissolved organic carbon reflect the activity of the biological community and chemical reactions that occur in seawater. From 2016 to 2019, we repeatedly sampled the oligotrophic northwest Sargasso Sea in the vicinity of the Bermuda Atlantic Time‐series Study site (BATS) to quantitatively follow known compounds within the pool of dissolved organic matter in the upper 1000 m of the water column. Most metabolites showed surface enrichment, and 83% of the metabolites had significantly lower concentrations with increasing depth. Dissolved metabolite concentrations most notably revealed temporal variability. Fourteen metabolites displayed seasonality that was repeated in each of the 4 yr sampled. Concentrations of vitamins, including pantothenic acid (vitamin B5) and riboflavin (vitamin B2), increased annually during winter periods when mixed layer depths were deepest. During diel sampling, light‐sensitive riboflavin decreased significantly during daylight hours. The temporal variability in metabolites at BATS was less than the spatial variability in metabolites from a previous sample set collected over a broad latitudinal range in the western Atlantic Ocean. The metabolites examined in this study are all components of central carbon metabolism. By examining these metabolites at finer resolution and in a time‐series, we begin to provide insights into the chemical compounds that may be exchanged by microorganisms in marine systems, data which are fundamental to understanding the chemical response of marine systems to future changes in climate.

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.

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.

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.

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.

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.

Machine learning identifies a strong association between warming and reduced primary productivity in an oligotrophic ocean gyre

Phytoplankton play key roles in the oceans by regulating global biogeochemical cycles and production in marine food webs. Global warming is thought to affect phytoplankton production both directly, by impacting their photosynthetic metabolism, and indirectly by modifying the physical environment in which they grow. In this respect, the Bermuda Atlantic Time-series Study (BATS) in the Sargasso Sea (North Atlantic gyre) provides a unique opportunity to explore effects of warming on phytoplankton production across the vast oligotrophic ocean regions because it is one of the few multidecadal records of measured net primary productivity (NPP). We analysed the time series of phytoplankton primary productivity at BATS site using machine learning techniques (ML) to show that increased water temperature over a 27-year period (1990–2016), and the consequent weakening of vertical mixing in the upper ocean, induced a negative feedback on phytoplankton productivity by reducing the availability of essential resources, nitrogen and light. The unbalanced availability of these resources with warming, coupled with ecological changes at the community level, is expected to intensify the oligotrophic state of open-ocean regions that are far from land-based nutrient sources.

Diel, seasonal, and interannual patterns in mesozooplankton abundance in the Sargasso Sea

AbstractTemporal changes in mesozooplankton abundance affect planktonic food web interactions and biogeochemistry. We enumerated mesozooplankton from monthly day and night tows in the epipelagic zone at the Bermuda Atlantic Time-series Study (BATS) site in the Sargasso Sea (1999–2010). Abundances of each taxon were determined using a ZooScan imaging system and microscopy. Generalized linear models were used to determine environmental parameters that best explained abundance patterns. Taxa with pronounced diel vertical migration included euphausiids, amphipods, Limacina spp. pteropods, and other shelled pteropods. Taxa with a pronounced spring abundance peak included euphausiids, appendicularians, and Limacina spp., while harpacticoid copepods peaked in late summer, and calanoid copepods in late winter/early spring and summer. Many taxa increased in 2003, coincident with a diatom bloom and the largest primary production peak in the time series. Long-term, increasing trends occurred in calanoid and oncaeid copepods, and ostracods, with barnacle nauplii significantly increasing. Sub-decadal-scale climate oscillations and long-term warming may be driving decreases in shelled pteropods and appendicularians. Chaetognath abundance increased in response to increased density of a major prey taxon, calanoid copepods. Calanoid copepods and ostracods increased with increasing water column stratification index and the Atlantic Multidecadal Oscillation index, indicating warmer sea surface temperatures favour these taxa.

Mixing regime-dependent causality between phytoplankton and bacteria in the subtropical North Atlantic Ocean ecosystem

MEPS Marine Ecology Progress Series Contact the journal Facebook Twitter RSS Mailing List Subscribe to our mailing list via Mailchimp HomeLatest VolumeAbout the JournalEditorsTheme Sections MEPS 600:41-53 (2018) - DOI: https://doi.org/10.3354/meps12643 Mixing regime-dependent causality between phytoplankton and bacteria in the subtropical North Atlantic Ocean ecosystem Hyewon Kim1,3,*, Dong Eun Lee2, Hugh W. Ducklow1 1Division of Biology and Paleo Environment, Lamont-Doherty Earth Observatory, Columbia University, Palisades, NY 10964, USA 2Division of Ocean and Climate Physics, Lamont-Doherty Earth Observatory, Columbia University, Palisades, NY 10964, USA 3Present address: Department of Environmental Sciences, University of Virginia, Charlottesville, VA 22904, USA *Corresponding author: hk8m@virginia.edu ABSTRACT: Ever since marine heterotrophic bacteria were understood to mediate carbon fluxes to upper trophic levels via the microbial loop, quantifying the coupling between bacteria and phytoplankton has been one of the main priorities for marine microbiologists. The complex nature of phytoplankton-bacterial interactions may lead to nonlinear and regime-dependent coupling, for which conventional statistical approaches such as cross-correlation provide an incomplete picture of the dynamics. Here, we employed a nonlinear method for detecting causality, called convergent cross mapping (CCM), to examine a causal linkage between phytoplankton (primary production) and bacteria (bacterial production) at the Bermuda Atlantic Time-series Study (BATS) site. First, we verified the robustness of our CCM models with synthetic time series output from the Fasham-Ducklow-McKelvie ecosystem model. Initially, we hypothesized a strong bi-directional causal link between phytoplankton and bacteria due to the importance of the microbial loop at BATS. However, the results from our CCM models highlight that phytoplankton-bacterial causal coupling depends on seasonally distinct mixing regimes. While there was no evidence for bi-directional causality between phytoplankton and bacteria during the mixing period, moderate unidirectional causality of bacteria on phytoplankton was detected during the stratification period. Our study reveals causal associations between the 2 major microbial loop processes, for which better quantification is needed to improve our understanding of carbon cycling and export via microbial food webs. KEY WORDS: Convergent cross mapping · CCM · Bermuda Atlantic Time-series Study · BATS · Primary production · Bacterial production · Sargasso Sea Full text in pdf format Supplementary material PreviousNextCite this article as: Kim H, Lee DE, Ducklow HW (2018) Mixing regime-dependent causality between phytoplankton and bacteria in the subtropical North Atlantic Ocean ecosystem. Mar Ecol Prog Ser 600:41-53. https://doi.org/10.3354/meps12643 Export citation RSS - Facebook - Tweet - linkedIn Cited by Published in MEPS Vol. 600. Online publication date: July 30, 2018 Print ISSN: 0171-8630; Online ISSN: 1616-1599 Copyright © 2018 Inter-Research.

Erratum to: Patterns of coccolithophore pigment change under global acidification conditions based on in-situ observations at BATS site between July 1990–Dec 2008

Coccolith production is an important part of the biogenic carbon cycle as the largest source of calcium carbonate on earth, accounting for about 75% of the deposition of carbon on the sea floor. Recent studies based on laboratory experiment results indicated that increasing anthropogenic CO2 in the atmosphere triggered global ocean acidification leading to a decrease of calcite or aragonite saturation and calcium carbonate, and to decreasing efficiency of carbon export/pumping to deep layers. In the present study, we analyzed about 20 years of field observations of coccolithophore pigment, dissolved inorganic carbon (DIC), nutrients, and temperatures from the Bermuda Atlantic Time-series Study (BATS) site and satellite remote sensing to investigate the variable tendency of the coccolithophore pigment, and to evaluate the influence of ocean acidification on coccolithophore biomass. The results indicated that there was a generally increasing tendency of coccolithophore pigment, coupled with increasing bicarbonate concentrations or decreasing carbonate ion concentration. The change of coccolithophore pigment was also closely associated with pH, nutrients, mixed layer depth (MLD), and temperature. Correlation analyses between coccolithophores and abiotic parameter imply that coccoliths production or coccolithophore pigment has increased with increasing acidification in the recent 20 years.

Twenty Years of Marine Carbon Cycle Observations at Devils Hole Bermuda Provide Insights into Seasonal Hypoxia, Coral Reef Calcification, and Ocean Acidification

Open–ocean observations have revealed gradual changes in seawater carbon dioxide (CO2) chemistry resulting from uptake of atmospheric CO2 and ocean acidification (OA), but, with few long–term records (>five years) of the coastal ocean that can reveal the pace and direction of environmental change. In this paper, observations collected from 1996 to 2016 at Harrington Sound, Bermuda, constitute one of the longest time–series of coastal ocean inorganic carbon chemistry. Uniquely, such changes can be placed into the context of contemporaneous offshore changes observed at the nearby Bermuda Atlantic Time-series Study (BATS) site. Onshore, surface dissolved inorganic carbon (DIC) and partial pressure of CO2 (pCO2; >10% change per decade) have increased and OA indicators such as pH and calcium carbonate (CaCO3) saturation state (Ω) decreased from 1996–2016 at a rate of two to three times that observed offshore at BATS. Such changes, combined with reduction of total alkalinity over time, reveal a complex interplay of biogeochemical processes influencing Bermuda reef metabolism, including net ecosystem production (NEP=gross primary production–autotrophic and heterotrophic respiration) and net ecosystem calcification (NEC=gross calcification–gross CaCO3 dissolution). These long–term data show ae seasonal shift between wintertime net heterotrophy and summertime net autotrophy for the entire Bermuda reef system. Over annual time-scales, the Bermuda reef system does not appear to be in trophic balance, but rather slightly net heterotrophic. In addition, the reef system is net accretive (i.e., gross calcification > gross CaCO3 dissolution), but there were occasional periods when the entire reef system appears to transiently shift to net dissolution. A previous five–year study of the Bermuda reef suggested that net calcification and net heterotrophy have both increased. Over the past twenty years, rates of net calcification and net heterotrophy determined for the Bermuda reef system have increased by ~30%, most likely due to increased coral nutrition occurring in concert with increased offshore productivity in the surrounding subtropical North Atlantic Ocean. Importantly, this long–term study reveals that other environmental factors can mitigate against the effects of ocean acidification on coral reef calcification, at least over the past couple of decades.

An Intercomparison of Dissolved Iron Speciation at the Bermuda Atlantic Time-series Study (BATS) Site: Results from GEOTRACES Crossover Station A

The organic complexation of dissolved iron (Fe) was determined in depth profile samples collected from GEOTRACES Crossover Station A, the Bermuda Atlantic Time-series Study (BATS) site, as part of the Dutch and U.S. GEOTRACES North Atlantic programs in June 2010 and November 2011, respectively. The two groups employed distinct competitive ligand exchange-adsorptive cathodic stripping voltammetry (CLE-AdCSV) methods, and resulting ligand concentrations and conditional stability constants from each profile were compared. Excellent agreement was found between the total ligand concentrations determined in June 2010 and the strongest, L1-type, ligand concentrations determined in November 2011. Yet a primary distinction between the datasets was the number of ligand classes observed: a single ligand class was characterized in the June 2010 profile while two ligand classes were observed in the November 2011 profile. To assess the role of differing interpretation approaches in determining final results, analysts exchanged titration data and accompanying parameters from the profiles for reinterpretation. The reinterpretation exercises highlighted the considerable influence of the sensitivity (S) parameter applied on interpretation results, consistent with recent intercalibration work on interpretation of copper speciation titrations. The potential role of titration data structure, humic-type substances, differing dissolved Fe concentrations, and seasonality are also discussed as possible drivers of the one versus two ligand class determinations between the two profiles, leading to recommendations for future studies of Fe-binding ligand cycling in the oceans.

Carbon flux from bio-optical profiling floats: Calibrating transmissometers for use as optical sediment traps

Our mechanistic understanding of the processes controlling the ocean's biological pump is limited, in part, by our lack of observational data at appropriate timescales. The “optical sediment trap” (OST) technique utilizes a transmissometer on a quasi-Lagrangian platform to collect sedimenting particles. This method could help fill the observational gap by providing autonomous measurements of particulate carbon (PC) flux in the upper mesopelagic ocean at high spatiotemporal resolution. Here, we used a combination of field measurements and laboratory experiments to test hydrodynamic and zooplankton-swimmer effects on the OST method, and we quantitatively calibrated this method against PC flux measured directly in same-platform, neutrally buoyant sediment traps (NBSTs) during 5 monthly cruises at the Bermuda Atlantic Time-series Study (BATS) site. We found a well-correlated, positive relationship (R2=0.66, n=15) between the OST proxy, and the PC flux measured directly using NBSTs. Laboratory tests showed that scattering of light from multiple particles between the source and detector was unlikely to affect OST proxy results. We found that the carbon-specific attenuance of sinking particles was larger than literature values for smaller, suspended particles in the ocean, and consistent with variable carbon: size relationships reported in the literature for sinking particles. We also found evidence for variability in PC flux at high spatiotemporal resolution. Our results are consistent with the literature on particle carbon content and optical properties in the ocean, and support more widespread use of the OST proxy, with proper site-specific and platform-specific calibration, to better understand variability in the ocean biological pump.

Seasonal to multi-decadal trends in apparent optical properties in the Sargasso Sea

Multi-decadal, monthly observations of optical and biogeochemical properties, made as part of the Bermuda Bio-Optics Project (BBOP) at the Bermuda Atlantic Time-series Study (BATS) site in the Sargasso Sea, allow for the examination of temporal trends in vertical light attenuation and their potential controls. Trends in the magnitude of the diffuse attenuation coefficient, Kd(λ), and a proxy for its spectral shape reflect changes in phytoplankton and chromophoric dissolved organic matter (CDOM) characteristics. The length and methodological consistency of this time series provide an excellent opportunity to extend analyses of seasonal cycles of apparent optical properties to interannual and decadal time scales. Here, we characterize changes in the magnitude and spectral shape proxy of diffuse attenuation coefficient spectra and compare them to available biological and optical data from the BATS time series program. The time series analyses reveal a 1.01%±0.18% annual increase of the magnitude of the diffuse attenuation coefficient at 443nm over the upper 75m of the water column while showing no significant change in selected spectral characteristics over the study period. These and other observations indicate that changes in phytoplankton rather than changes in CDOM abundance are the primary driver for the diffuse attenuation trends on multi-year timescales for this region. Our findings are inconsistent with previous decadal-scale global ocean water clarity and global satellite ocean color analyses yet are consistent with recent analyses of the BATS time series and highlight the value of long-term consistent observation at ocean time series sites.

Salp contributions to vertical carbon flux in the Sargasso Sea

We developed a one-dimensional model to estimate salp contributions to vertical carbon flux at the Bermuda Atlantic Time-series Study (BATS) site in the North Atlantic subtropical gyre for a 17-yr period (April 1994 to December 2011). We based the model parameters on published rates of salp physiology and experimentally determined sinking and decomposition rates of salp carcasses. Salp grazing was low during non-bloom conditions, but routinely exceeded 100% of chlorophyll standing stock and primary production during blooms. Fecal pellet production was the largest source of salp carbon flux (78% of total), followed by respiration below 200m (19%), sinking of carcasses (3%), and DOC excretion below 200m (<0.1%). Thalia democratica, Salpa fusiformis, Salpa aspera, Wheelia cylindrica, and Iasis zonaria were the five highest contributors, accounting for 95% of total salp-mediated carbon flux. Seasonally, salp flux was higher during spring-summer than fall-winter, due to seasonal changes in species composition and abundance. Salp carbon export to 200m was on average 2.3mgCm−2d−1 across the entire time series. This is equivalent to 11% of the mean 200m POC flux measured by sediment traps in the region. During years with significant salp blooms, however, annually-averaged salp carbon export was the equivalent of up to 60% of trap POC flux at 200m. Salp carbon flux attenuated slowly, and at 3200m the average modeled carbon from salps was 109% of the POC flux measured in sediment traps at that depth. Migratory and carcass carbon export pathways should also be considered (alongside fecal pellet flux) as facilitating carbon export to sequestration depths in future studies.

Apparent increase in coccolithophore abundance in the subtropical North Atlantic from 1990 to 2014

Abstract. As environmental conditions evolve with rapidly increasing atmospheric CO2, biological communities will change as species reorient their distributions, adapt, or alter their abundance. In the surface ocean, dissolved inorganic carbon (DIC) has been increasing over the past several decades as anthropogenic CO2 dissolves into seawater, causing acidification (decreases in pH and carbonate ion concentration). Calcifying phytoplankton, such as coccolithophores, are thought to be especially vulnerable to ocean acidification. How coccolithophores will respond to increasing carbon input has been a subject of much speculation and inspired numerous laboratory and mesocosm experiments, but how they are currently responding in situ is less well documented. In this study, we use coccolithophore (haptophyte) pigment data collected at the Bermuda Atlantic Time-series Study (BATS) site together with satellite estimates (1998–2014) of surface chlorophyll and particulate inorganic carbon (PIC) as a proxy for coccolithophore abundance to show that coccolithophore populations in the North Atlantic subtropical gyre have been increasing significantly over the past 2 decades. Over 1990–2012, we observe a 37 % increase in euphotic zone-integrated coccolithophore pigment abundance at BATS, though we note that this is sensitive to the period being analyzed. We further demonstrate that variability in coccolithophore chlorophyll a here is positively correlated with variability in nitrate and DIC (and especially the bicarbonate ion) in the upper 30 m of the water column. Previous studies have suggested that coccolithophore photosynthesis may benefit from increasing CO2, but calcification may eventually be hindered by low pHT (&lt; 7.7). Given that DIC has been increasing at BATS by ∼ 1.4 µmol kg−1 yr−1 over the period of 1991–2012, we speculate that coccolithophore photosynthesis and perhaps calcification may have increased in response to anthropogenic CO2 input.