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We described new sea surface CO2 observations in the south-western Indian Ocean obtained in January 2020 when a strong bloom event occurred south-east of Madagascar and extended eastward in the oligotrophic Indian Ocean subtropical domain. Compared to previous years (1991–2019) we observed very low fCO2 and dissolved inorganic carbon concentrations (CT) in austral summer 2020, indicative of a biologically driven process. In the bloom, the anomaly of fCO2 and CT reached respectively −33 µatm and −42 µmol kg−1, whereas no change is observed for alkalinity (AT). In January 2020 we estimated a local maximum of air–sea CO2 flux at 27∘ S of −6.9 mmol m−2 d−1 (ocean sink) and −4.3 mmol m−2 d−1 when averaging the flux in the band 26–30∘ S. In the domain 25–30∘ S, 50–60∘ E we estimated that the bloom led to a regional carbon uptake of about −1 TgC per month in January 2020, whereas this region was previously recognized as an ocean CO2 source or near equilibrium during this season. Using a neural network approach that reconstructs the monthly fCO2 fields, we estimated that when the bloom was at peak in December 2019 the CO2 sink reached −3.1 (±1.0) mmol m−2 d−1 in the band 25–30∘ S; i.e. the model captured the impact of the bloom. Integrated in the domain restricted to 25–30∘ S, 50–60∘ E, the region was a CO2 sink in December 2019 of −0.8 TgC per month compared to a CO2 source of +0.12 (±0.10) TgC per month in December when averaged over the period 1996–2018. Consequently in 2019 this region was a stronger CO2 annual sink of −8.8 TgC yr−1 compared to −7.0 (±0.5) TgC yr−1 averaged over 1996–2018. In austral summer 2019–2020, the bloom was likely controlled by a relatively deep mixed-layer depth during the preceding winter (July–September 2019) that would supply macro- and/or micro-nutrients such as iron to the surface layer to promote the bloom that started in November 2019 in two large rings in the Madagascar Basin. Based on measurements in January 2020, we observed relatively high N2 fixation rates (up to 18 nmol N L−1 d−1), suggesting that diazotrophs could play a role in the bloom in the nutrient-depleted waters. The bloom event in austral summer 2020, along with the new carbonate system observations, represents a benchmark case for complex biogeochemical model sensitivity studies (including the N2 fixation process and iron supplies) for a better understanding of the origin and termination of this still “mysterious” sporadic bloom and its impact on ocean carbon uptake in the future.

The oceanic phosphorus cycle describes how phosphorus moves through the ocean, accumulates with the sediments on the seafloor, and participates in biogeochemical reactions. We propose a new two-reservoir scenario of the glacial–interglacial phosphorus cycle. It relies on diagenesis in methane hydrate-bearing sediments to mobilize sedimentary phosphorus and transfer it to the oceanic reservoir during times when falling sea level lowers the hydrostatic pressure on the seafloor and destabilizes methane hydrates. The stock of solid phase phosphorus mobilizable by this process is of the same order of magnitude as the dissolved phosphate inventory of the current oceanic reservoir. The potential additional flux of phosphate during the glacial period is of the same order of magnitude as pre-agricultural, riverine dissolved phosphate fluxes to the ocean. Throughout the cycle, primary production assimilates phosphorus and inorganic carbon into biomass, which, upon settling and burial, returns phosphorus to the sedimentary reservoir. Primary production also lowers the partial pressure of CO2 in the surface ocean, potentially drawing down CO2 from the atmosphere. Concurrent with this slow “biological pump”, but operating in the opposite direction, a “physical pump” brings metabolic CO2-enriched waters from deep-ocean basins to the upper ocean. The two pumps compete, but the direction of the CO2 flux at the air–sea interface depends on the nutrient content of the deep waters. Because of the transfer of reactive phosphorus to the sedimentary reservoir throughout a glaciation cycle, low-phosphorus and high-CO2 deep waters reign at the beginning of a deglaciation, resulting in rapid transfer of CO2 to the atmosphere. The new scenario provides another element to the suite of processes that may have contributed to the rapid glacial–interglacial climate transitions documented in paleo-records.

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.

Continental shelves are thought to be affected disproportionately by climate change and are a large contributor to global air–sea carbon dioxide (CO2) fluxes. It is often reported that low-latitude shelves tend to act as net sources of CO2, whereas mid- and high-latitude shelves act as net sinks. Here, we combine a high-resolution regional model with surface water time series and repeat transect observations from the Scotian Shelf, a mid-latitude region in the northwest North Atlantic, to determine what processes are driving the temporal and spatial variability of partial pressure of CO2 (pCO2) on a seasonal scale. In contrast to the global trend, the Scotian Shelf acts as a net source. Surface pCO2 undergoes a strong seasonal cycle with an amplitude of ∼ 200–250 µatm. These changes are associated with both a strong biological drawdown of dissolved inorganic carbon (DIC) in spring (corresponding to a decrease in pCO2 of 100–200 µatm) and pronounced effects of temperature, which ranges from 0 ∘C in the winter to near 20 ∘C in the summer, resulting in an increase in pCO2 of ∼ 200–250 µatm. Throughout the summer, events with low surface water pCO2 occur associated with coastal upwelling. This effect of upwelling on pCO2 is also in contrast to the general assumption that upwelling increases surface pCO2 by delivering DIC-enriched water to the surface. Aside from these localized events, pCO2 is relatively uniform across the shelf. Our model agrees with regional observations, reproduces seasonal patterns of pCO2, and simulates annual outgassing of CO2 from the ocean of +1.7±0.2 mol C m−2 yr−1 for the Scotian Shelf, net uptake of CO2 by the ocean of -0.5±0.2 mol C m−2 yr−1 for the Gulf of Maine, and uptake by the ocean of -1.3±0.3 mol C m−2 yr−1 for the Grand Banks.

Exchange of dissolved substances at the sediment–water interface provides an important link between the short-term and long-term geochemical cycles in the ocean. A second, as yet poorly understood sediment–water exchange is supported by low-temperature circulation of seawater through the oceanic basement underneath the sediments. From the basement, upwards diffusing oxygen and other dissolved species modify the sediment, whereas reaction products diffuse from the sediment down into the basement where they are transported by the basement fluid and released to the ocean. Here, we investigate the impact of this “second” route with respect to transport, release and consumption of oxygen, nitrate, manganese, nickel and cobalt on the basis of sediment cores retrieved from the Clarion Clipperton Zone (CCZ) in the equatorial Pacific Ocean. We show that in this abyssal ocean region characterised by low organic carbon burial and sedimentation rates vast areas exist where the downward- and upward-directed diffusive fluxes of oxygen meet so that the sediments are oxic throughout. This is especially the case where sediments are thin or in the proximity of faults. Oxygen diffusing upward from the basaltic crust into the sediment contributes to the degradation of sedimentary organic matter. Where the sediments are entirely oxic, nitrate produced in the upper sediment by nitrification is lost both by upward diffusion into the bottom water and by downward diffusion into the fluids circulating within the basement. Where the oxygen profiles do not meet, they are separated by a suboxic sediment interval characterised by Mn2+ in the porewater. Where porewater Mn2+ in the suboxic zones remains low, nitrate consumption is low and the sediment continues to deliver nitrate to the ocean bottom waters and basement fluid. We observe that at elevated porewater manganese concentrations, nitrate consumption exceeds production and nitrate diffuses from the basement fluid into the sediment. Within the suboxic zone, not only manganese but also cobalt and nickel are released into the porewater by reduction of Mn oxides, diffusing towards the oxic–suboxic fronts above and below where they precipitate, effectively removing these metals from the suboxic zone and concentrating them at the two oxic–suboxic redox boundaries. We show that not only do diffusive fluxes in the top part of deep-sea sediments modify the geochemical composition over time but also diffusive fluxes of dissolved constituents from the basement into the bottom layers of the sediment. Hence, the palaeoceanographic interpretation of sedimentary layers should carefully consider such deep secondary modifications in order to prevent the misinterpretation of primary signatures.

The flow of Pacific water to the North Atlantic exerts a globally significant control on nutrient balances between the two ocean basins and strongly influences biological productivity in the northwest Atlantic. Nutrient ratios of nitrate (NO3-) versus phosphate (PO43-) have previously been used to complement salinity characteristics in tracing the distribution of Pacific water in the North Atlantic. We expand on this premise and demonstrate that the fraction of Pacific water as determined by NO3- : PO43- ratios can be quantitatively predicted from the isotopic composition of sub-euphotic nitrate in the northwest Atlantic. Our linear model thus provides a critically important framework for interpreting δ15N signatures incorporated into both modern marine biomass and organic material in historical and paleoceanographic archives along the northwest Atlantic margin.

The West Antarctic Ice Sheet (WAIS) is one of the largest potential sources of future sea-level rise, with glaciers draining the WAIS thinning at an accelerating rate over the past 40 years. Due to complexities in calibrating palaeoceanographic proxies for the Southern Ocean, it remains difficult to assess whether similar changes have occurred earlier during the Holocene or whether there is underlying centennial- to millennial-scale forcing in oceanic variability. Archaeal lipid-based proxies, specifically glycerol dialkyl glycerol tetraether (GDGT; e.g. TEX86 and TEX86L), are powerful tools for reconstructing ocean temperature, but these proxies have been shown previously to be difficult to apply to the Southern Ocean. A greater understanding of the parameters that control Southern Ocean GDGT distributions would improve the application of these biomarker proxies and thus help provide a longer-term perspective on ocean forcing of Antarctic ice sheet changes. In this study, we characterised intact polar lipid (IPL)-GDGTs, representing (recently) living archaeal populations in suspended particulate matter (SPM) from the Amundsen Sea and the Scotia Sea. SPM samples from the Amundsen Sea were collected from up to four water column depths representing the surface waters through to Circumpolar Deep Water (CDW), whereas the Scotia Sea samples were collected along a transect encompassing the sub-Antarctic front through to the southern boundary of the Antarctic Circumpolar Current. IPL-GDGTs with low cyclic diversity were detected throughout the water column with high relative abundances of hydroxylated IPL-GDGTs identified in both the Amundsen and Scotia seas. Results from the Scotia Sea show shifts in IPL-GDGT signatures across well-defined fronts of the Southern Ocean. Indicating that the physicochemical parameters of these water masses determine changes in IPL-GDGT distributions. The Amundsen Sea results identified GDGTs with hexose-phosphohexose head groups in the CDW, suggesting active GDGT synthesis at these depths. These results suggest that GDGTs synthesised at CDW depths may be a significant source of GDGTs exported to the sedimentary record and that temperature reconstructions based on TEX86 or TEX86L proxies may be significantly influenced by the warmer waters of the CDW.

A key challenge for biological oceanography is relating the physiological mechanisms controlling phytoplankton growth to the spatial distribution of those phytoplankton. Physiological mechanisms are often isolated by varying one driver of growth, such as nutrient or light, in a controlled laboratory setting producing what we call “intrinsic relationships”. We contrast these with the “apparent relationships” which emerge in the environment in climatological data. Although previous studies have found machine learning (ML) can find apparent relationships, there has yet to be a systematic study examining when and why these apparent relationships diverge from the underlying intrinsic relationships found in the lab and how and why this may depend on the method applied. Here we conduct a proof-of-concept study with three scenarios in which biomass is by construction a function of time-averaged phytoplankton growth rate. In the first scenario, the inputs and outputs of the intrinsic and apparent relationships vary over the same monthly timescales. In the second, the intrinsic relationships relate averages of drivers that vary on hourly timescales to biomass, but the apparent relationships are sought between monthly averages of these inputs and monthly-averaged output. In the third scenario we apply ML to the output of an actual Earth system model (ESM). Our results demonstrated that when intrinsic and apparent relationships operate on the same spatial and temporal timescale, neural network ensembles (NNEs) were able to extract the intrinsic relationships when only provided information about the apparent relationships, while colimitation and its inability to extrapolate resulted in random forests (RFs) diverging from the true response. When intrinsic and apparent relationships operated on different timescales (as little separation as hourly versus daily), NNEs fed with apparent relationships in time-averaged data produced responses with the right shape but underestimated the biomass. This was because when the intrinsic relationship was nonlinear, the response to a time-averaged input differed systematically from the time-averaged response. Although the limitations found by NNEs were overestimated, they were able to produce more realistic shapes of the actual relationships compared to multiple linear regression. Additionally, NNEs were able to model the interactions between predictors and their effects on biomass, allowing for a qualitative assessment of the colimitation patterns and the nutrient causing the most limitation. Future research may be able to use this type of analysis for observational datasets and other ESMs to identify apparent relationships between biogeochemical variables (rather than spatiotemporal distributions only) and identify interactions and colimitations without having to perform (or at least performing fewer) growth experiments in a lab. From our study, it appears that ML can extract useful information from ESM output and could likely do so for observational datasets as well.

We present a new natural carbon dioxide (CO 2 ) system located off the southern coast of the island of La Palma (Canary Islands, Spain). Like CO 2 seeps, these CO 2 submarine groundwater discharges (SGDs) can be used as an analogue to study the effects of ocean acidification (OA) on the marine realm. With this aim, we present the chemical characterization of the area, describing the carbon system dynamics, by measuring pH, AT and CT and calculating Ω aragonite and calcite. Our explorations of the area have found several emission points with similar chemical features. Here, the CT varies from 2120.10 to 10 784.84 µ mol kg −1 , AT from 2415.20 to 10 817.12 µ mol kg −1 , pH from 7.12 to 8.07, Ω aragonite from 0.71 to 4.15 and Ω calcite from 1.09 to 6.49 units. Also, the CO 2 emission flux varies between 2.8 and 28 kg CO 2 d −1 , becoming a significant source of carbon. These CO 2 emissions, which are of volcanic origin, acidify the brackish groundwater that is discharged to the coast and alter the local seawater chemistry. Although this kind of acidified system is not a perfect image of future oceans, this area of La Palma is an exceptional spot to perform studies aimed at understanding the effect of different levels of OA on the functioning of marine ecosystems. These studies can then be used to comprehend how life has persisted through past eras, with higher atmospheric CO 2 , or to predict the consequences of present fossil fuel usage on the marine ecosystem of the future oceans.

We developed a simple method to refine existing open-ocean maps and extend them towards different coastal seas. Using a multi-linear regression we produced monthly maps of surface ocean fCO2 in the northern European coastal seas (the North Sea, the Baltic Sea, the Norwegian Coast and the Barents Sea) covering a time period from 1998 to 2016. A comparison with gridded Surface Ocean CO2 Atlas (SOCAT) v5 data revealed mean biases and standard deviations of 0 ± 26 µatm in the North Sea, 0 ± 16 µatm along the Norwegian Coast, 0 ± 19 µatm in the Barents Sea and 2 ± 42 µatm in the Baltic Sea. We used these maps to investigate trends in fCO2, pH and air–sea CO2 flux. The surface ocean fCO2 trends are smaller than the atmospheric trend in most of the studied regions. The only exception to this is the western part of the North Sea, where sea surface fCO2 increases by 2 µatm yr−1, which is similar to the atmospheric trend. The Baltic Sea does not show a significant trend. Here, the variability was much larger than the expected trends. Consistently, the pH trends were smaller than expected for an increase in fCO2 in pace with the rise of atmospheric CO2 levels. The calculated air–sea CO2 fluxes revealed that most regions were net sinks for CO2. Only the southern North Sea and the Baltic Sea emitted CO2 to the atmosphere. Especially in the northern regions the sink strength increased during the studied period.

Oxygen-depleted regions of the global ocean are rapidly expanding, with important implications for global biogeochemical cycles. However, our ability to make projections about the future of oxygen in the ocean is limited by a lack of empirical data with which to test and constrain the behavior of global climatic and oceanographic models. We use depth-stratified plankton tows to demonstrate that some species of planktic foraminifera are adapted to life in the heart of the pelagic oxygen minimum zone (OMZ). In particular, we identify two species, Globorotaloides hexagonus and Hastigerina parapelagica, living within the eastern tropical North Pacific OMZ. The tests of the former are preserved in marine sediments and could be used to trace the extent and intensity of low-oxygen pelagic habitats in the fossil record. Additional morphometric analyses of G. hexagonus show that tests found in the lowest oxygen environments are larger, more porous, less dense, and have more chambers in the final whorl. The association of this species with the OMZ and the apparent plasticity of its test in response to ambient oxygenation invites the use of G. hexagonus tests in sediment cores as potential proxies for both the presence and intensity of overlying OMZs.

Typhoons are assumed to stimulate primary ocean production through the upward mixing of nutrients into the ocean surface. This assumption is based largely on observations of increased surface chlorophyll concentrations following the passage of typhoons. This surface chlorophyll enhancement, occasionally detected by satellites, is often undetected due to intense cloud coverage. Daily data from a BGC-Argo profiling float revealed the upper-ocean response to Typhoon Trami in the northwest Pacific Ocean. Temperature and chlorophyll changed rapidly, with a significant drop in sea surface temperature and a surge in surface chlorophyll associated with strong vertical mixing, which was only partially captured by satellite observations. However, no net increase in vertically integrated chlorophyll was observed during Typhoon Trami or in its wake. In contrast to the prevailing dogma, the result shows that typhoons likely have a limited effect on net primary ocean production. Observed surface chlorophyll enhancements during and immediately following typhoons in tropical and subtropical waters are more likely to be associated with surface entrainment of deep chlorophyll maxima. Moreover, the findings demonstrate that remote sensing data alone can overestimate the impact of storms on primary production in all oceans. Full understanding of the impact of storms on upper-ocean productivity can only be achieved with ocean-observing robots dedicated to high-resolution temporal sampling in the path of storms.

Marine calcification is an important component of the global carbon cycle. The mechanism by which some organisms take up inorganic carbon for the production of their shells or skeletons, however, remains only partly known. Although foraminifera are responsible for a large part of the global calcium carbonate production, the process by which they concentrate inorganic carbon is debated. Some evidence suggests that seawater is taken up by vacuolization and participates relatively unaltered in the process of calcification, whereas other results suggest the involvement of transmembrane transport and the activity of enzymes like carbonic anhydrase. Here, we tested whether inorganic-carbon uptake relies on the activity of carbonic anhydrase using incubation experiments with the perforate, large benthic, symbiont-bearing foraminifer Amphistegina lessonii. Calcification rates, determined by the alkalinity anomaly method, showed that inhibition of carbonic anhydrase by acetazolamide (AZ) stopped most of the calcification process. Inhibition of photosynthesis either by 3-(3,4-Dichlorophenyl)-1,1-dimethylurea (DCMU) or by incubating the foraminifera in the dark also decreased calcification rates but to a lesser degree than with AZ. Results from this study show that carbonic anhydrase plays a key role in biomineralization of Amphistegina lessonii and indicates that calcification of those perforate, large benthic foraminifera might, to a certain extent, benefit from the extra dissolved inorganic carbon (DIC), which causes ocean acidification.

The shallower oxygen-poor water masses of the ocean confine a majority of the microbial communities that can produce up to 90 % of oceanic N2. This effective N2-yielding section encloses a suspended small-particle layer, inferred from particle backscattering (bbp) measurements. It is thus hypothesized that this layer (hereafter, the bbp-layer) is linked to microbial communities involved in N2 yielding such as nitrate-reducing SAR11 as well as sulfur-oxidizing, anammox, and denitrifying bacteria – a hypothesis yet to be evaluated. Here, data collected by three BGC-Argo floats deployed in the Black Sea are used to investigate the origin of this bbp-layer. To this end, we evaluate how the key drivers of N2-yielding bacteria dynamics impact the vertical distribution of bbp and the thickness of the bbp-layer. In conjunction with published data on N2 excess, our results suggest that the bbp-layer is at least partially composed of the bacteria driving N2 yielding for three main reasons: (1) strong correlations are recorded between bbp and nitrate; (2) the top location of the bbp-layer is driven by the ventilation of oxygen-rich subsurface waters, while its thickness is modulated by the amount of nitrate available to produce N2; and (3) the maxima of both bbp and N2 excess coincide at the same isopycnals where bacteria involved in N2 yielding coexist. We thus advance that bbp and O2 can be exploited as a combined proxy to delineate the N2-yielding section of the Black Sea. This proxy can potentially contribute to refining delineation of the effective N2-yielding section of oxygen-deficient zones via data from the growing BGC-Argo float network.

Cold-water corals (CWCs) constitute important deep-water ecosystems that are under increasing environmental pressure due to ocean acidification and global warming. The sensitivity of these deep-water ecosystems to environmental change is demonstrated by abundant paleorecords drilled through CWC mounds that reveal characteristic alterations between rapid formation and dormant or erosive phases. Previous studies have identified several central parameters for driving or inhibiting CWC growth such as food supply, oxygenation, and the carbon saturation state of bottom water, yet there are still large uncertainties about the relative importance of the different environmental parameters. To advance this debate we have performed a multiproxy study on a sediment core retrieved from the 25 m high Bowie Mound, located at 866 m water depth on the continental slope off southeastern Brazil, a structure built up mainly by the CWC Solenosmilia variabilis. Our results indicate a multifactorial control on CWC growth at Bowie Mound during the past ∼ 160 kyr, which reveals distinct formation pulses during northern high-latitude glacial cold events (Heinrich stadials, HSs) largely associated with anomalously strong monsoonal rainfall over the continent. The ensuing enhanced runoff elevated the terrigenous nutrient and organic-matter supply to the continental margin and likely boosted marine productivity. The dispersal of food particles towards the CWC colonies during HSs was facilitated by the highly dynamic hydraulic conditions along the continental slope that prevailed throughout glacial periods. These conditions caused the emplacement of a pronounced nepheloid layer above Bowie Mound, thereby aiding the concentration and along-slope dispersal of organic matter. Our study thus emphasizes the impact of continental climate variability on a highly vulnerable deep-marine ecosystem. ispartof: Biogeosciences vol:17 issue:23 pages:5883-5908 status: Published online

A three-dimensional physical-biological model of the marginal seas of China was used to analyze interannual and intra-seasonal variations in hypoxic conditions and identify the main processes controlling their generation off the Changjiang (or Yangtze River) estuary. The model was compared against available observations and reproduces the observed temporal and spatial variability of physical and biological properties including bottom oxygen. Interannual variations of hypoxic extent in the simulation are partly explained by variations in river discharge but not nutrient load. As riverine inputs of freshwater and nutrients are consistently high, promoting large productivity and subsequent oxygen consumption in the region affected by the river plume, wind forcing is important in modulating interannual and short-term variability. Wind direction is relevant because it determines the spatial extent and distribution of the freshwater plume, which is strongly affected by either upwelling or downwelling conditions. High-wind events can lead to partial reoxygenation of bottom waters and, when occurring in succession throughout the hypoxic season, can effectively suppress the development of hypoxic conditions, thus influencing interannual variability. A model-derived oxygen budget is presented and suggests that sediment oxygen consumption is the dominant oxygen sink below the pycnocline and that advection of oxygen in the bottom waters acts as an oxygen sink in spring but becomes a source during hypoxic conditions in summer, especially in the southern part of the hypoxic region, which is influenced by open-ocean intrusions.

Cobalt (Co) is an important bioactive trace metal that is the metal cofactor in cobalamin (vitamin B12) which can limit or co-limit phytoplankton growth in many regions of the ocean. Total dissolved and labile Co measurements in the Canadian sector of the Arctic Ocean during the U.S. GEOTRACES Arctic expedition (GN01) and the Canadian International Polar Year GEOTRACES expedition (GIPY14) revealed a dynamic biogeochemical cycle for Co in this basin. The major sources of Co in the Arctic were from shelf regions and rivers, with only minimal contributions from other freshwater sources (sea ice, snow) and eolian deposition. The most striking feature was the extremely high concentrations of dissolved Co in the upper 100 m, with concentrations routinely exceeding 800 pmol L−1 over the shelf regions. This plume of high Co persisted throughout the Arctic basin and extended to the North Pole, where sources of Co shifted from primarily shelf-derived to riverine, as freshwater from Arctic rivers was entrained in the Transpolar Drift. Dissolved Co was also strongly organically complexed in the Arctic, ranging from 70 % to 100 % complexed in the surface and deep ocean, respectively. Deep-water concentrations of dissolved Co were remarkably consistent throughout the basin (∼55 pmol L−1), with concentrations reflecting those of deep Atlantic water and deep-ocean scavenging of dissolved Co. A biogeochemical model of Co cycling was used to support the hypothesis that the majority of the high surface Co in the Arctic was emanating from the shelf. The model showed that the high concentrations of Co observed were due to the large shelf area of the Arctic, as well as to dampened scavenging of Co by manganese-oxidizing (Mn-oxidizing) bacteria due to the lower temperatures. The majority of this scavenging appears to have occurred in the upper 200 m, with minimal additional scavenging below this depth. Evidence suggests that both dissolved Co (dCo) and labile Co (LCo) are increasing over time on the Arctic shelf, and these limited temporal results are consistent with other tracers in the Arctic. These elevated surface concentrations of Co likely lead to a net flux of Co out of the Arctic, with implications for downstream biological uptake of Co in the North Atlantic and elevated Co in North Atlantic Deep Water. Understanding the current distributions of Co in the Arctic will be important for constraining changes to Co inputs resulting from regional intensification of freshwater fluxes from ice and permafrost melt in response to ongoing climate change.

A massive mangrove dieback event occurred in 2015–2016 along ∼1000 km of pristine coastline in the Gulf of Carpentaria, Australia. Here, we use sediment and wood chronologies to gain insights into geochemical and climatic changes related to this dieback. The unique combination of low rainfall and low sea level observed during the dieback event had been unprecedented in the preceding 3 decades. A combination of iron (Fe) chronologies in wood and sediment, wood density and estimates of mangrove water use efficiency all imply lower water availability within the dead mangrove forest. Wood and sediment chronologies suggest a rapid, large mobilization of sedimentary Fe, which is consistent with redox transitions promoted by changes in soil moisture content. Elemental analysis of wood cross sections revealed a 30- to 90-fold increase in Fe concentrations in dead mangroves just prior to their mortality. Mangrove wood uptake of Fe during the dieback is consistent with large apparent losses of Fe from sediments, which potentially caused an outwelling of Fe to the ocean. Although Fe toxicity may also have played a role in the dieback, this possibility requires further study. We suggest that differences in wood and sedimentary Fe between living and dead forest areas reflect sediment redox transitions that are, in turn, associated with regional variability in groundwater flows. Overall, our observations provide multiple lines of evidence that the forest dieback was driven by low water availability coinciding with a strong El Niño–Southern Oscillation (ENSO) event and was associated with climate change.

Sediment–water oxygen fluxes are widely used as a proxy for organic carbon production and mineralization at the seafloor. In situ fluxes can be measured non-invasively with the aquatic eddy covariance technique, but a critical requirement is that the sensors of the instrument are able to correctly capture the high-frequency variations in dissolved oxygen concentration and vertical velocity. Even small changes in sensor characteristics during deployment as caused, e.g. by biofouling can result in erroneous flux data. Here we present a dual-optode eddy covariance instrument (2OEC) with two fast oxygen fibre sensors and document how erroneous flux interpretations and data loss can effectively be reduced by this hardware and a new data analysis approach. With deployments over a carbonate sandy sediment in the Florida Keys and comparison with parallel benthic advection chamber incubations, we demonstrate the improved data quality and data reliability facilitated by the instrument and associated data processing. Short-term changes in flux that are dubious in measurements with single oxygen sensor instruments can be confirmed or rejected with the 2OEC and in our deployments provided new insights into the temporal dynamics of benthic oxygen flux in permeable carbonate sands. Under steady conditions, representative benthic flux data can be generated with the 2OEC within a couple of hours, making this technique suitable for mapping sediment–water, intra-water column, or atmosphere–water fluxes.

Light-use efficiency defines the ability of primary producers to convert sunlight energy to primary production and is computed as the ratio between the gross primary production and the intercepted photosynthetic active radiation. While this measure has been applied broadly within terrestrial ecology to investigate habitat resource-use efficiency, it remains underused within the aquatic realm. This report provides a conceptual framework to compute hourly and daily light-use efficiency using underwater O2 eddy covariance, a recent technological development that produces habitat-scale rates of primary production under unaltered in situ conditions. The analysis, tested on two benthic flux datasets, documents that hourly light-use efficiency may approach the theoretical limit of 0.125 O2 per photon under low-light conditions, but it decreases rapidly towards the middle of the day and is typically 10-fold lower on a 24 h basis. Overall, light-use efficiency provides a useful measure of habitat functioning and facilitates site comparison in time and space.

As the primary energy and carbon source in aquatic food webs, phytoplankton generally display spatial heterogeneity due to complicated biotic and abiotic controls; however our understanding of the causes of this spatial heterogeneity is challenging, as it involves multiple regulatory mechanisms. We applied a combination of field observation, numerical modeling, and remote sensing to display and interpret the spatial gradient of phytoplankton biomass in a Dutch tidal bay (the Eastern Scheldt) on the east coast of the North Sea. The 19 years (1995–2013) of monitoring data reveal a seaward increasing trend in chlorophyll-a (chl a) concentrations during the spring bloom. Using a calibrated and validated three-dimensional hydrodynamic–biogeochemical model, two idealized model scenarios were run: switching off the suspension feeders and halving the open-boundary nutrient and phytoplankton loading. Results reveal that bivalve grazing exerts a dominant control on phytoplankton in the bay and that the tidal import mainly influences algal biomass near the mouth. Satellite data captured a post-bloom snapshot that indicated the temporally variable phytoplankton distribution. Based on a literature review, we found five common spatial phytoplankton patterns in global estuarine–coastal ecosystems for comparison with the Eastern Scheldt case: seaward increasing, seaward decreasing, concave with a chlorophyll maximum, weak spatial gradients, and irregular patterns. We highlight the temporal variability of these spatial patterns and the importance of anthropogenic and environmental influences.

The coastal ocean is strongly affected by ocean acidification because of its shallow water depths, low volume, and the closeness to terrestrial dynamics. Earlier observations of dissolved inorganic carbon (DIC) and total alkalinity (TA) in the southern part of the North Sea, a northwest European shelf sea, revealed lower acidification effects than expected. It has been assumed that anaerobic degradation and subsequent TA release in the adjacent back-barrier tidal areas (Wadden Sea) in summertime is responsible for this phenomenon. In this study the exchange rates of TA and DIC between the Wadden Sea tidal basins and the North Sea and the consequences for the carbonate system in the German Bight are estimated using a 3D ecosystem model. The aim of this study is to differentiate the various sources contributing to observed high summer TA in the southern North Sea. Measured TA and DIC in the Wadden Sea are considered as model boundary conditions. This procedure acknowledges the dynamic behaviour of the Wadden Sea as an area of effective production and decomposition of organic material. According to the modelling results, 39 Gmol TA yr−1 were exported from the Wadden Sea into the North Sea, which is less than a previous estimate but within a comparable range. The interannual variabilities in TA and DIC, mainly driven by hydrodynamic conditions, were examined for the years 2001–2009. Dynamics in the carbonate system are found to be related to specific weather conditions. The results suggest that the Wadden Sea is an important driver for the carbonate system in the southern North Sea. On average 41 % of TA inventory changes in the German Bight were caused by riverine input, 37 % by net transport from adjacent North Sea sectors, 16 % by Wadden Sea export, and 6 % were caused by internal net production of TA. The dominant role of river input for the TA inventory disappears when focusing on TA concentration changes due to the corresponding freshwater fluxes diluting the marine TA concentrations. The ratio of exported TA versus DIC reflects the dominant underlying biogeochemical processes in the Wadden Sea. Whereas aerobic degradation of organic matter played a key role in the North Frisian Wadden Sea during all seasons of the year, anaerobic degradation of organic matter dominated in the East Frisian Wadden Sea. Despite the scarcity of high-resolution field data, it is shown that anaerobic degradation in the Wadden Sea is one of the main contributors of elevated summer TA values in the southern North Sea.

Boron isotope systematics of planktonic foraminifera from core-top sediments and culture experiments have been studied to investigate the sensitivity of δ11B of calcite tests to seawater pH. However, our knowledge of the relationship between δ11B and pH remains incomplete for many taxa. Thus, to expand the potential scope of application of this proxy, we report δ11B data for seven different species of planktonic foraminifera from sediment core tops. We utilize a method for the measurement of small samples of foraminifera and calculate the δ11B -calcite sensitivity to pH for Globigerinoides ruber, Trilobus sacculifer (sacc or without sacc), Orbulina universa, Pulleniatina obliquiloculata, Neogloboquadrina dutertrei, Globorotalia menardii, and Globorotalia tumida, including for unstudied core tops and species. These taxa have diverse ecological preferences and are from sites that span a range of oceanographic regimes, including some that are in regions of air–sea equilibrium and others that are out of equilibrium with the atmosphere. The sensitivity of δ11B carbonate to δ11B borate (e.g., Δδ11Bcarbonate∕Δδ11Bborate ) in core tops is consistent with previous studies for T. sacculifer and G. ruber and close to unity for N. dutertrei, O. universa, and combined deep-dwelling species. Deep-dwelling species closely follow the core-top calibration for O. universa, which is attributed to respiration-driven microenvironments likely caused by light limitation and/or symbiont–host interactions. Our data support the premise that utilizing boron isotope measurements of multiple species within a sediment core can be utilized to constrain vertical profiles of pH and pCO2 at sites spanning different oceanic regimes, thereby constraining changes in vertical pH gradients and yielding insights into the past behavior of the oceanic carbon pumps.

The coastal ecosystem of the Gulf of Alaska (GOA) is especially vulnerable to the effects of ocean acidification and climate change. Detection of these long-term trends requires a good understanding of the system’s natural state. The GOA is a highly dynamic system that exhibits large inorganic carbon variability on subseasonal to interannual timescales. This variability is poorly understood due to the lack of observations in this expansive and remote region. We developed a new model setup for the GOA that couples the three-dimensional Regional Oceanic Model System (ROMS) and the Carbon, Ocean Biogeochemistry and Lower Trophic (COBALT) ecosystem model. To improve our conceptual understanding of the system, we conducted a hindcast simulation from 1980 to 2013. The model was explicitly forced with temporally and spatially varying coastal freshwater discharges from a high-resolution terrestrial hydrological model, thereby affecting salinity, alkalinity, dissolved inorganic carbon, and nutrient concentrations. This represents a substantial improvement over previous GOA modeling attempts. Here, we evaluate the model on seasonal to interannual timescales using the best available inorganic carbon observations. The model was particularly successful in reproducing observed aragonite oversaturation and undersaturation of near-bottom water in May and September, respectively. The largest deficiency in the model is its inability to adequately simulate springtime surface inorganic carbon chemistry, as it overestimates surface dissolved inorganic carbon, which translates into an underestimation of the surface aragonite saturation state at this time. We also use the model to describe the seasonal cycle and drivers of inorganic carbon parameters along the Seward Line transect in under-sampled months. Model output suggests that the majority of the near-bottom water along the Seward Line is seasonally undersaturated with respect to aragonite between June and January, as a result of upwelling and remineralization. Such an extensive period of reoccurring aragonite undersaturation may be harmful to ocean acidification-sensitive organisms. Furthermore, the influence of freshwater not only decreases the aragonite saturation state in coastal surface waters in summer and fall, but it simultaneously decreases the surface partial pressure of carbon dioxide (pCO2), thereby decoupling the aragonite saturation state from pCO2. The full seasonal cycle and geographic extent of the GOA region is under-sampled, and our model results give new and important insights for months of the year and areas that lack in situ inorganic carbon observations.

Sediments in oxygen-depleted marine environments can be an important sink or source of bio-essential trace metals in the ocean. However, the key mechanisms controlling the release from or burial of trace metals in sediments are not exactly understood. Here, we investigate the benthic biogeochemical cycling of iron (Fe) and cadmium (Cd) in the oxygen minimum zone off Peru. We combine bottom water and pore water concentrations, as well as benthic fluxes determined from pore water profiles and from in situ benthic chamber incubations, along a depth transect at 12∘ S. In agreement with previous studies, both concentration–depth profiles and in situ benthic fluxes indicate a release of Fe from sediments to the bottom water. Diffusive Fe fluxes and Fe fluxes from benthic chamber incubations (−0.3 to −17.5 mmol m−2 yr−1) are broadly consistent at stations within the oxygen minimum zone, where the flux magnitude is highest, indicating that diffusion is the main transport mechanism of dissolved Fe across the sediment–water interface. The occurrence of mats of sulfur-oxidizing bacteria on the seafloor represents an important control on the spatial distribution of Fe fluxes by regulating hydrogen sulfide (H2S) concentrations and, potentially, Fe sulfide precipitation within the surface sediment. Rapid removal of dissolved Fe after its release to anoxic bottom waters hints at oxidative removal by nitrite and interactions with particles in the near-bottom water column. Benthic flux estimates of Cd suggest a flux into the sediment within the oxygen minimum zone. Fluxes from benthic chamber incubations (up to 22.6 µmol m−2 yr−1) exceed diffusive fluxes ( µmol m−2 yr−1) by a factor of more than 25, indicating that downward diffusion of Cd across the sediment–water interface is of subordinate importance for Cd removal from benthic chambers. As Cd removal in benthic chambers covaries with H2S concentrations in the pore water of surface sediments, we argue that Cd removal is mediated by precipitation of cadmium sulfide (CdS) within the chamber water or directly at the sediment–water interface. A mass balance approach, taking the contributions of diffusive and chamber fluxes as well as Cd delivery with organic material into account, suggests that CdS precipitation in the near-bottom water could make an important contribution to the overall Cd mass accumulation in the sediment solid phase. According to our results, the solubility of trace metal sulfide minerals (Cd ≪ Fe) is a key factor controlling trace metal removal and, consequently, the magnitude and the temporal and spatial heterogeneity of sedimentary fluxes. We argue that, depending on their sulfide solubility, sedimentary source or sink fluxes of trace metals will change differentially as a result of declining oxygen concentrations and the associated expansion of sulfidic surface sediments. Such a trend could cause a change in the trace metal stoichiometry of upwelling water masses with potential consequences for marine ecosystems in the surface ocean.

Around thirty species of marine bacteria can emit light, a critical characteristic in the oceanic environment where the major part is deprived of sunlight. In this article, we first review current knowledge on bioluminescent bacteria symbiosis in light organs. Then, focusing on gut-associated bacteria, we highlight that recent works, based on omics methods, confirm previous claims about the prominence of bioluminescent bacterial species in fish guts. Such host-symbiont relationships are relatively well established and represent important knowledge in the bioluminescence field. However, the consequences of bioluminescent bacteria continuously released from light organ and through the digestive tracts to the seawater have been barely taken into account at the ecological and biogeochemical level. For too long neglected, we propose to consider the role of bioluminescent bacteria, and to reconsider the biological carbon pump taking into account the bioluminescence effect (bioluminescence shunt hypothesis). Indeed, it has been shown that marine snow and fecal pellets are often luminous due to microbial colonization, which makes them a visual target. These luminous particles seem preferentially consumed by organisms of higher trophic levels in comparison to non-luminous ones. As a consequence, the sinking rate of consumed particles could be either increased (due to repackaging) or reduced (due to sloppy feeding or coprophagy/coprorhexy) which can imply a major impact on global biological carbon fluxes. Finally, we propose a strategy, at a worldwide scale, relying on recently developed instrumentation and methodological tools to quantify the impact of bioluminescent bacteria in the biological carbon pump.

The dynamics of the physicochemical and biological parameters were followed during the decline of a Cymodocea nodosa meadow in the northern Adriatic Sea from July 2017 to October 2018. During the regular growth of C. nodosa from July 2017 to March 2018, the species successfully adapted to the changes in environmental conditions and prevented H2S accumulation by its reoxidation, supplying the sediment with O2 from the water column and/or leaf photosynthesis. The C. nodosa decline was most likely triggered in April 2018 when light availability to the plant was drastically reduced due to increased seawater turbidity that resulted from increased terrigenous input, indicated by a decrease in salinity accompanied with a substantial increase in particulate matter concentration, combined with resuspension of sediment and elevated autotrophic biomass. Light reduction impaired photosynthesis of C. nodosa and the oxidation capability of belowground tissue. Simultaneously, a depletion of oxygen due to intense oxidation of H2S occurred in the sediment, thus creating anoxic conditions in most of the rooted areas. These linked negative effects on the plant performance caused an accumulation of H2S in the sediments of the C. nodosa meadow. During the decay of aboveground and belowground tissues, culminating in August 2018, high concentrations of H2S were reached and accumulated in the sediment as well as in bottom waters. The influx of oxygenated waters in September 2018 led to the re-establishment of H2S oxidation in the sediment and remainder of the belowground tissue. Our results indicate that if disturbances of environmental conditions, particularly those compromising the light availability, take place during the recruitment phase of plant growth when metabolic needs are at a maximum and stored reserves minimal, a sudden and drastic decline of the seagrass meadow occurs.

The Campanian age (Late Cretaceous) is characterized by a warm greenhouse climate with limited land-ice volume. This makes this period an ideal target for studying climate dynamics during greenhouse periods, which are essential for predictions of future climate change due to anthropogenic greenhouse gas emissions. Well-preserved fossil shells from the Campanian (±78 Ma) high mid-latitude (50ĝˆ  N) coastal faunas of the Kristianstad Basin (southern Sweden) offer a unique snapshot of short-term climate and environmental variability, which complements existing long-term climate reconstructions. In this study, we apply a combination of high-resolution spatially resolved trace element analyses (micro-X-ray fluorescence - μXRF - and laser ablation inductively coupled plasma mass spectrometry - LA-ICP-MS), stable isotope analyses (IRMS) and growth modeling to study short-term (seasonal) variations recorded in the oyster species Rastellum diluvianum from the Ivö Klack locality. Geochemical records through 12 specimens shed light on the influence of specimen-specific and ontogenetic effects on the expression of seasonal variations in shell chemistry and allow disentangling vital effects from environmental influences in an effort to refine paleoseasonality reconstructions of Late Cretaceous greenhouse climates. Growth models based on stable oxygen isotope records yield information on the mode of life, circadian rhythm and reproductive cycle of these extinct oysters. This multi-proxy study reveals that mean annual temperatures in the Campanian higher mid-latitudes were 17 to 19 ĝˆ C, with winter minima of ĝˆ1/413 ĝˆ C and summer maxima of 26 ĝˆ C, assuming a Late Cretaceous seawater oxygen isotope composition of -1 ‰ VSMOW (Vienna standard mean ocean water). These results yield smaller latitudinal differences in temperature seasonality in the Campanian compared to today. Latitudinal temperature gradients were similar to the present, contrasting with previous notions of "equable climate" during the Late Cretaceous. Our results also demonstrate that species-specific differences and uncertainties in the composition of Late Cretaceous seawater prevent trace element proxies (Mgĝˆ•Ca, Srĝˆ•Ca, Mgĝˆ•Li and Srĝˆ•Li) from being used as reliable temperature proxies for fossil oyster shells. However, trace element profiles can serve as a quick tool for diagenesis screening and investigating seasonal growth patterns in ancient shells.

Dimethyl sulfide and volatile organic compounds (VOCs) are important for atmospheric chemistry. The emissions of biogenically derived organic gases, including dimethyl sulfide and especially isoprene, are not well constrained in the Southern Ocean. Due to a paucity of measurements, the role of the ocean in the atmospheric budgets of atmospheric methanol, acetone, and acetaldehyde is even more poorly known. In order to quantify the air–sea fluxes of these gases, we measured their seawater concentrations and air mixing ratios in the Atlantic sector of the Southern Ocean, along a ∼ 11 000 km long transect at approximately 60∘ S in February–April 2019. Concentrations, oceanic saturations, and estimated fluxes of five simultaneously sampled gases (dimethyl sulfide, isoprene, methanol, acetone, and acetaldehyde) are presented here. Campaign mean (±1σ) surface water concentrations of dimethyl sulfide, isoprene, methanol, acetone, and acetaldehyde were 2.60 (±3.94), 0.0133 (±0.0063), 67 (±35), 5.5 (±2.5), and 2.6 (±2.7) nmol dm−3 respectively. In this dataset, seawater isoprene and methanol concentrations correlated positively. Furthermore, seawater acetone, methanol, and isoprene concentrations were found to correlate negatively with the fugacity of carbon dioxide, possibly due to a common biological origin. Campaign mean (±1σ) air mixing ratios of dimethyl sulfide, isoprene, methanol, acetone, and acetaldehyde were 0.17 (±0.09), 0.053 (±0.034), 0.17 (±0.08), 0.081 (±0.031), and 0.049 (±0.040) ppbv. We observed diel changes in averaged acetaldehyde concentrations in seawater and ambient air (and to a lesser degree also for acetone and isoprene), which suggest light-driven production. Campaign mean (±1σ) fluxes of 4.3 (±7.4) µmol m−2 d−1 DMS and 0.028 (±0.021) µmol m−2 d−1 isoprene are determined where a positive flux indicates from the ocean to the atmosphere. Methanol was largely undersaturated in the surface ocean with a mean (±1σ) net flux of −2.4 (±4.7) µmol m−2 d−1, but it also had a few occasional episodes of outgassing. This section of the Southern Ocean was found to be a source and a sink for acetone and acetaldehyde this time of the year, depending on location, resulting in a mean net flux of −0.55 (±1.14) µmol m−2 d−1 for acetone and −0.28 (±1.22) µmol m−2 d−1 for acetaldehyde. The data collected here will be important for constraining the air–sea exchange, cycling, and atmospheric impact of these gases, especially over the Southern Ocean.

Trichodesmium is a globally important marine microbe that provides fixed nitrogen (N) to otherwise N-limited ecosystems. In nature, nitrogen fixation is likely regulated by iron or phosphate availability, but the extent and interaction of these controls are unclear. From metaproteomics analyses using established protein biomarkers for nutrient stress, we found that iron–phosphate co-stress is the norm rather than the exception for Trichodesmium colonies in the North Atlantic Ocean. Counterintuitively, the nitrogenase enzyme was more abundant under co-stress as opposed to single nutrient stress. This is consistent with the idea that Trichodesmium has a specific physiological state during nutrient co-stress. Organic nitrogen uptake was observed and occurred simultaneously with nitrogen fixation. The quantification of the phosphate ABC transporter PstA combined with a cellular model of nutrient uptake suggested that Trichodesmium is generally confronted by the biophysical limits of membrane space and diffusion rates for iron and phosphate acquisition in the field. Colony formation may benefit nutrient acquisition from particulate and organic sources, alleviating these pressures. The results highlight that to predict the behavior of Trichodesmium, both Fe and P stress must be evaluated simultaneously.

Coastal systems can act as filters for anthropogenic nutrient input into marine environments. Here, we assess the processes controlling the removal of phosphorus (P) and nitrogen (N) for four sites in the eutrophic Stockholm archipelago. Bottom water concentrations of oxygen (O2) and P are inversely correlated. This is attributed to the seasonal release of P from iron-oxide-bound (Fe-oxide-bound) P in surface sediments and from degrading organic matter. The abundant presence of sulfide in the pore water and its high upward flux towards the sediment surface (∼4 to 8 mmol m−2 d−1), linked to prior deposition of organic-rich sediments in a low-O2 setting (“legacy of hypoxia”), hinder the formation of a larger Fe-oxide-bound P pool in winter. This is most pronounced at sites where water column mixing is naturally relatively low and where low bottom water O2 concentrations prevail in summer. Burial rates of P are high at all sites (0.03–0.3 mol m−2 yr−1), a combined result of high sedimentation rates (0.5 to 3.5 cm yr−1) and high sedimentary P at depth (∼30 to 50 µmol g−1). Sedimentary P is dominated by Fe-bound P and organic P at the sediment surface and by organic P, authigenic Ca-P and detrital P at depth. Apart from one site in the inner archipelago, where a vivianite-type Fe(II)-P mineral is likely present at depth, there is little evidence for sink switching of organic or Fe-oxide-bound P to authigenic P minerals. Denitrification is the major benthic nitrate-reducing process at all sites (0.09 to 1.7 mmol m−2 d−1) with rates decreasing seaward from the inner to outer archipelago. Our results explain how sediments in this eutrophic coastal system can remove P through burial at a relatively high rate, regardless of whether the bottom waters are oxic or (frequently) hypoxic. Our results suggest that benthic N processes undergo annual cycles of removal and recycling in response to hypoxic conditions. Further nutrient load reductions are expected to contribute to the recovery of the eutrophic Stockholm archipelago from hypoxia. Based on the dominant pathways of P and N removal identified in this study, it is expected that the sediments will continue to remove part of the P and N loads.

The largest and commercially appealing mineral deposits can be found in the abyssal sea floor of the Clarion-Clipperton Zone (CCZ), a polymetallic nodule province, in the NE Pacific Ocean, where experimental mining is due to take place. In anticipation of deep-sea mining impacts, it has become essential to rapidly and accurately assess biodiversity. For this reason, ophiuroid material collected during eight scientific cruises from five exploration licence areas within CCZ, one area being protected from mining (APEI3, Area of Particular Environmental Interest) in the periphery of CCZ and the DISturbance and re-COLonisation (DISCOL) Experimental Area (DEA), in the SE Pacific Ocean, was examined. Specimens were genetically analysed using a fragment of the mitochondrial cytochrome c oxidase subunit I (COI). Maximum-likelihood and neighbour-joining trees were constructed, while four tree-based and distance-based methods of species delineation (automatic barcode gap discovery, ABGD; barcode index numbers, BINs; general mixed Yule–coalescent, GMYC; multi-rate Poisson tree process, mPTP) were employed to propose secondary species hypotheses (SSHs) within the ophiuroids collected. The species delimitation analyses' concordant results revealed the presence of 43 deep-sea brittle star SSHs, revealing an unexpectedly high diversity and showing that the most conspicuous invertebrates in abyssal plains have been so far considerably underestimated. The number of SSHs found in each area varied from five (IFREMER area) to 24 (BGR (Federal Institute for Geosciences and Natural Resources, Germany) area) while 13 SSHs were represented by singletons. None of the SSHs were found to be present in all seven areas while the majority of species (44.2 %) had a single-area presence (19 SSHs). The most common species were Ophioleucidae sp. (Species 29), Amphioplus daleus (Species 2) and Ophiosphalma glabrum (Species 3), present in all areas except APEI3. The biodiversity patterns could be mainly attributed to particulate organic carbon (POC) fluxes that could explain the highest species numbers found in BGR (German contractor area) and UKSRL (UK Seabed Resources Ltd, UK contractor area) areas. The five exploration contract areas belong to a mesotrophic province, while conversely the APEI3 is located in an oligotrophic province, which could explain the lowest diversity as well as very low similarity with the other six study areas. Based on these results the representativeness and the appropriateness of APEI3 to meet its purpose of preserving the biodiversity of the CCZ fauna are questioned. Finally, this study provides the foundation for biogeographic and functional analyses that will provide insight into the drivers of species diversity and its role in ecosystem function.

Arctic sea ice is retreating and thinning and its rate of decline has steepened in the last decades. While phytoplankton blooms are known to seasonally propagate along the ice edge as it recedes from spring to summer, the substitution of thick multiyear ice (MYI) with thinner, ponded first-year ice (FYI) represents an unequal exchange when considering the roles sea ice plays in the ecology and climate of the Arctic. Consequences of this shifting sea ice on the phenology of phytoplankton and the associated cycling of the climate-relevant gas dimethylsulfide (DMS) and its precursor dimethylsulfoniopropionate (DMSP) remain ill constrained. In July–August 2014, two contrasting ice edges in the Canadian High Arctic were explored: a FYI-dominated ice edge in Barrow Strait and a MYI-dominated ice edge in Nares Strait. Our results reveal two distinct planktonic systems and associated DMS dynamics in connection to these diverging ice types. The surface waters exiting the ponded FYI in Barrow Strait were characterized by moderate chlorophyll a (Chl a,  µg L−1) as well as high DMSP (115 nmol L−1) and DMS (12 nmol L−1), suggesting that a bloom had already started to develop under the markedly melt-pond-covered (ca. 40 %) FYI. Heightened DMS concentrations at the FYI edge were strongly related to ice-associated seeding of DMS in surface waters and haline-driven stratification linked to ice melt (Spearman's rank correlation between DMS and salinity, rs=-0.91, p, n=20). However, surface waters exiting the MYI edge at the head of Nares Strait were characterized by low concentrations of Chl a ( µg L−1), DMSP ( nmol L−1), and DMS ( nmol L−1), despite the nutrient-replete conditions characterizing the surface waters. The increase in autotrophic biomass and methylated sulfur compounds took place several kilometers (ca. 100 km) away from the MYI edge, suggesting the requisite for ice-free, light-sufficient conditions for a phytoplankton bloom to fully develop and for sulfur compound dynamics to follow and expand. In light of the ongoing and projected climate-driven changes to Arctic sea ice, results from this study suggest that the early onset of autotrophic blooms under thinner, melt-pond-covered ice may have vast implications for the timing and magnitude of DMS pulses in the Arctic.

The extracellular concentration of H2O2 in surface aquatic environments is controlled by a balance between photochemical production and the microbial synthesis of catalase and peroxidase enzymes to remove H2O2 from solution. In any kind of incubation experiment, the formation rates and equilibrium concentrations of reactive oxygen species (ROSs) such as H2O2 may be sensitive to both the experiment design, particularly to the regulation of incident light, and the abundance of different microbial groups, as both cellular H2O2 production and catalase–peroxidase enzyme production rates differ between species. Whilst there are extensive measurements of photochemical H2O2 formation rates and the distribution of H2O2 in the marine environment, it is poorly constrained how different microbial groups affect extracellular H2O2 concentrations, how comparable extracellular H2O2 concentrations within large-scale incubation experiments are to those observed in the surface-mixed layer, and to what extent a mismatch with environmentally relevant concentrations of ROS in incubations could influence biological processes differently to what would be observed in nature. Here we show that both experiment design and bacterial abundance consistently exert control on extracellular H2O2 concentrations across a range of incubation experiments in diverse marine environments. During four large-scale ( >1000 L) mesocosm experiments (in Gran Canaria, the Mediterranean, Patagonia and Svalbard) most experimental factors appeared to exert only minor, or no, direct effect on H2O2 concentrations. For example, in three of four experiments where pH was manipulated to 0.4–0.5 below ambient pH, no significant change was evident in extracellular H2O2 concentrations relative to controls. An influence was sometimes inferred from zooplankton density, but not consistently between different incubation experiments, and no change in H2O2 was evident in controlled experiments using different densities of the copepod Calanus finmarchicus grazing on the diatom Skeletonema costatum ( % change in [ H2O2 ] comparing copepod densities from 1 to 10 L −1 ). Instead, the changes in H2O2 concentration contrasting high- and low-zooplankton incubations appeared to arise from the resulting changes in bacterial activity. The correlation between bacterial abundance and extracellular H2O2 was stronger in some incubations than others ( R2 range 0.09 to 0.55), yet high bacterial densities were consistently associated with low H2O2 . Nonetheless, the main control on H2O2 concentrations during incubation experiments relative to those in ambient, unenclosed waters was the regulation of incident light. In an open (lidless) mesocosm experiment in Gran Canaria, H2O2 was persistently elevated (2–6-fold) above ambient concentrations; whereas using closed high-density polyethylene mesocosms in Crete, Svalbard and Patagonia H2O2 within incubations was always reduced (median 10 %–90 %) relative to ambient waters.

The thriving interest in harvesting deep-sea mineral resources, such as polymetallic nodules, calls for environmental impact studies and, ultimately, for regulations for environmental protection. Industrial-scale deep-sea mining of polymetallic nodules most likely has severe consequences for the natural environment. However, the effects of mining activities on deep-sea ecosystems, sediment geochemistry and element fluxes are still poorly understood. Predicting the environmental impact is challenging due to the scarcity of environmental baseline studies as well as the lack of mining trials with industrial mining equipment in the deep sea. Thus, currently we have to rely on small-scale disturbances simulating deep-sea mining activities as a first-order approximation to study the expected impacts on the abyssal environment. Here, we investigate surface sediments in disturbance tracks of seven small-scale benthic impact experiments, which have been performed in four European contract areas for the exploration of polymetallic nodules in the Clarion–Clipperton Zone (CCZ) in the NE Pacific. These small-scale disturbance experiments were performed 1 d to 37 years prior to our sampling program in the German, Polish, Belgian and French contract areas using different disturbance devices. We show that the depth distribution of solid-phase Mn in the upper 20 cm of the sediments in the CCZ provides a reliable tool for the determination of the disturbance depth, which has been proposed in a previous study from the SE Pacific (Paul et al., 2018). We found that the upper 5–15 cm of the sediments was removed during various small-scale disturbance experiments in the different exploration contract areas. Transient transport-reaction modeling for the Polish and German contract areas reveals that the removal of the surface sediments is associated with the loss of the reactive labile total organic carbon (TOC) fraction. As a result, oxygen consumption rates decrease significantly after the removal of the surface sediments, and, consequently, oxygen penetrates up to 10-fold deeper into the sediments, inhibiting denitrification and Mn(IV) reduction. Our model results show that the return to steady-state geochemical conditions after the disturbance is controlled by diffusion until the reactive labile TOC fraction in the surface sediments is partly re-established and the biogeochemical processes commence. While the re-establishment of bioturbation is essential, steady-state geochemical conditions are ultimately controlled by the delivery rate of organic matter to the seafloor. Hence, under current depositional conditions, new steady-state geochemical conditions in the sediments of the CCZ are reached only on a millennium scale even for these small-scale disturbances simulating deep-sea mining activities.

Rising concentrations of atmospheric carbon dioxide are causing ocean acidification and will influence marine processes and trace metal biogeochemistry. In June 2012, in the Raunefjord (Bergen, Norway), we performed a mesocosm experiment, comprised of a fully factorial design of ambient and elevated pCO2 and/or an addition of the siderophore desferrioxamine B (DFB). In addition, the macronutrient concentrations were manipulated to enhance a bloom of the coccolithophore Emiliania huxleyi. We report the changes in particulate trace metal concentrations during this experiment. Our results show that particulate Ti and Fe were dominated by lithogenic material, while particulate Cu, Co, Mn, Zn, Mo and Cd had a strong biogenic component. Furthermore, significant correlations were found between particulate concentrations of Cu, Co, Zn, Cd, Mn, Mo and P in seawater and phytoplankton biomass ( µ gC L −1 ), supporting a significant influence of the bloom in the distribution of these particulate elements. The concentrations of these biogenic metals in the E. huxleyi bloom were ranked as follows: Zn Cu ≈ Mn Mo Co Cd. Changes in CO2 affected total particulate concentrations and biogenic metal ratios (Me : P) for some metals, while the addition of DFB only significantly affected the concentrations of some particulate metals (mol L −1 ). Variations in CO2 had the most clear and significant effect on particulate Fe concentrations, decreasing its concentration under high CO2 . Indeed, high CO2 and/or DFB promoted the dissolution of particulate Fe, and the presence of this siderophore helped in maintaining high dissolved Fe. This shift between particulate and dissolved Fe concentrations in the presence of DFB, promoted a massive bloom of E. huxleyi in the treatments with ambient CO2 . Furthermore, high CO2 decreased the Me : P ratios of Co, Zn and Mn while increasing the Cu : P ratios. These findings support theoretical predictions that the molar ratios of metal to phosphorous (Me : P ratios) of metals whose seawater dissolved speciation is dominated by free ions (e.g., Co, Zn and Mn) will likely decrease or stay constant under ocean acidification. In contrast, high CO2 is predicted to shift the speciation of dissolved metals associated with carbonates such as Cu, increasing their bioavailability and resulting in higher Me : P ratios.

Compared to the rest of the globe, the Arctic Ocean is affected disproportionately by climate change. Despite these fast environmental changes, we currently know little about the effects of ocean acidification (OA) on marine key species in this area. Moreover, the existing studies typically test the effects of OA under constant, hence artificial, light fields. In this study, the abundant Arctic picoeukaryote Micromonas pusilla was acclimated to current (400 µatm) and future (1000 µatm) pCO2 levels under a constant as well as a dynamic light, simulating more realistic light fields as experienced in the upper mixed layer. To describe and understand the responses to these drivers, growth, particulate organic carbon (POC) production, elemental composition, photophysiology and reactive oxygen species (ROS) production were analysed. M. pusilla was able to benefit from OA on various scales, ranging from an increase in growth rates to enhanced photosynthetic capacity, irrespective of the light regime. These beneficial effects were, however, not reflected in the POC production rates, which can be explained by energy partitioning towards cell division rather than biomass build-up. In the dynamic light regime, M. pusilla was able to optimize its photophysiology for effective light usage during both low- and high-light periods. This photoacclimative response, which was achieved by modifications to photosystem II (PSII), imposed high metabolic costs leading to a reduction in growth and POC production rates when compared to constant light. There were no significant interactions observed between dynamic light and OA, indicating that M. pusilla is able to maintain effective photoacclimation without increased photoinactivation under high pCO2. Based on these findings, M. pusilla is likely to cope well with future conditions in the Arctic Ocean.

Micronekton – small marine pelagic organisms mostly in the size range 1–10 cm – is a key component of the ocean ecosystem, as it constitutes the main source of forage for all larger predators. Moreover, the mesopelagic component of micronekton that undergoes Diel Vertical Migration (DVM) likely plays a key role in the transfer and storage of CO2 in the deep ocean: the so-called ‘biological pump’ mechanism. SEAPODYM-MTL is a spatially explicit dynamical model of micronekton. It simulates six functional groups of migrant and non-migrant micronekton, in the epipelagic and mesopelagic layers. Coefficients of energy transfer efficiency between primary production and each group are unknown. But they are essential as they control the predicted biomass. Since these coefficients are not directly measurable, a data assimilation method is used to estimate them. In this study, Observing System Simulation Experiments (OSSE) in the framework of twin experiments are used to test various observation networks at a global scale regarding energy transfer coefficients estimation. Observational networks show a variety of performances. It appears that environmental conditions are crucial to determine network efficiency. According to our study, ideal sampling areas are warm, non-dynamic and productive waters like the eastern side of tropical Oceans. These regions are found to reduce the error of estimated coefficients by 20 % compared to cold and dynamic sampling regions. The results are discussed in term of interactions between physical and biological processes.

[EN]Southern Ocean waters are projected to undergo profound changes in their physical and chemical properties in the coming decades. Coccolithophore blooms in the Southern Ocean are thought to account for a major fraction of the global marine calcium carbonate (CaCO3) production and export to the deep sea. Therefore, changes in the composition and abundance of Southern Ocean coccolithophore populations are likely to alter the marine carbon cycle, with feedbacks to the rate of global climate change. However, the contribution of coccolithophores to CaCO3 export in the Southern Ocean is uncertain, particularly in the circumpolar subantarctic zone that represents about half of the areal extent of the Southern Ocean and where coccolithophores are most abundant. Here, we present measurements of annual CaCO3 flux and quantitatively partition them amongst coccolithophore species and heterotrophic calcifiers at two sites representative of a large portion of the subantarctic zone. We find that coccolithophores account for a major fraction of the annual CaCO3 export, with the highest contributions in waters with low algal biomass accumulations. Notably, our analysis reveals that although Emiliania huxleyi is an important vector for CaCO3 export to the deep sea, less abundant but larger species account for most of the annual coccolithophore CaCO3 flux. This observation contrasts with the generally accepted notion that high particulate inorganic carbon accumulations during the austral summer in the subantarctic Southern Ocean are mainly caused by E. huxleyi blooms. It appears likely that the climate-induced migration of oceanic fronts will initially result in the poleward expansion of large coccolithophore species increasing CaCO3 production. However, subantarctic coccolithophore populations will eventually diminish as acidification overwhelms those changes. Overall, our analysis emphasizes the need for species-centred studies to improve our ability to project future changes in phytoplankton communities and their influence on marine biogeochemical cycles., European Union's Horizon 2020, Marie Skłodowska-Curie Individual fellowship

Rivers are a major source of nutrients, carbon and alkalinity to the global ocean. In this study, we firstly estimate pre-industrial riverine loads of nutrients, carbon and alkalinity based on a hierarchy of weathering and terrestrial organic matter export models, while identifying regional hotspots of the riverine exports. Secondly, we implement the riverine loads into a global ocean biogeochemical model to describe their implications for oceanic nutrient concentrations, net primary production (NPP) and air–sea CO2 fluxes globally, as well as in an analysis of coastal regions. Thirdly, we quantitatively assess the terrestrial origins and the long-term fate of riverine carbon in the ocean. We quantify annual bioavailable pre-industrial riverine loads of 3.7 Tg P, 27 Tg N, 158 Tg Si and 603 Tg C delivered to the ocean globally. We thereby identify the tropical Atlantic catchments (20 % of global C), Arctic rivers (9 % of global C) and Southeast Asian rivers (15 % of global C) as dominant suppliers of carbon for the ocean. The riverine exports lead to a simulated net global oceanic CO2 source of 231 Tg C yr−1 to the atmosphere, which is mainly caused by inorganic carbon (source of 183 Tg C yr−1) and by organic carbon (source of 128 Tg C yr−1) riverine loads. Additionally, a sink of 80 Tg C yr−1 is caused by the enhancement of the biological carbon uptake from dissolved inorganic nutrient inputs from rivers and the resulting alkalinity production. While large outgassing fluxes are simulated mostly in proximity to major river mouths, substantial outgassing fluxes can be found further offshore, most prominently in the tropical Atlantic. Furthermore, we find evidence for the interhemispheric transfer of carbon in the model; we detect a larger relative outgassing flux (49 % of global riverine-induced outgassing) in the Southern Hemisphere in comparison to the hemisphere's relative riverine inputs (33 % of global C inputs), as well as an outgassing flux of 17 Tg C yr−1 in the Southern Ocean. The addition of riverine loads in the model leads to a strong NPP increase in the tropical west Atlantic, Bay of Bengal and the East China Sea (+166 %, +377 % and +71 %, respectively). On the light-limited Arctic shelves, the NPP is not strongly sensitive to riverine loads, but the CO2 flux is strongly altered regionally due to substantial dissolved inorganic and organic carbon supplies to the region. While our study confirms that the ocean circulation remains the main driver for biogeochemical distributions in the open ocean, it reveals the necessity to consider riverine inputs for the representation of heterogeneous features in the coastal ocean and to represent riverine-induced pre-industrial carbon outgassing in the ocean. It also underlines the need to consider long-term CO2 sources from volcanic and shale oxidation fluxes in order to close the framework's atmospheric carbon budget.

Global warming will be combined with predicted increases in thermal variability in the future surface ocean, but how temperature dynamics will affect phytoplankton biology and biogeochemistry is largely unknown. Here, we examine the responses of the globally important marine coccolithophore Emiliania huxleyi to thermal variations at two frequencies (one-day and two-day) at low (18.5 °C) and high (25.5 °C) mean temperatures. Elevated temperature and thermal variation decreased growth, calcification and physiological rates, both individually and interactively. One-day thermal variation frequencies were less inhibitory than two-day variations under high temperature, indicating that high frequency thermal fluctuations may reduce heat‐induced mortality and mitigate some impacts of extreme high temperature events. Cellular elemental composition and calcification was significantly affected by both thermal variation treatments relative to each other, and to the constant temperature controls. The negative effects of thermal variation on E. huxleyi growth rate and physiology are especially pronounced at high temperatures. These responses of the key marine calcifier E. huxleyi to warmer, more variable temperature regimes have potentially large implications for ocean productivity and marine biogeochemical cycles under a future changing climate.

The biogenic sulfur compounds dimethyl sulfide (DMS), dimethyl sulfoniopropionate (DMSP) and dimethyl sulfoxide (DMSO) are produced and transformed by diverse populations of marine microorganisms and have substantial physiological, ecological and biogeochemical importance spanning organism to global scales. Understanding the production and transformation dynamics of these compounds under shifting environmental conditions is important for predicting their roles in a changing ocean. Here, we report the physiological and biochemical response of a robust strain of Alexandrium minutum, a dinoflagellate with the highest reported intracellular DMSP content, exposed to a 6 d increase in temperature mimicking mild and extreme coastal marine heatwave conditions (+4 and +12 ∘C). Under mild temperature increases (+4 ∘C), A. minutum growth was enhanced, with no measurable physiological stress response. However, under a very acute increase in temperature (+12 ∘C) triggering thermal stress, A. minutum growth declined, photosynthetic efficiency (FV∕FM) was impaired, and enhanced oxidative stress was observed. These physiological responses indicative of thermal stress were accompanied by increased DMS and DMSO concentrations followed by decreased DMSP concentration. At this temperature extreme, we observed a cascading stress response in A. minutum, which was initiated 6 h after the start of the experiment by a spike in DMS and DMSO concentrations and a rapid decrease in FV∕FM. This was followed by an increase in reactive oxygen species (ROS) and an abrupt decline in DMS and DMSO on day 2 of the experiment. A subsequent decrease in DMSP coupled with a decline in the growth rate of both A. minutum and its associated total bacterial assemblage coincided with a shift in the composition of the A. minutum microbiome. Specifically, an increase in the relative abundance of the operational taxonomic units (OTUs) matching Oceanicaulis (17.0 %), Phycisphaeraceae SM1A02 (8.8 %) and Balneola (4.9 %) as well as a decreased relative abundance of Maribacter (24.4 %), Marinoscillum (4.7 %) and Seohaeicola (2.7 %) were primarily responsible for differences in microbiome structure observed between temperature treatments. These shifts in microbiome structure are likely to have been driven by either the temperature itself, the changing physiological state of A. minutum cells, shifts in biogenic sulfur concentrations, the presence of other solutes, or a combination of all. Nevertheless, we suggest that these results point to the significant effect of extreme heatwaves on the physiology, growth and microbiome composition of the red-tide causing dinoflagellate A. minutum, as well as potential implications for biogenic sulfur cycling processes and marine DMS emissions.

Nitric oxide (NO) is a short-lived compound of the marine nitrogen cycle; however, our knowledge about its oceanic distribution and turnover is rudimentary. Here we present the measurements of dissolved NO in the surface and bottom layers at 75 stations in the Bohai Sea (BS) and the Yellow Sea (YS) in June 2011. Moreover, NO photoproduction rates were determined at 27 stations in both seas. The NO concentrations in the surface and bottom layers were highly variable and ranged from below the limit of detection (i.e., 32 pmol L−1) to 616 pmol L−1 in the surface layer and 482 pmol L−1 in the bottom layer. There was no significant difference (p>0.05) between the mean NO concentrations in the surface (186±108 pmol L−1) and bottom (174±123 pmol L−1) layers. A decreasing trend of NO in bottom-layer concentrations with salinity indicates a NO input by submarine groundwater discharge. NO in the surface layer was supersaturated at all stations during both day and night and therefore the BS and YS were a persistent source of NO to the atmosphere at the time of our measurements. The average flux was about 4.5×10-16 mol cm−2 s−1 and the flux showed significant positive relationship with the wind speed. The accumulation of NO during daytime was a result of photochemical production, and photoproduction rates were correlated to illuminance. The persistent nighttime NO supersaturation pointed to an unidentified NO dark production. NO sea-to-air flux densities were much lower than the NO photoproduction rates. Therefore, we conclude that the bulk of the NO produced in the mixed layer was rapidly consumed before its release to the atmosphere.

Rapidly changing climate in the Northern Hemisphere and associated socio-economic impacts require reliable understanding of lake systems as important freshwater resources and sensitive sentinels of environmental change. To better understand time-series data in lake sediment cores, it is necessary to gain information on within-lake spatial variabilities of environmental indicator data. Therefore, we retrieved a set of 38 samples from the sediment surface along spatial habitat gradients in the boreal, deep, and yet pristine Lake Bolshoe Toko in southern Yakutia, Russia. Our methods comprise laboratory analyses of the sediments for multiple proxy parameters, including diatom and chironomid taxonomy, oxygen isotopes from diatom silica, grain-size distributions, elemental compositions (XRF), organic carbon content, and mineralogy (XRD). We analysed the lake water for cations, anions, and isotopes. Our results show that the diatom assemblages are strongly influenced by water depth and dominated by planktonic species, i.e. Pliocaenicus bolshetokoensis. Species richness and diversity are higher in the northern part of the lake basin, associated with the availability of benthic, i.e. periphytic, niches in shallower waters. δ18Odiatom values are higher in the deeper south-western part of the lake, probably related to water temperature differences. The highest amount of the chironomid taxa underrepresented in the training set used for palaeoclimate inference was found close to the Utuk River and at southern littoral and profundal sites. Abiotic sediment components are not symmetrically distributed in the lake basin, but vary along restricted areas of differential environmental forcing. Grain size and organic matter are mainly controlled by both river input and water depth. Mineral (XRD) data distributions are influenced by the methamorphic lithology of the Stanovoy mountain range, while elements (XRF) are intermingled due to catchment and diagenetic differences. We conclude that the lake represents a valuable archive for multiproxy environmental reconstruction based on diatoms (including oxygen isotopes), chironomids, and sediment–geochemical parameters. Our analyses suggest multiple coring locations preferably at intermediate depth in the northern basin and the deep part in the central basin, to account for representative bioindicator distributions and higher temporal resolution, respectively.

The intertidal bivalve Katelysia rhytiphora, endemic to south Australia and Tasmania, is used here for pulsed Sr-labelling experiments in aquaculture experiments to visualize shell growth at the micro- to nanoscale. The ventral margin area of the outer shell layer composed of (i) an outermost outer shell layer (oOSL) with compound composite prismatic architecture with three hierarchical orders of prisms and (ii) an innermost outer shell layer (iOSL) with crossed-acicular architecture consisting of intersecting lamellae bundles. All structural orders in both layers are enveloped by an organic sheath and the smallest mineralized units are nano-granules. Electron backscatter diffraction reveals a strong preferred orientation of the aragonite c axes perpendicular to the growth layers, while the a and b axes are scattered within a plane normal to the local growth direction and >46 % twin grain boundaries are detected. The Young's modulus shows a girdle-like maximum of elastically stiffer orientations for the shell following the inner shell surface. For 6 d, the bivalves were subjected twice to seawater with an increased Sr concentration of 18× mean ocean water by dissolving 144 µ g g −1 Sr (159.88 Sr∕Ca mmol ∕ mol) in seawater. The pulse labelling intervals in the shell are 17× (oOSL) and 12× (iOSL) enriched in Sr relative to the Sr-spiked seawater. All architectural units in the shell are transected by the Sr label, demonstrating shell growth to progress homogeneously instead of forming one individual architectural unit after the other. Distribution coefficients, DSr ∕ Ca , for labelled and unlabelled shells are similar to shell proportions formed in the wild (0.12 to 0.15). All DSr ∕ Ca values are lower than values for equilibrium partitioning of Sr in synthetic aragonite.

Mining of polymetallic nodules in abyssal seafloor sediments promises to address the growing worldwide demand for metallic minerals. Given that prospective mining operations are likely to have profound impacts on deep seafloor communities, industrial investment has been accompanied by scientific involvement for the assessment of baseline conditions and provision of guidelines for environmentally sustainable mining practices. Benthic meiofaunal communities were studied in four prospective mining areas of the Clarion–Clipperton Zone (CCZ) in the eastern Pacific Ocean, arranged in a southeast–northwest fashion coinciding with the productivity gradient in the area. Additionally, samples were collected from the Area of Particular Environmental Interest no. 3 (APEI-3) in the northwest of the CCZ, where mining will be prohibited and which should serve as a “source area” for the biota within the larger CCZ. Total densities in the 0–5 cm upper layer of the sediment were influenced by sedimentary characteristics, water depth and nodule density at the various sampling locations, indicating the importance of nodules for meiofaunal standing stock. Nematodes were the most abundant meiobenthic taxon, and their assemblages were typically dominated by a few genera (generally 2–6) accounting for 40 %–70 % of all individuals, which were also widely spread along the CCZ and shared among all sampled license areas. However, almost half of the communities consisted of rare genera, each contributing less than 5 % to the overall abundances and displaying a distribution which was usually restricted to a single license area. The same observations (dominant and widely spread versus rare and scattered) could be made for the species of one of the dominant genera, Halalaimus, implying that it might be mainly these rare genera and species that will be vulnerable to mining-induced changes in their habitat.

Particle aggregation determines the particle flux length scale and affects the marine oxygen concentration and thus the volume of oxygen minimum zones (OMZs) that are of special relevance for ocean nutrient cycles and marine ecosystems and that have been found to expand faster than can be explained by current state-of-the-art models. To investigate the impact of particle aggregation on global model performance, we carried out a sensitivity study with different parameterisations of marine aggregates and two different model resolutions. Model performance was investigated with respect to global nutrient and oxygen concentrations, as well as extent and location of OMZs. Results show that including an aggregation model improves the representation of OMZs. Moreover, we found that besides a fine spatial resolution of the model grid, the consideration of porous particles, an intermediate-to-high particle sinking speed and a moderate-to-high stickiness improve the model fit to both global distributions of dissolved inorganic tracers and regional patterns of OMZs, compared to a model without aggregation. Our model results therefore suggest that improvements not only in the model physics but also in the description of particle aggregation processes can play a substantial role in improving the representation of dissolved inorganic tracers and OMZs on a global scale. However, dissolved inorganic tracers are apparently not sufficient for a global model calibration, which could necessitate global model calibration against a global observational dataset of marine organic particles.

Industrial interest in deep-sea mineral extraction began decades ago, and today it is at an all-time high, accelerated by global demand for metals. Several seafloor ecosystem disturbance experiments began in the 1970s, including the Disturbance and Recolonization experiment (DISCOL) conducted in the Peru Basin in 1989. A large seafloor disturbance was created by repeatedly ploughing the seafloor over an area of ∼10.8 km2. Though a number of studies in abyssal mining regions have evaluated megafaunal biodiversity and ecosystem responses, few have included quantitative and detailed data on fishes or scavengers despite their ecological importance as top predators. We used towed camera transects (1989–1996, 2015) and baited camera data (1989–1992) to evaluate the fish community at the DISCOL site. The abyssal fish community included 16 taxa and was dominated by Ipnops meadi. Fish density was lower in ploughed habitat at 6 months and 3 years following disturbance but thereafter increased over time. Twenty-six years after disturbance there were no differences in overall total fish densities between reference and experimental areas, but the dominant fish, I. meadi, still exhibited much lower densities in ploughed habitat, likely avoiding these areas and suggesting that the fish community remains affected after decades. At the scale of industrial mining, these results could translate to population-level effects. The scavenging community was dominated by eelpouts (Pachycara spp.), hermit crabs (Probeebei mirabilis) and shrimp. The large contribution of hermit crabs appears to be unique amongst abyssal scavenger studies worldwide. The abyssal fish community at DISCOL was similar to that in the more northerly Clarion–Clipperton Zone (CCZ), though some species have only been observed at DISCOL thus far. Also, further species-level identifications are required to refine this assessment. Additional studies across the polymetallic nodule provinces of the Pacific are required to further evaluate the environmental drivers of fish density, diversity and species biogeographies. This information will be important for the development of appropriate management plans aimed at minimizing human impact from deep-sea mining.

The concentration of CO2 in the atmosphere is sensitive to changes in the depth at which sinking particulate organic matter is remineralized: often described as a change in the exponent “b” of the Martin curve. Sediment trap observations from deep and intermediate depths suggest there is a spatially heterogeneous pattern of b, particularly varying with latitude, but disagree over the exact spatial patterns. Here we use a biogeochemical model of the phosphorus cycle coupled with a steady-state representation of ocean circulation to explore the sensitivity of preformed phosphate and atmospheric CO2 to spatial variability in remineralization depths. A Latin hypercube sampling method is used to simultaneously vary the Martin curve independently within 15 different regions, as a basis for a regression-based analysis used to derive a quantitative measure of sensitivity. Approximately 30 % of the sensitivity of atmospheric CO2 to changes in remineralization depths is driven by changes in the subantarctic region (36 to 60∘ S) similar in magnitude to the Pacific basin despite the much smaller area and lower export production. Overall, the absolute magnitude of sensitivity is controlled by export production, but the relative spatial patterns in sensitivity are predominantly constrained by ocean circulation pathways. The high sensitivity in the subantarctic regions is driven by a combination of high export production and the high connectivity of these regions to regions important for the export of preformed nutrients such as the Southern Ocean and North Atlantic. Overall, regionally varying remineralization depths contribute to variability in CO2 of between around 5 and 15 ppm, relative to a global mean change in remineralization depth. Future changes in the environmental and ecological drivers of remineralization, such as temperature and ocean acidification, are expected to be most significant in the high latitudes where CO2 sensitivity to remineralization is also highest. The importance of ocean circulation pathways to the high sensitivity in subantarctic regions also has significance for past climates given the importance of circulation changes in the Southern Ocean.

Biochemical markers developed initially for food-web studies of terrestrial and shallow-water environments have only recently been applied to deep-sea ecosystems (i.e., in the early 2000s). For the first time since their implementation, this review took a close look at the existing literature in the field of deep-sea trophic ecology to synthesize current knowledge. Furthermore, it provided an opportunity for a preliminary analysis of global geographic (i.e., latitudinal, along a depth gradient) trends in the isotopic (δ15N, δ13C) and fatty acid composition of deep-sea macro- and megafauna from heterotrophic systems. Results revealed significant relationships along the latitudinal and bathymetric gradients. Deep-sea animals sampled at temperate and polar latitudes displayed lower isotopic ratios and greater proportions of essential ω3 long-chain polyunsaturated fatty acids (LC-PUFAs) than did tropical counterparts. Furthermore, δ15N and δ13C ratios as well as proportions of arachidonic acid increased with increasing depth. Since similar latitudinal trends in the isotopic and fatty acid composition were found in surface water phytoplankton and particulate organic matter, these results highlight the link across latitudes between surface primary production and deep-water communities. Because global climate change may affect quantity and quality (e.g., levels of essential ω3 PUFAs) of surface primary productivity, and by extension those of its downward flux, the dietary intake of deep-sea organisms may likely be altered. In addition, because essential ω3 PUFAs play a major role in the response to temperature variations, climate change may interfere with the ability of deep-sea species to cope with potential temperature shifts. Importantly, methodological disparities were highlighted that prevented in-depth analyses, indicating that further studies should be conducted using standardized methods in order to generate more reliable global predictions.