ALBERT

Description: The northern Humboldt Current upwelling system (HCS) belongs to the most productive marine ecosystems, providing five to eight times higher fisheries landings per unit area than other coastal upwelling systems. To solve this “Peruvian puzzle”, to elucidate the pelagic food-web structure and to better understand trophic interactions in the HCS, a combined stable isotope and fatty acid trophic biomarker approach was adopted for key zooplankton taxa and higher trophic positions with an extensive spatial coverage from 8.5 to 16°S and a vertical range down to 1,000 m depth. A pronounced regional shift by up to ∼5‰ in the δ15N baseline of the food web occurred from North to South. Besides regional shifts, δ15N ratios of particulate organic matter (POM) also tended to increase with depth, with differences of up to 3.8‰ between surface waters and the oxygen minimum zone. In consequence, suspension-feeding zooplankton permanently residing at depth had up to ∼6‰ higher δ15N signals than surface-living species or diel vertical migrants. The comprehensive data set covered over 20 zooplankton taxa and indicated that three crustacean species usually are key in the zooplankton community, i.e., the copepods Calanus chilensis at the surface and Eucalanus inermis in the pronounced OMZ and the krill Euphausia mucronata, resulting in an overall low number of major trophic pathways toward anchovies. In addition, the semi-pelagic squat lobster Pleuroncodes monodon appears to play a key role in the benthic-pelagic coupling, as indicated by highest δ13C’ ratios of −14.7‰. If feeding on benthic resources and by diel vertical migration, they provide a unique pathway for returning carbon and energy from the seafloor to the epipelagic layer, increasing the food supply for pelagic fish. Overall, these mechanisms result in a very efficient food chain, channeling energy toward higher trophic positions and partially explaining the “Peruvian puzzle” of enormous fish production in the HCS.

Description: Fucoidanases are endo-fucoidanases (also known as endo-fucanases) that catalyze hydrolysis of α-glycosidic linkages in fucoidans, a family of sulfated fucose-rich polysaccharides primarily found in the cell walls of brown seaweeds. Fucoidanases are promising tools for producing bioactive fucoidan oligosaccharides for a range of biomedical applications. High sulfation degree has been linked to high bioactivity of fucoidans. In this study, a novel fucoidanase, Fhf2, was identified in the genome of the aerobic, Gram-negative marine bacterium Formosa haliotis. Fhf2 was found to share sequence similarity to known endo-α(1,4)-fucoidanases (EC 3.2.1.212) from glycoside hydrolase family 107. A C-terminal deletion mutant Fhf2∆484, devoid of 484 amino acids at the C-terminus, with a molecular weight of approximately 46 kDa, was constructed and found to be more stable than the full-length Fhf2 protein. Fhf2∆484 showed endo-fucoidanase activity on fucoidans from different seaweed species including Fucus evanescens, Fucus vesiculosus, Sargassum mcclurei, and Sargassum polycystum. The highest activity was observed on fucoidan from F. evanescens. The Fhf2∆484 enzyme was active at 20–45°C and at pH 6–9 and had optimal activity at 37°C and pH 8. Additionally, Fhf2∆484 was found to be calcium-dependent. NMR analysis showed that Fhf2∆484 catalyzed hydrolysis of α(1,4) linkages between L-fucosyl moieties sulfated on C2 (similar to Fhf1 from Formosa haliotis), but Fhf2∆484 in addition released oligosaccharides containing a substantial amount of 2,4-disulfated fucose residues. The data thus suggest that the Fhf2∆484 enzyme could be a valuable candidate for producing highly sulfated oligosaccharides applicable for fucoidan bioactivity investigations.

Description: Trichodesmium spp. is a colonial diazotrophic cyanobacterium found in the oligotrophic (sub)tropical oceans, where dissolved inorganic phosphorus (DIP) can be depleted. To cope with low P concentrations, P can be scavenged from the dissolved organic P (DOP) pool. This requires the deployment of multiple enzymes activated by trace metals, potentially enhancing metal requirements under stronger P limitations. To test this, we grew Trichodesmium under trace-metal-controlled conditions, where P was supplied as either DIP or DOP (methylphosphonic acid). Mean steady-state biomass under the DOP treatment was only 40% of that grown under equivalent DIP supply, carbon normalized alkaline phosphorus activity was elevated 4-fold, and the zinc (Zn)–carbon ratio was elevated 3.5-fold. Our finding matches the known, dominant Zn requirement across a diversity of enzymes involved in P stress responses and supports an important interaction in the oceanic cycles of these two nutrients.

Description: Paleoclimate and paleoenvironmental reconstructions from increment-yielding archives strongly depend on precise age models. Like bivalves, corals, trees, and speleothems, the coralline alga Clathromorphum compactum produces annual growth increments and shows considerable promise as an environmental archive for arctic and subarctic regions. Though their growth increment widths correlate with temperature and sea ice cover in high Arctic regions, existing timeseries have not been crossdated. In fact, previous studies have shown a lack of inter-sample correlation in non-crossdated timeseries suggesting possible age model dating errors. Here, we use dendrochronology crossdating techniques and COFECHA software to ensure and validate synchrony between C. compactum timeseries (〈141 years) from three specimens collected near Beechey Island, Nunavut, Canada. Results showed that non-crossdated timeseries constructed by four coralline red algae researchers using annual increments of the same C. compactum samples were highly variable and showcase the likelihood of dating errors in non-crossdated timeseries. Crossdating improved inter-series correlations, and correlations to sea ice-related records, suggesting that at least three crossdated timeseries are required to isolate paleoclimate signals. Our findings suggest that future reconstructions with C. compactum should employ crossdating techniques to reduce dating errors and allow for more precise climate reconstructions.

Description: The picophytoplankton Synechococcus is a globally abundant autotroph that contributes significantly to primary production in the oceans and coastal areas. These cyanobacteria constitute a diverse genus of organisms that have developed independent niche spaces throughout aquatic environments. Here, we use the 16S V3–V4 rRNA gene region and flow cytometry to explore the diversity of Synechococcus within the picophytoplankton community in the Gullmar Fjord, on the west coast of Sweden. We conducted a station-based 1-year time series and two transect studies of the fjord. Our analysis revealed that within the large number of Synechococcus amplicon sequence variants (ASVs; 239 in total), prevalent ASVs phylogenetically clustered with clade representatives in both marine subcluster 5.1 and 5.2. The near-surface composition of ASVs shifted from spring to summer, when a 5.1 subcluster dominated community developed along with elevated Synechococcus abundances up to 9.3 × 104 cells ml–1. This seasonal dominance by subcluster 5.1 was observed over the length of the fjord (25 km), where shifts in community composition were associated with increasing depth. Unexpectedly, the community shift was not associated with changes in salinity. Synechococcus abundance dynamics also differed from that of the photosynthetic picoeukaryote community. These results highlight how seasonal variations in environmental conditions influence the dynamics of Synechococcus clades in a high latitude threshold fjord.

Description: Anthropogenic CO2 emissions cause a drop in seawater pH and shift the inorganic carbon speciation. Collectively, the term ocean acidification (OA) summarizes these changes. Few studies have examined OA effects on predatory plankton, e.g. Hydrozoa and fish larvae as well as their interaction in complex natural communities. Because Hydrozoa can seriously compete with and prey on other higher-level predators such as fish, changes in their abundances may have significant consequences for marine food webs and ecosystem services. To investigate the interaction between Hydrozoa and fish larvae influenced by OA, we enclosed a natural plankton community in Raunefjord, Norway, for 53 days in eight ≈ 58 m³ pelagic mesocosms. CO2 levels in four mesocosms were increased to ≈ 2000 µatm pCO2, whereas the other four served as untreated controls. We studied OA-induced changes at the top of the food web by following ≈2000 larvae of Atlantic herring (Clupea harengus) hatched inside each mesocosm during the first week of the experiment, and a Hydrozoa population that had already established inside the mesocosms. Under OA, we detected 20% higher abundance of hydromedusae staged jellyfish, but 25% lower biomass. At the same time, survival rates of Atlantic herring larvae were higher under OA (control pCO2: 0.1%, high pCO2: 1.7%) in the final phase of the study. These results indicate that a decrease in predation pressure shortly after hatch likely shaped higher herring larvae survival, when hydromedusae abundance was lower in the OA treatment compared to control conditions. We conclude that indirect food-web mediated OA effects drove the observed changes in the Hydrozoa – fish relationship, based on significant changes in the phyto-, micro-, and mesoplankton community under high pCO2. Ultimately, the observed immediate consequences of these changes for fish larvae survival and the balance of the Hydrozoa – fish larvae predator – prey relationship has important implications for the functioning of oceanic food webs.

Description: Oxygen minimum zones (OMZs) have been suggested as a suitable niche for the oxygen-sensitive process of biological fixation of dinitrogen (N2) gas. However, most N2 fixation rates reported from such waters are low. This low N2 fixation activity has been proposed to result from the unusual community of N2 fixers, in which cyanobacteria were typically underrepresented. The Northern Benguela Upwelling System (North BUS) is part of one of the most productive marine ecosystems and hosts a well-developed OMZ. Although previous observations indicated low to absent N2 fixation rates, the community composition of diazotrophs needed to understand the North BUS has not been described. Here, we present a first detailed analysis of the diazotrophic diversity in the North BUS OMZ and the Angola tropical zone (ATZ), based on genetic data and isotope speciation. Consistent with a previous study, we detected a slight N deficit in the OMZ, but isotope data did not indicate any active or past N2 fixation. The diazotroph community in the North BUS was dominated by non-cyanobacterial microbes clustering with members of gamma-proteobacteria, as is typical for other OMZ regions. However, we found a strikingly high diversity of Cluster III diazotrophs not yet described in other OMZs. In contrast to previous observations, we could also identify cyanobacteria of the clades Trichodesmium sp., UCYN-A and Cyanothece sp., in surface waters connected to or above the OMZ, which were potentially active as shown by the presence of genes and transcripts of the key functional marker gene for N2 fixation, nifH. While the detection of diazotrophs and the absence of active N2 fixation (based on isotopic speciation) are consistent with other OMZ observations, the detected regional variation in the diversity and presence of cyanobacteria indicate that we still are far from understanding the role of diazotrophs in OMZs, which, however, is relevant for understanding the N cycle in OMZ waters, as well for predicting the future development of OMZ biogeochemistry in a changing ocean.

Description: The abyssal seafloor in the Clarion-Clipperton Zone (CCZ) in the NE Pacific hosts the largest abundance of polymetallic nodules in the deep sea and is being targeted as an area for potential deep-sea mining. During nodule mining, seafloor sediment will be brought into suspension by mining equipment, resulting in the formation of sediment plumes, which will affect benthic and pelagic life not naturally adapted to any major sediment transport and deposition events. To improve our understanding of sediment plume dispersion and to support the development of plume dispersion models in this specific deep-sea area, we conducted a small-scale, 12-hour disturbance experiment in the German exploration contract area in the CCZ using a chain dredge. Sediment plume dispersion and deposition was monitored using an array of optical and acoustic turbidity sensors and current meters placed on platforms on the seafloor, and by visual inspection of the seafloor before and after dredge deployment. We found that seafloor imagery could be used to qualitatively visualise the redeposited sediment up to a distance of 100 m from the source, and that sensors recording optical and acoustic backscatter are sensitive and adequate tools to monitor the horizontal and vertical dispersion of the generated sediment plume. Optical backscatter signals could be converted into absolute mass concentration of suspended sediment to provide quantitative data on sediment dispersion. Vertical profiles of acoustic backscatter recorded by current profilers provided qualitative insight into the vertical extent of the sediment plume. Our monitoring setup proved to be very useful for the monitoring of this small-scale experiment and can be seen as an exemplary strategy for monitoring studies of future, upscaled mining trials. We recommend that such larger trials include the use of AUVs for repeated seafloor imaging and water column plume mapping (optical and acoustical), as well as the use of in-situ particle size sensors and/or particle cameras to better constrain the effect of suspended particle aggregation on optical and acoustic backscatter signals.

Description: The increased fraction of first year ice (FYI) at the expense of old ice (second-year ice (SYI) and multi-year ice (MYI)) likely affects the permeability of the Arctic ice cover. This in turn influences the pathways of gases circulating therein and the exchange at interfaces with the atmosphere and ocean. We present sea ice temperature and salinity time series from different ice types relevant to temporal development of sea ice permeability and brine drainage efficiency from freeze-up in October to the onset of spring warming in May. Our study is based on a dataset collected during the Multidisciplinary drifting Observatory for the Study of Arctic Climate (MOSAiC) Expedition in 2019 and 2020. These physical properties were used to derive sea ice permeability and Rayleigh numbers. The main sites included FYI and SYI. The latter was composed of an upper layer of residual ice that had desalinated but survived the previous summer melt and became SYI. Below this ice a layer of new first-year ice formed. As the layer of new first-year ice has no direct contact with the atmosphere, we call it insulated first-year ice (IFYI). The residual/SYI-layer also contained refrozen melt ponds in some areas. During the freezing season, the residual/SYI-layer was consistently impermeable, acting as barrier for gas exchange between the atmosphere and ocean. While both FYI and SYI temperatures responded similarly to atmospheric warming events, SYI was more resilient to brine volume fraction changes because of its low salinity (〈 2). Furthermore, later bottom ice growth during spring warming was observed for SYI in comparison to FYI. The projected increase in the fraction of more permeable FYI in autumn and spring in the coming decades may favor gas exchange at the atmosphere-ice interface when sea ice acts as a source relative to the atmosphere. While the areal extent of old ice is decreasing, so is its thickness at the onset of freeze-up. Our study sets the foundation for studies on gas dynamics within the ice column and the gas exchange at both ice interfaces, i.e. with the atmosphere and the ocean.

Description: Measuring plankton and associated variables as part of ocean time-series stations has the potential to revolutionize our understanding of ocean biology and ecology and their ties to ocean biogeochemistry. It will open temporal scales (e.g., resolving diel cycles) not typically sampled as a function of depth. In this review we motivate the addition of biological measurements to time-series sites by detailing science questions they could help address, reviewing existing technology that could be deployed, and providing examples of time-series sites already deploying some of those technologies. We consider here the opportunities that exist through global coordination within the OceanSITES network for long-term (climate) time series station in the open ocean. Especially with respect to data management, global solutions are needed as these are critical to maximize the utility of such data. We conclude by providing recommendations for an implementation plan.

Description: Fast repetition rate fluorometry (FRRf) based on active chlorophyll fluorescence is a powerful, noninvasive tool for studying phytoplankton physiological status at high spatial and temporal resolution. The South China Sea (SCS) is one of the largest tropical–subtropical marginal seas in the world, which plays an important role in modulating regional carbon budget and climate. In this study, underway in situ FRRf measurements were carried out throughout the outer continental shelf of the northern SCS, the basin of the northern SCS, the cyclonic eddy influenced domain in the western SCS, and the basin of the southeastern SCS. Pronounced diurnal variability of FRRf-derived parameters were observed, characterized by a large midday depression and slight nocturnal depression of the maximum quantum yield of photosystem II ( F v /F m ) and a slight increase in the functional absorption cross-section of photosystem II photochemistry (σ PSII ) at noon. F v /F m at the surface was typically as low as 0.1–0.3 and exhibited higher values (~0.4) where internal waves occurred. The cyclonic eddy increased F v /F m slightly, implying that it had a limited impact on surface phytoplankton photophysiology. With proper interpretation, FRRf has been a powerful tool to assess the physiological status of phytoplankton in the sea and to correlate that to ocean dynamics in an unprecedented fine scale.

Description: In the Equatorial Atlantic nitrogen availability is assumed to control phytoplankton dynamics. However, in situ measurements of phytoplankton physiology and productivity are surprisingly sparse in comparison with the North Atlantic. In addition to the formation of the Equatorial cold tongue in the boreal summer, tropical instability waves (TIWs) and related short-term processes may locally cause episodic events of enhanced nutrient supply to the euphotic layer. Here, we assess changes in phytoplankton photophysiology in response to such episodic events as well as short-term nutrient addition experiments using a pair of custom-built fluorometers that measure chlorophyll a (Chl a) variable fluorescence and fluorescence lifetimes. The fluorometers were deployed during a transatlantic cruise along the Equator in the fall of 2019. We hypothesized that the Equatorial Atlantic is nitrogen-limited, with an increasing degree of limitation to the west where the cold tongue is not prominent, and that infrequent nitrate injection by TIW related processes are the primary source alleviating this limitation. We further hypothesized phytoplankton are well acclimated to the low levels of nitrogen, and once nitrogen is supplied, they can rapidly utilize it to stimulate growth and productivity. Across three TIW events encountered, we observed increased productivity and chlorophyll a concentration concurrent with a decreased photochemical conversion efficiency and overall photophysiological competency. Moreover, the observed decrease in photosynthetic turnover rates toward the western section suggested a 70% decrease in growth rates compared to their maximum values under nutrient-replete conditions. This decrease aligned with the increased growth rates observed following 24 h incubation with added nitrate in the western section. These results support our hypotheses that nitrogen is the limiting factor in the region and that phytoplankton are in a state of balanced growth, waiting to “body surf” waves of nutrients which fuel growth and productivity.

Description: Artificial upwelling of nutrient-rich waters and the corresponding boost in primary productivity harbor the potential to enhance marine fishery yields and strengthen the biological pump for sequestration of atmospheric CO2. There is increasing urgency to understand this technology as a “ocean-based solution” for counteracting two major challenges of the 21st century—climate change and overfishing. Yet, little is known about the actual efficacy and/or possible side effects of artificial upwelling. We conducted a large-scale off-shore mesocosm study (∼44 m3) in the oligotrophic waters of the Canary Islands to identify the community-level effects of artificial upwelling on a natural oligotrophic plankton community. Four upwelling intensities were simulated (approx. 1.5/3/5.7/10 μmol L–1 of nitrate plus phosphate and silicate) via two different upwelling modes (a singular deep-water pulse vs. recurring supply every 4 days) for 37 days. Here we present results on the response of net community production (NCP), metabolic balance and phytoplankton community composition (〈250 μm). Higher upwelling intensities yielded higher cumulative NCP. Following upwelling onset, the phytoplankton community became dominated by diatoms in all treatments, but other taxa such as Coccolithophores increased later in the experiment. The magnitude of effects on the metabolic balance scaled with the amount of added deep water, leading to (i) a balanced to net-heterotrophic system in the singular and (ii) a net-autotrophic system in the recurring upwelling treatments. Accordingly, the mode in which nutrients are supplied to an oligotrophic system plays a crucial role in the ecosystem response, with recurring upwelling leading to higher long-term positive NCP than singular upwelling. These results highlight the importance of empirically measured local responses to upwelling such as community structure and metabolism, with major implications for the potential employment of artificial upwelling as an ocean-based solution to generate (primary) production.

Description: Researchers have recognized the potential of enzymes and metabolic pathways hidden among the unseen majority of Earth’s microorganisms for decades now. Most of the microbes expected to colonize the seafloor and its subsurface are currently uncultured. Thus, their ability and contribution to element cycling remain enigmatic. Given that the seafloor covers ∼70% of our planet, this amounts to an uncalled potential of unrecognized metabolic properties and interconnections catalyzed by this microbial dark matter. Consequently, a tremendous black box awaits discovery of novel enzymes, catalytic abilities, and metabolic properties in one of the largest habitats on Earth. This mini review summarizes the current knowledge of cultivation-dependent and -independent techniques applied to seafloor habitats to unravel the role of the microbial dark matter. It highlights the great potential that combining microbiological and biogeochemical data from in situ experiments with molecular tools has for providing a holistic understanding of bio-geo-coupling in seafloor habitats and uses hydrothermal vent systems as a case example.

Description: Global estimates see river deltas and estuaries contributing about equally to CO2 and CH4 emissions as lakes and reservoirs, despite a factor 6 smaller surface area. Assessing the horizontal gradients in dissolved gas concentrations from large river reaches to connecting canals and wetland lakes remains a challenge in many deltaic systems. To elucidate the processes affecting local CO2 and CH4 concentrations in the Romanian part of the Danube Delta, we mapped dissolved O2, N2, He and Ar using a portable gas-equilibration membrane-inlet mass spectrometer (GE-MIMS), along with CO2, CH4, water temperature and conductivity. We measured the concentrations along the aquatic continuum from a small houseboat during two campaigns, in spring and autumn, to capture different hydrological and plant growth conditions. Delta-scale concentration patterns were comparably stable across seasons. Small connecting channels were highly influenced by the riparian wetland, which was strongest in the eastern part of the biosphere reserve. These sites represented the delta’s CO2 and CH4 hotspots and showed clear signs of excess air, i.e., supersaturation of dissolved noble gases with respect to air-saturated water. As the adjacent wetland was permanently inundated, this signal was likely caused by root aeration of Phragmites australis, as opposed to traditional excess air formation via water table fluctuations in the unsaturated zone. The special vegetation setting with reed growing on floating peat coincided with the highest CO2 and CH4 concentrations (〉700 μmol/L CO2 and 13 μmol/L CH4, respectively) observed in an adjacent channel. Shallow lakes, on the other hand, were major sites of photosynthetic production with O2 oversaturation reaching up to 150% in spring. The observed deficit in non-reactive gases (He, Ar and N2) indicated that the lakes were affected by O2 ebullition from macrophytes. According to our estimations, this ebullitive flux decreased O2 concentrations by up to 2 mg/L. This study highlights the effect of plant-mediated gas transfer on dissolved gas concentrations and supports recent studies stressing the need to account for ebullitive gas exchange when assessing metabolism parameters from O2 in shallow, productive settings.

Description: Submarine hydrothermal systems along active volcanic ridges and arcs are highly dynamic, responding to both oceanographic (e.g., currents, tides) and deep-seated geological forcing (e.g., magma eruption, seismicity, hydrothermalism, and crustal deformation, etc.). In particular, volcanic and hydrothermal activity may also pose profoundly negative societal impacts (tsunamis, the release of climate-relevant gases and toxic metal(loid)s). These risks are particularly significant in shallow (〈1000m) coastal environments, as demonstrated by the January 2022 submarine paroxysmal eruption by the Hunga Tonga-Hunga Ha’apai Volcano that destroyed part of the island, and the October 2011 submarine eruption of El Hierro (Canary Islands) that caused vigorous upwelling, floating lava bombs, and natural seawater acidification. Volcanic hazards may be posed by the Kolumbo submarine volcano, which is part of the subduction-related Hellenic Volcanic Arc at the intersection between the Eurasian and African tectonic plates. There, the Kolumbo submarine volcano, 7 km NE of Santorini and part of Santorini’s volcanic complex, hosts an active hydrothermal vent field (HVF) on its crater floor (~500m b.s.l.), which degasses boiling CO 2 –dominated fluids at high temperatures (~265°C) with a clear mantle signature. Kolumbo’s HVF hosts actively forming seafloor massive sulfide deposits with high contents of potentially toxic, volatile metal(loid)s (As, Sb, Pb, Ag, Hg, and Tl). The proximity to highly populated/tourist areas at Santorini poses significant risks. However, we have limited knowledge of the potential impacts of this type of magmatic and hydrothermal activity, including those from magmatic gases and seismicity. To better evaluate such risks the activity of the submarine system must be continuously monitored with multidisciplinary and high resolution instrumentation as part of an in-situ observatory supported by discrete sampling and measurements. This paper is a design study that describes a new long-term seafloor observatory that will be installed within the Kolumbo volcano, including cutting-edge and innovative marine-technology that integrates hyperspectral imaging, temperature sensors, a radiation spectrometer, fluid/gas samplers, and pressure gauges. These instruments will be integrated into a hazard monitoring platform aimed at identifying the precursors of potentially disastrous explosive volcanic eruptions, earthquakes, landslides of the hydrothermally weakened volcanic edifice and the release of potentially toxic elements into the water column.

Description: Deep coral-dominated communities play paramount roles in benthic environments by increasing their complexity and biodiversity. Coral-associated microbes are crucial to maintain fitness and homeostasis at the holobiont level. However, deep-sea coral biology and their associated microbiomes remain largely understudied, and less from remote and abyssal environments such as those in the Clarion-Clipperton Fracture Zone (CCZ) in the tropical Northeast (NE) Pacific Ocean. Here, we study microbial-associated communities of abyssal gorgonian corals and anemones (〉4,000 m depth) in the CCZ; an area harboring the largest known global reserve of polymetallic nodules that are commercially interesting for the deep-sea nodule mining. Coral samples (n = 25) belonged to Isididae and Primnoidae families, while anemones (n = 4) to Actinostolidae family. Significant differences in bacterial community compositions were obtained between these three families, despite sharing similar habitats. Anemones harbored bacterial microbiomes composed mainly of Hyphomicrobiaceae, Parvibaculales, and Pelagibius members. Core microbiomes of corals were mainly dominated by different Spongiibacteraceae and Terasakiellaceae bacterial members, depending on corals’ taxonomy. Moreover, the predicted functional profiling suggests that deep-sea corals harbor bacterial communities that allow obtaining additional energy due to the scarce availability of nutrients. This study presents the first report of microbiomes associated with abyssal gorgonians and anemones and will serve as baseline data and crucial insights to evaluate and provide guidance on the impacts of deep-sea mining on these key abyssal communities.

Description: Assessing the status of ocean acidification across ocean and coastal waters requires standardized procedures at all levels of data collection, dissemination, and analysis. Standardized procedures for assuring quality and accessibility of ocean carbonate chemistry data are largely established, but a common set of best practices for ocean acidification trend analysis is needed to enable global time series comparisons, establish accurate records of change, and communicate the current status of ocean acidification within and outside the scientific community. Here we expand upon several published trend analysis techniques and package them into a set of best practices for assessing trends of ocean acidification time series. These best practices are best suited for time series capable of characterizing seasonal variability, typically those with sub-seasonal (ideally monthly or more frequent) data collection. Given ocean carbonate chemistry time series tend to be sparse and discontinuous, additional research is necessary to further advance these best practices to better address uncharacterized variability that can result from data discontinuities. This package of best practices and the associated open-source software for computing and reporting trends is aimed at helping expand the community of practice in ocean acidification trend analysis. A broad community of practice testing these and new techniques across different data sets will result in improvements and expansion of these best practices in the future.

Description: Climate change is expected to alter the intensity and frequency of upwelling in high productive coastal regions, thus impacting nutrient fluxes, primary productivity and consequently carbon cycling. However, it is unknown how these changes will impact the planktonic (phytoplankton and bacteria) community structure, which affects community respiration (CR) and hence the carbon available for sequestration or transfer to upper trophic levels. Here we present results from a 37-day mesocosm experiment where we examined the response of CR to nutrient additions by simulating upwelling events at different intensities (low, medium, high and extreme) and modes (singular and recurring additions). We also analysed the potential contribution of different plankton size classes and functional groups to CR. The trend in accumulated CR with respect to nutrient fertilisation (total nitrogen added during the experiment) was linear in the two modes. Microplankton (mostly diatoms) and nanoplankton (small flagellates) dominated under extreme upwelling intensities and high CR in both singular and recurring upwelling modes, explaining 〉65% of the observed variability in CR. In contrast, prokaryotic picoplankton (heterotrophic bacteria and autotrophic cyanobacteria) explained 〈43% of the variance in CR under the rest of the upwelling intensities and modes tested. Changes in planktonic community structure, while modulating CR variability, would regulate the metabolic balance of the ecosystem, shifting it towards net-heterotrophy when the community is dominated by small heterotrophs and to net-autotrophy when large autotrophs prevail; although depending on the mode in which nutrients are supplied to the system. This shift in the dominance of planktonic organism will hence affect not only CR but also carbon sequestration in upwelling regions

Description: Marine sedimentary archives are routinely used to reconstruct past environmental changes. In many cases, bioturbation and sedimentary mixing affect the proxy time-series and the age-depth relationship. While idealized models of bioturbation exist, they usually assume homogeneous mixing, thus that a single sample is representative for the sediment layer it is sampled from. However, it is largely unknown to which extent this assumption holds for sediments used for paleoclimate reconstructions. To shed light on 1) the age-depth relationship and its full uncertainty, 2) the magnitude of mixing processes affecting the downcore proxy variations, and 3) the representativity of the discrete sample for the sediment layer, we designed and performed a case study on South China Sea sediment material which was collected using a box corer and which covers the last glacial cycle. Using the radiocarbon content of foraminiferal tests as a tracer of time, we characterize the spatial age-heterogeneity of sediments in a three-dimensional setup. In total, 118 radiocarbon measurements were performed on defined small- and large-volume bulk samples ( ∼ 200 specimens each) to investigate the horizontal heterogeneity of the sediment. Additionally, replicated measurements on small numbers of specimens (10 × 5 specimens) were performed to assess the heterogeneity within a sample volume. Visual assessment of X-ray images and a quantitative assessment of the mixing strength show typical mixing from bioturbation corresponding to around 10 cm mixing depth. Notably, our 3D radiocarbon distribution reveals that the horizontal heterogeneity (up to 1,250 years), contributing to the age uncertainty, is several times larger than the typically assumed radiocarbon based age-model error (single errors up to 250 years). Furthermore, the assumption of a perfectly bioturbated layer with no mixing underneath is not met. Our analysis further demonstrates that the age-heterogeneity might be a function of sample size; smaller samples might contain single features from the incomplete mixing and are thus less representative than larger samples. We provide suggestions for future studies, optimal sampling strategies for quantitative paleoclimate reconstructions and realistic uncertainty in age models, as well as discuss possible implications for the interpretation of paleoclimate records.

Description: Hydrothermal vents are a source of many trace metals to the oceans. Compared to mid-ocean ridges, hydrothermal vent systems at arcs occur in shallower water depth and are much more diverse in fluid composition, resulting in highly variable water column trace metal concentrations. However, only few studies have focused on trace metal dynamics in hydrothermal plumes at volcanic arcs. During R/V Sonne cruise SO253 in 2016/2017, hydrothermal plumes from two hydrothermally active submarine volcanoes along the Kermadec arc in the Southwest Pacific Ocean were sampled: (1) Macauley, a magmatic dominated vent site located in water depths between 300 and 680 m, and (2) Brothers, located between 1,200 and 1,600 m water depth, where hydrothermalism influenced by water rock interactions and magmatically influenced vent sites occur near each other. Surface currents estimated from satellite-altimeter derived currents and direct measurements at the sites using lowered acoustic Doppler current profilers indicate the oceanic regime is dominated by mesoscale eddies. At both volcanoes, results indicated strong plumes of dissolved trace metals, notably Mn, Fe, Co, Ni, Cu, Zn, Cd, La, and Pb, some of which are essential micronutrients. Dissolved metal concentrations commonly decreased with distance from the vents, as to be expected, however, certain element/Fe ratios increased, suggesting a higher solubility of these elements and/or their stronger stabilization (e.g., for Zn compared to Fe). Our data indicate that at the magmatically influenced Macauley and Brothers cone sites, the transport of trace metals is strongly controlled by sulfide nanoparticles, while at the Brothers NW caldera wall site iron oxyhydroxides seem to dominate the trace metal transport over sulfides. Solution stabilization of trace metals by organic complexation appears to compete with particle adsorption processes. As well as extending the generally sparse data set for hydrothermal plumes at volcanic arc systems, our study presents the first data on several dissolved trace metals in the Macauley system, and extends the existing plume dataset of Brothers volcano. Our data further indicate that chemical signatures and processes at arc volcanoes are highly diverse, even on small scales.

Description: Understanding and sustainably managing complex environments such as marine ecosystems benefits from an integrated approach to ensure that information about all relevant components and their interactions at multiple and nested spatiotemporal scales are considered. This information is based on a wide range of ocean observations using different systems and approaches. An integrated approach thus requires effective collaboration between areas of expertise in order to improve coordination at each step of the ocean observing value chain, from the design and deployment of multi-platform observations to their analysis and the delivery of products, sometimes through data assimilation in numerical models. Despite significant advances over the last two decades in more cooperation across the ocean observing activities, this integrated approach has not yet been fully realized. The ocean observing system still suffers from organizational silos due to independent and often disconnected initiatives, the strong and sometimes destructive competition across disciplines and among scientists, and the absence of a well-established overall governance framework. Here, we address the need for enhanced organizational integration among all the actors of ocean observing, focusing on the occidental systems. We advocate for a major evolution in the way we collaborate, calling for transformative scientific, cultural, behavioral, and management changes. This is timely because we now have the scientific and technical capabilities as well as urgent societal and political drivers. The ambition of the United Nations Decade of Ocean Science for Sustainable Development (2021–2030) and the various efforts to grow a sustainable ocean economy and effective ocean protection efforts all require a more integrated approach to ocean observing. After analyzing the barriers that currently prevent this full integration within the occidental systems, we suggest nine approaches for breaking down the silos and promoting better coordination and sharing. These recommendations are related to the organizational framework, the ocean science culture, the system of recognition and rewards, the data management system, the ocean governance structure, and the ocean observing drivers and funding. These reflections are intended to provide food for thought for further dialogue between all parties involved and trigger concrete actions to foster a real transformational change in ocean observing

Description: Seabed methane gas emissions occur worldwide at cold seeps located along most continental margins. Fluxes of methane gas released from the seabed in the form of bubbles can be extremely variable even over short time intervals. Some factors controlling the variability are still poorly understood. Here, we report on the results of continuous long-term sonar monitoring of bubble emissions at a depth of 1,260 m on the Clayoquot Slope, northern Cascadia margin. With a total monitoring duration of 4 years and a sampling period of 1 h, this is by far the longest high temporal resolution monitoring of seabed methane gas release ever conducted. Our results provide evidence that the diurnal and semi-diurnal tides influence the timing of the onset and cessation of bubble emissions. However, gas emissions within the monitoring area are active more than 84% of the time, indicating that tides alone are not sufficient to make venting pause. We hypothesize that the gas fluxes are transient but generally sufficiently high to maintain ebullition independently of the tidally-induced bottom pressure variations. Results also show that the tides do not seem to modulate the vigor of active gas emissions.

Description: Enhanced weathering of mafic and ultra-mafic minerals has been suggested as a strategy for carbon dioxide removal (CDR) and a contribution to achieve a balance between global CO2 sources and sinks (net zero emission). This study was designed to assess CDR by dissolution of ultramafic sand (UMS) in artificial seawater (ASW). Fine grained UMS with an olivine content of ~75% was reacted in ASW for up to 134 days at 1 bar and 21.5–23.9°C. A decline in total alkalinity (TA) was observed over the course of the experiments. This unexpected result indicates that TA removal via precipitation of cation-rich authigenic phases exceeded the production of TA induced by olivine dissolution. The TA decline was accompanied by a decrease in dissolved inorganic carbon and Ca concentrations presumably induced by CaCO3 precipitation. Temporal changes in dissolved Si, Ca, Mg, and TA concentrations observed during the experiments were evaluated by a numerical model to identify secondary mineral phases and quantify rates of authigenic phase formation. The modeling indicates that CaCO3, FeOOH and a range of Mg-Si-phases were precipitated during the experiments. Chemical analysis of precipitates and reacted UMS surfaces confirmed that these authigenic phases accumulated in the batch reactors. Nickel released during olivine dissolution, a potential toxic element for certain organisms, was incorporated in the secondary phases and is thus not a suitable proxy for dissolution rates as proposed by earlier studies. The overall reaction stoichiometry derived from lab experiments was applied in a box model simulating atmospheric CO2 uptake in a continental shelf setting induced by olivine addition. The model results indicate that CO2 uptake is reduced by a factor of 5 due to secondary mineral formation and the buffering capacity of seawater. In comparable natural settings, olivine addition may thus be a less efficient CDR method than previously believed.

Description: The highest concentration of cold seep sites worldwide has been observed along convergent margins, where fluid migration through sedimentary sequences is enhanced by tectonic deformation and dewatering of marine sediments. In these regions, gas seeps support thriving chemosynthetic ecosystems increasing productivity and biodiversity along the margin. In this paper, we combine seismic reflection, multibeam and split-beam hydroacoustic data to identify, map and characterize five known sites of active gas seepage. The study area, on the southern Hikurangi Margin off the North Island of Aotearoa/New Zealand, is a well-established gas hydrate province and has widespread evidence for methane seepage. The combination of seismic and hydroacoustic data enable us to investigate the geological structures underlying the seep sites, the origin of the gas in the subsurface and the associated distribution of gas flares emanating from the seabed. Using multi-frequency split-beam echosounder (EK60) data we constrain the volume of gas released at the targeted seep sites that lie between 1,110 and 2,060 m deep. We estimate the total deep-water seeps in the study area emission between 8.66 and 27.21 × 10 6 kg of methane gas per year. Moreover, we extrpolate methane fluxes for the whole Hikurangi Margin based on an existing gas seep database, that range between 2.77 × 10 8 and 9.32 × 10 8 kg of methane released each year. These estimates can result in a potential decrease of regional pH of 0.015–0.166 relative to the background value of 7.962. This study provides the most quantitative assessment to date of total methane release on the Hikurangi Margin. The results have implications for understanding what drives variation in seafloor biological communities and ocean biogeochemistry in subduction margin cold seep sites.

Description: The products of eruptive and mass-wasting processes that built island arc volcanoes are better preserved in marine deposits than on land. Holes U1397A and U1399A drilled during IODP Expedition 340 provide a 1.5 Ma record of the volcanic history of Martinique. 14C dating and δ18O patterns are used to reconstitute the chronostratigraphy of tephra, volcaniclastic turbidites, and mass-wasting events (traced by debris avalanches, debrites, and duplication and deformation of pre-existing sediments), leading to a new volcanic history of Montagne Pelée and Pitons du Carbet volcanoes. The top 50 m of core U1397A provides a continuous high-resolution sedimentation record over the last ∼130 ka. The sedimentation record deeper than 50 m in core U1397A and in the whole core U1399A is discontinuous because of the numerous sliding and deformation events triggered by debris avalanches related to flank collapses. Three successive activity periods are identified since ∼190 ka: the “Old Pelée” until 50 ka, the “Grand Rivière” (50–20 ka), and the “Recent Pelée” (20 ka—present day). The first two periods have the highest volcanic deposition rates offshore but very little outcrop on land. The whole magmatic activity of Mt Pelée comprises silicic andesites, but mafic andesites were also emitted during the whole “Grand Rivière.” At ∼115 ka, a major flank collapse (“Le Prêcheur”) produced a debris avalanche and submarine landslide that affected sea floor sediments by erosion and deformation up to ∼70 km from the shore. The Pitons du Carbet volcano was active from 1.2 Ma to 260 ka with numerous large flank collapses at a mean rate of 1 event every 100 ka. The average deposition rate of tephra fall offshore is much less than that at Mt Pelée. Our data show that correlations between the timing of large landslides or emission of mafic magmas and rapid sea level rise or lowstands suggested by previous studies are not systematic. The reconstituted chronostratigraphy of cores U1397A and U1399A provides the framework necessary for further studies of the magma petrology and production rates and timing of the mechanisms triggering flank collapses and related submarine landslides of Mt Pelée and Pitons du Carbet.

Description: The occurrence of mantle-derived peridotite xenoliths in phonolitic melts is a rare phenomenon, and is commonly ascribed to a mantle origin of the phonolite. The alternative possibility, that xenoliths are transported into evolving phonolite melts by mafic magmas, has received little attention. A unique tephriphonolite lava with phonolitic groundmass composition, from the active Cumbre Vieja volcano of La Palma (Canary Islands), allows to test these models. The lava contains abundant inclusions that represent the island’s major xenolith types: kaersutite-dominated cumulates, gabbros from the lower oceanic crust, and peridotites from the mantle. Our petrological investigations indicate that the tephriphonolite magma contained 3–4 wt% H 2 O and was stored in the lower crust at around 250–350 MPa and 900–950°C, at oxidized conditions (∆NNO of 2–3). The peridotite xenoliths are mantled by complex polyphase selvages, with adjacent up to 1.6 mm wide zonations where olivine compositions change from Fo 78-86 at the selvage contact to Fo 89-91 inside the xenoliths. We carried out diffusion modelling for Fe-Mg exchange and found that the peridotites had contact with intermediate to evolved alkaline melts over decades to centuries. This timescale is comparable to that inferred for basanite-hosted peridotite xenoliths from Cumbre Vieja. The following model is proposed: differentiation of evolved melts occurs in a magma accumulation zone in the lowermost oceanic crust beneath La Palma. The evolving melts receive periodic recharge by mantle-derived mafic magmas at intervals on the order of decades to a few centuries, comparable to historic eruption recurrences (80 years on average). Some of these recharge pulses carry mantle peridotite fragments that become deposited in the accumulation zone. Thus, these xenoliths do not reflect formation of the evolved melts in the mantle. Final ascent of the tephriphonolite was triggered by magma recharge some weeks before its eruption, resulting in entrainment and thorough mingling of a mixed xenolith population (cumulates, oceanic crust gabbros, peridotites). We infer that formation of phonolites in the lower crust beneath oceanic island volcanoes, and subsequent eruption, requires a balance between rates and volumes of magma recharge pulses and of eruptive events.

Description: The simultaneous limitation of productivity by two or more nutrients, commonly referred to as nutrient co-limitation, affects microbial communities throughout the marine environment and is of profound importance because of its impacts on various biogeochemical cycles. Multiple types of co-limitation have been described, enabling distinctions based on the hypothesized mechanisms of co-limitation at a biochemical level. These definitions usually pertain to individuals and do not explicitly, or even implicitly, consider complex ecological dynamics found within a microbial community. However, limiting and co-limiting nutrients can be produced in situ by a subset of microbial community members, suggesting that interactions within communities can underpin co-limitation. To address this, we propose a new category of nutrient co-limitation, community interaction co-limitation (CIC). During CIC, one part of the community is limited by one nutrient, which results in the insufficient production or transformation of a biologically produced nutrient that is required by another part of the community, often primary producers. Using cobalamin (vitamin B12) and nitrogen fixation as our models, we outline three different ways CIC can arise based on current literature and discuss CIC’s role in biogeochemical cycles. Accounting for the inherent and complex roles microbial community interactions play in generating this type of co-limitation requires an expanded toolset – beyond the traditional approaches used to identify and study other types of co-limitation. We propose incorporating processes and theories well-known in microbial ecology and evolution to provide meaningful insight into the controls of community-based feedback loops and mechanisms that give rise to CIC in the environment. Finally, we highlight the data gaps that limit our understanding of CIC mechanisms and suggest methods to overcome these and further identify causes and consequences of CIC. By providing this framework for understanding and identifying CIC, we enable systematic examination of the impacts this co-limitation can have on current and future marine biogeochemical processes.

Description: Regional climate variability in the tropical Atlantic, from interannual to decadal time scales, is inevitably connected to changes in the strength and position of the individual components of the tropical current system with impacts on societally relevant climate hazards such as anomalous rainfall or droughts over the surrounding continents (Bourlès et al., 2019; Foltz et al., 2019). Furthermore, the lateral supply of dissolved oxygen in the tropical Atlantic upper-ocean is closely linked to the zonal current bands (Brandt et al., 2008; Brandt et al., 2012; Burmeister et al., 2020) and especially to the Equatorial Undercurrent (EUC) and its long-term variations with potential implications for regional marine ecosystems (Brandt et al., 2021). The eastward flowing EUC is located between 70 to 200 m depth and forms one of the strongest tropical currents with maximum velocities of up to 1 m s-1 and maximum variability on seasonal time scales (Brandt et al., 2014; Johns et al., 2014). In the intermediate to deep equatorial Atlantic, variability on longer time scales is mainly governed by alternating, vertically-stacked, zonal currents (equatorial deep jets (EDJs); Johnson and Zhang, 2003). At a fixed location, the phases of these jets are propagating downward with time, implying that parts of their energy must propagate upward towards the surface (Brandt et al., 2011). In fact, a pronounced interannual cycle of about 4.5 years, that is associated with EDJs, is projected onto surface parameters such as sea surface temperature or precipitation (Brandt et al., 2011) further demonstrating the importance of understanding equatorial circulation variability and its role in tropical climate variability. While variability in the zonal velocity component on the equator is focused on seasonal to interannual time scales (Brandt et al., 2016; Claus et al., 2016; Kopte et al., 2018), meridional velocity fluctuations dominate the intraseasonal period range (20 to 50 days) due to the presence and passage of westward propagating Tropical Instability Waves (TIWs; Grodsky et al., 2005; Bunge et al., 2007; Wenegrat and McPhaden, 2015; Tuchen et al., 2018; Specht et al., 2021). In general, intraseasonal variability in the central equatorial Atlantic is mainly attributed to TIWs in the upper ocean (Athie and Marin, 2008), while intraseasonal variability in the deep ocean is associated with the signature of equatorial Yanai waves (Ascani et al., 2015; Tuchen et al., 2018, Körner et al., 2022). The observed and modelled interaction between intraseasonal equatorial waves and the aforementioned EDJs was found to maintain the deep equatorial circulation against dissipation (Greatbatch et al., 2018; Bastin et al., 2020) pointing toward the importance of intraseasonal variability for equatorial ocean dynamics. These findings are largely based on, or underpinned by a unique and steadily expanding data set of current velocity observations in the central equatorial Atlantic Ocean. Since 2001, current velocities have been measured almost continuously as part of a multilateral collaboration, the Prediction and Research Moored Array in the Tropical Atlantic (PIRATA), that regularly services a moored observatory located at 0°N/23°W (Bourlès et al., 2019). The significance of this data set is characterized by the length of the time series and by the full-depth coverage of current velocity observations which allow for a detailed analysis of both upper-ocean and deep-ocean dynamics on a wide range of time scales and frequencies. For instance, it enables the decomposition of the current velocity time series into vertical modes pointing toward the existence of resonant basin modes and identifying different sources of deep intraseasonal variability (Brandt et al., 2016; Claus et al., 2016; Greatbatch et al., 2018; Tuchen et al., 2018, Körner et al. under review). Here, we present 20 years of full-depth current velocity observations at 0°N/23°W. The aim of this study is to provide the scientific community with a publicly available reference data set that could be used in manifold ways, including, for instance, the validation of ocean models or reanalysis products.

Description: Marine Heatwaves (MHWs) are ocean extreme events, characterized by anomalously high temperatures, which can have significant ecological impacts. The Northeast U.S. continental shelf is of great economical importance as it is home to a highly productive ecosystem. Local warming rates exceed the global average and the region experienced multiple MHWs in the last decade with severe consequences for regional fisheries. Due to the lack of subsurface observations, the depth-extent of MHWs is not well-known, which hampers the assessment of impacts on pelagic and benthic ecosystems. This study utilizes a global ocean circulation model with a high-resolution (1/20°) nest in the Atlantic to investigate the depth structure of MHWs and associated drivers on the Northeast U.S. continental shelf. It is shown that MHWs exhibit varying spatial extents, with some only occurring at depth. The highest intensities are found around 100 m depth with temperatures exceeding the climatological mean by up to 7°C, while surface intensities are typically smaller (around 3°C). Distinct vertical structures are associated with different spatial MHW patterns and drivers. Investigation of the co-variability of temperature and salinity reveals that over 80% of MHWs at depth (〉50 m) coincide with extreme salinity anomalies. Two case studies provide insight into opposing MHW patterns at the surface and at depth, being forced by anomalous air-sea heat fluxes and Gulf Stream warm core ring interaction, respectively. The results highlight the importance of local ocean dynamics and the need to realistically represent them in climate models.

Description: Ecosystem models need to capture biodiversity, because it is a fundamental determinant of food web dynamics and consequently of the cycling of energy and matter in ecosystems. In oceanic food webs, the plankton compartment encompasses by far most of the biomass and diversity. Therefore, capturing plankton diversity is paramount for marine ecosystem modelling. In recent years, many models have been developed, each representing different aspects of plankton diversity, but a systematic comparison remains lacking. Here we present established modelling approaches to study plankton ecology and diversity, discussing the limitations and strengths of each approach. We emphasize their different spatial and temporal resolutions and consider the potential of these approaches as tools to address societal challenges. Finally, we make suggestions as to how better integration of field and experimental data with modelling could advance understanding of both plankton biodiversity specifically and more broadly the response of marine ecosystems to environmental change, including climate change.

Description: The Norway lobster, Nephrops norvegicus, supports a key European fishery. Stock assessments for this species are mostly based on trawling and UnderWater TeleVision (UWTV) surveys. However, N. norvegicus are burrowing organisms and these survey methods are unable to sample or observe individuals in their burrows. To account for this, UWTV surveys generally assume that “1 burrow system = 1 animal”, due to the territorial behavior of N. norvegicus. Nevertheless, this assumption still requires in-situ validation. Here, we outline how to improve the accuracy of current stock assessments for N. norvegicus with novel ecological monitoring technologies, including: robotic fixed and mobile camera-platforms, telemetry, environmental DNA (eDNA), and Artificial Intelligence (AI). First, we outline the present status and threat for overexploitation in N. norvegicus stocks. Then, we discuss how the burrowing behavior of N. norvegicus biases current stock assessment methods. We propose that state-of-the-art stationary and mobile robotic platforms endowed with innovative sensors and complemented with AI tools could be used to count both animals and burrows systems in-situ, as well as to provide key insights into burrowing behavior. Next, we illustrate how multiparametric monitoring can be incorporated into assessments of physiology and burrowing behavior. Finally, we develop a flowchart for the appropriate treatment of multiparametric biological and environmental data required to improve current stock assessment methods.

Description: This paper presents the results of a study on the response of living benthic foraminifera to progressing environmental successions in a cold-seep ecosystem. Sediment samples were collected from Vestnesa Ridge (79°N, Fram Strait) at ~1200 m water depth. The distribution of live (Rose Bengal-stained) foraminifera were analyzed in the upper sediment layers in relation to pore water biogeochemical data together with the distribution of sulfur-bacterial mats and Siboglinidae tubeworms. At methane cold seeps, the process of environmental succession is strongly connected to the duration and strength of methane seepage and the intensity of methane-related biological processes, e.g, aerobic and anaerobic oxidation of methane (MOx and AOM, respectively). The results show that the distribution patterns of benthic foraminifera change according to the progressing environmental succession. The benthic foraminifera seemed to thrive in sediments with a moderate activity of seepage, dominated by MOx, i.e, at an early stage of seepage or when seepage decreases at a late stage of the succession. Species composition of the foraminiferal fauna under these conditions was similar to the control sites (outside of pockmarks with no seepage); the dominant species being Melonis barleeanus and Cassidulina neoteretis. In sediments with strong seepage and high AOM activity, the hostile environmental conditions due to the presence of toxic sulfide caused a reduction in the foraminiferal population, and samples were almost barren of foraminifera. In environments of moderate methane seepage, the presence of chemosynthetic Siboglinidae tube worms potentially support communities of the epibenthic species Cibicidoides wuellerstorfi. Despite the very different environmental conditions, the foraminiferal assemblages were very similar (or nearly absent). Therefore, the foraminiferal faunas cannot be used as exclusive indicators of past strength of methane seepage in palaeoceanographic interpretations.

Description: Sponges host dense and diverse communities of microbes (known as the microbiome) beneficial for the host nutrition and defense. Symbionts in turn receive shelter and metabolites from the sponge host, making their relationship beneficial for both partners. Given that sponge-microbes associations are fundamental for the survival of both, especially the sponge, such relationship is maintained through their life and even passed on to the future generations. In many organisms, the microbiome has profound effects on the development of the host, but the influence of the microbiome on the reproductive and developmental pathways of the sponges are less understood. In sponges, microbes are passed on to oocytes, sperm, embryos, and larvae (known as vertical transmission), using a variety of methods that include direct uptake from the mesohyl through phagocytosis by oocytes to indirect transmission to the oocyte by nurse cells. Such microbes can remain in the reproductive elements untouched, for transfer to offspring, or can be digested to make the yolky nutrient reserves of oocytes and larvae. When and how those decisions are made are fundamentally unanswered questions in sponge reproduction. Here we review the diversity of vertical transmission modes existent in the entire phylum Porifera through detailed imaging using electron microscopy, available metabarcoding data from reproductive elements, and macroevolutionary patterns associated to phylogenetic constraints. Additionally, we examine the fidelity of this vertical transmission and possible reasons for the observed variability in some developmental stages. Our current understanding in marine sponges, however, is that the adult microbial community is established by a combination of both vertical and horizontal (acquisition from the surrounding environment in each new generation) transmission processes, although the extent in which each mode shapes the adult microbiome still remains to be determined. We also assessed the fundamental role of filtration, the cellular structures for acquiring external microbes, and the role of the host immune system, that ultimately shapes the stable communities of prokaryotes observed in adult sponges.

Description: In the face of climate change there is a need to reduce atmospheric CO2 concentrations. Artificial upwelling of nutrient-rich deep waters has been proposed as a method to enhance the biological carbon pump in oligotrophic oceanic regions in order to increase carbon sequestration. Here we examine the effect of different artificial upwelling intensities and modes (single pulse versus recurring pulses) on the dynamics of the dissolved organic matter pool (DOM). We introduced nutrient-rich deep water to large scale mesocosms (~44 m3) in the oligotrophic subtropical North Atlantic and found that artificial upwelling strongly increased DOM concentrations and changed its characteristics. The magnitude of the observed changes was related to the upwelling intensity: more intense treatments led to higher accumulation of dissolved organic carbon (〉70 μM of excess DOC over ambient waters for the most intense) and to comparatively stronger changes in DOM characteristics (increased proportions of chromophoric DOM (CDOM) and humic-like fluorescent DOM), suggesting a transformation of the DOM pool at the molecular level. Moreover, the single upwelling pulse resulted in higher CDOM quantities with higher molecular weight than the recurring upwelling mode. Together, our results indicate that under artificial upwelling, large DOM pools may accumulate in the surface ocean without being remineralized in the short-term. Possible reasons for this persistence could be a combination of the molecular diversification of DOM due to microbial reworking, nutrient limitation and reduced metabolic capabilities of the prokaryotic communities within the mesocosms. Our study demonstrates the importance of the DOC pool when assessing the carbon sequestration potential of artificial upwelling.

Description: Legal requirement in Europe asks for Ecosystem-Based Fisheries Management (EBFM) in European seas, including consideration of trophic interactions and minimization of negative impacts of fishing on food webs and ecosystem functioning. This study presents the first mass-balanced ecosystem model focused on the western Baltic Sea (WBS). Results show that heavy fishing pressure exerted on the WBS has forced top predators such as harbour porpoise and cod to cover their dietary needs by shifting from forage fish to other prey or find food outside of the model area. The model was then developed to explore the dynamics of four future fishery scenarios: (1) business as usual (BAU), (2) maximum sustainable fishing (F = FMSY), (3) half of FMSY, and (4) EBFM with F = 0.5 FMSY for forage fish and F = 0.8 FMSY for other fish. Simulations show that BAU would perpetuate low catches from depleted stocks with a high risk of extinction for harbour porpoise. In contrast, the EBFM scenario would allow the recovery of harbour porpoise, forage fish and cod with increases in catch of herring and cod. EBFM promotes ecosystem resilience to eutrophication and ocean warming, and through the rebuilding of commercial stocks increases by more than three times carbon sequestration compared to BAU. The model provides an interrelated assessment of trophic guilds in the WBS, as required by European law to assess whether European seas are in good environmental status.

Description: Artificial upwelling brings nutrient-rich deep water to the sun-lit surface to boost fisheries or carbon sequestration. Deep water sources under consideration range widely in inorganic silicon (Si) relative to nitrogen (N). Yet, little is known about how such differences in nutrient composition may influence the effectiveness of the fertilization. Si is essential primarily for diatoms that may increase food web and export efficiency via their large size and ballasting mineral shells, respectively. With a month-long mesocosm study in the subtropical North Atlantic, we tested the biological response to artificial upwelling with varying Si:N ratios (0.07-1.33). Community biomass increased 10-fold across all mesocosms, indicating that basic bloom dynamics were upheld despite the wide range in nutrient composition. Key properties of these blooms, however, were influenced by Si. Photosynthetic capacity and nutrient-use efficiency doubled from Si-poor to Si-rich upwelling, leading to C:N ratios as high as 17, well beyond Redfield. Si-rich upwelling also resulted in 6-fold higher diatom abundance and mineralized Si and a corresponding shift from smaller towards larger phytoplankton. The pronounced change in both plankton quantity (biomass) and quality (C:N ratio, size and mineral ballast) for trophic transfer and export underlines the pivotal role of Si in shaping the response of oligotrophic regions to upwelled nutrients. Our findings indicate a benefit of active Si management during artificial upwelling with the potential to optimize fisheries production and CO2 removal.

Description: Anthropogenic activities are modifying the oceanic environment rapidly and are causing ocean warming and deoxygenation, affecting biodiversity, productivity, and biogeochemical cycling. In coastal sediments, anaerobic organic matter degradation essentially fuels the production of hydrogen sulfide and methane. The release of these compounds from sediments is detrimental for the (local) environment and entails socio-economic consequences. Therefore, it is vital to understand which microbes catalyze the re-oxidation of these compounds under environmental dynamics, thereby mitigating their release to the water column. Here we use the seasonally dynamic Boknis Eck study site (SW Baltic Sea), where bottom waters annually fall hypoxic or anoxic after the summer months, to extrapolate how the microbial community and its activity reflects rising temperatures and deoxygenation. During October 2018, hallmarked by warmer bottom water and following a hypoxic event, modeled sulfide and methane production and consumption rates are higher than in March at lower temperatures and under fully oxic bottom water conditions. The microbial populations catalyzing sulfide and methane metabolisms are found in shallower sediment zones in October 2018 than in March 2019. DNA-and RNA profiling of sediments indicate a shift from primarily organotrophic to (autotrophic) sulfide oxidizing Bacteria, respectively. Previous studies using data collected over decades demonstrate rising temperatures, decreasing eutrophication, lower primary production and thus less fresh organic matter transported to the Boknis Eck sediments. Elevated temperatures are known to stimulate methanogenesis, anaerobic oxidation of methane, sulfate reduction and essentially microbial sulfide consumption, likely explaining the shift to a phylogenetically more diverse sulfide oxidizing community based on RNA.

Description: Introduction: Historically considered to be a single cosmopolitan species, the so called Octopus vulgaris species complex (OVSC) is now recognized to be a group of (at least) six cryptic species: O. americanus (in the west Atlantic), O. vulgaris (in the northeast Atlantic and Mediterranean Sea), O. aff. vulgaris (in the region of South Africa), O. tetricus (southeastern Oceania), O. sinensis (northwestern Pacific), and O. djinda (western Australia). The potentially different environmental preferences of this highly cryptic species complex may result in distinct consequences under future environmental conditions. Methods: The present study employed species distribution models (SDM) using MaxEnt to investigate potential changes in habitat suitability and geographical distribution of the OVSC in the future (i.e., 2050, and 2100), across four representative concentration pathway scenarios (RCP-2.6, 4.5, 6.0, and 8.5, CMIP5). Results: Differential responses were observed in the OVSC species analyzed. Specifically, O. vulgaris and O. tetricus exhibited a severe loss in distribution across their predicted range; O. americanus exhibited projected extirpation close to the equator, with limited expansion towards the poles; O. aff. vulgaris was projected to lose half of its current distribution; O. sinensis exhibited moderate losses, with projected increases in northern areas; and finally, O. djinda exhibited limited losses to its distribution. Except for O. sinensis, increasing RCP severity exacerbated changes in mean habitat suitability and projected distribution gains and losses. Discussion: Ultimately, this study provides information on the potential biogeographical effects of marine climate change on a key worldwide ecological and economic resource to further disentangle the effects over each OVSC species, with the goal of assisting toward the sustainable management of octopus species at the global scale.

Description: Ocean warming and acidification will be most pronounced in the Arctic. Both phenomena severely threat thecosome pteropods (holoplanktonic marine gastropods) by reducing their survival (warming) and causing dissolution of their aragonitic shell (acidification). Lipids, particularly phospholipids, play a major role in veligers and juveniles of the polar thecosome pteropod \textit{Limacina helicina} (Phipps 1774) comprising over two thirds of their total lipids. Membrane lipids (phospholipids) are important in temperature acclimation of ectotherms. Hence, we experimentally investigated ocean warming and acidification effects on total lipids, lipid classes and fatty acids of Arctic early-stage \textit{L. helicina}. Temperature and pCO\textsubscript{2} treatments chosen resembled Representative Concentration Pathway model scenarios for this century. We found a massive decrease of total lipids at elevated temperature and at the highest CO\textsubscript{2} concentration (1100 $\mu$atm) of the \textit{in situ} temperature. Clearly, temperature was the overriding factor. Total lipids were reduced by 47--70\%, mainly caused by a reduction of phospholipids by up to 60\%. Further, based on pH\textsubscript{T} development in the incubation water of pteropods during the experiment, some evidence exists for metabolic downregulation (shutdown?) in pteropods at high factor levels of temperature and pCO\textsubscript{2}. Consequently, cell differentiation and energy balance of early-stage larvae was probably severely compromised. Comparison of our experimental with 'wild' organisms suggests phospholipid reduction to values clearly outside natural variability. Based on the well-known significance of phospholipids for membranogenesis, early development, and reproduction, negative warming effects on such a basal metabolic function may be a much more immediate threat for pteropods than so far anticipated shell dissolution effects due to acidification.

Description: The recent rapid growth in aquaculture production reported by the United Nations Food and Agriculture Organization may have inadvertently generated what we denote here as aquaculture over-optimism. An extreme form of this is the notion that we need not worry about sustaining wild fish stocks because we can meet the global need through farming. Here we investigate whether the recent growth in aquaculture production can be maintained, and we compare aquaculture production projections with the future need for fish to find out whether aquaculture over-optimism can be justified. We show relevant evidence suggesting that aquaculture growth rates in all the cases studied have already reached their peak and have begun declining. Also, our results indicate that without wild fish, the world will face a fish food shortage of about 71 million tonnes annually by 2030, and the aquaculture production growth rate would have to be 3 times current average projected production by the FAO, the World Bank and the OECD in 2030. Finally, the current geographical distribution of farmed fish production suggests that even if aquaculture over-optimism is physically, economically, technically and ecologically feasible, its socio-economic cost to low-income coastal countries could be devastating.

Description: The use of cephalopod beaks in ecological and population dynamics studies has allowed major advances of our knowledge on the role of cephalopods in marine ecosystems in the last 60 years. Since the 1960’s, with the pioneering research by Malcolm Clarke and colleagues, cephalopod beaks (also named jaws or mandibles) have been described to species level and their measurements have been shown to be related to cephalopod body size and mass, which permitted important information to be obtained on numerous biological and ecological aspects of cephalopods in marine ecosystems. In the last decade, a range of new techniques has been applied to cephalopod beaks, permitting new kinds of insight into cephalopod biology and ecology. The workshop on cephalopod beaks of the Cephalopod International Advisory Council Conference (Sesimbra, Portugal) in 2022 aimed to review the most recent scientific developments in this field and to identify future challenges, particularly in relation to taxonomy, age, growth, chemical composition (i.e., DNA, proteomics, stable isotopes, trace elements) and physical (i.e., structural) analyses. In terms of taxonomy, new techniques (e.g., 3D geometric morphometrics) for identifying cephalopods from their beaks are being developed with promising results, although the need for experts and reference collections of cephalopod beaks will continue. The use of beak microstructure for age and growth studies has been validated. Stable isotope analyses on beaks have proven to be an excellent technique to get valuable information on the ecology of cephalopods (namely habitat and trophic position). Trace element analyses is also possible using beaks, where concentrations are significantly lower than in other tissues (e.g., muscle, digestive gland, gills). Extracting DNA from beaks was only possible in one study so far. Protein analyses can also be made using cephalopod beaks. Future challenges in research using cephalopod beaks are also discussed.

Description: It has long been hypothesized that the last glacial maximum (LGM) oversaw cold, arid, windy climates across southern Australia, and that these were driven by intensification and northward expansion of mid-latitude westerly circulation. Moreover, it was recently suggested that Australia experienced an extended LGM which began several millennia before the global peak. Aeolian sedimentary deposits provide key evidence for these hypotheses, and climate modelling an alternative means to test them. As yet, however, combined approaches to reconstructing glacial environments on the continent are scarce. Here we provide new evidence for westerly wind regimes across glacial southeastern Australia. We confirm active transverse lunette deposition at c. 29 ka and c. 23–19 ka in the semi-arid Willandra Lakes, and identify aeolian sand incursions to Spring Creek on the temperate Western Victorian Volcanic Plains from c. 29 ka. The Spring Creek deposits contain a surprising quantity of sand-sized quartz given the basalt setting, which we propose to be allochthonous and likely transported some distance. The site lies more than 50 km east and south of dunefields which were active at the same time and may have contributed sediment via long distance transport. We investigate the hypothesis for northward glacial expansion of westerly winds by combining our sediment records with aeolian particle transport simulations. We find that LGM near-surface winds were dominated by stronger, more focussed westerly air flow across southeastern Australia, compared with presently more diffuse wind regimes. Our results suggest stronger potential for LGM eastward distal sand transport onto the basalt plains, coeval with enhanced aeolian activity in the semi-arid Australian dunefields. Our combined reconstruction of aeolian deposition and trajectory modelling confirms the extended LGM hypothesis and indicates a northward migration of westerly winds over southeastern Australia during this period.

Description: Regrowth of body parts occurs in almost every phylum of the animal kingdom, but variation in this process across environmental, morphological, and behavioral gradients remains poorly understood. We examined regeneration patterns in feather stars – a group known for a wide range of morphologies and behaviors and up to a forty-fold difference in arm regeneration rates – and found that the variation in arm regeneration rates is best explained by swimming ability, not temperature, food supply, morphology (total number of arms and number of regenerating arms), or degree of injury. However, there were significant interactive effects of morphology on rates of regeneration of the main effect (swimming ability). Notably, swimmers grew up to three-fold faster than non-swimmers. The temperate feather star Florometra serratissima regenerated faster under warmer scenarios, but its rates fell within that of the tropical species suggesting temperature can account for intraspecific but not interspecific differences. We urge comparative molecular investigations of crinoid regeneration to identify the mechanisms responsible for the observed interspecific differences, and potentially address gaps in stem cell research.

Description: Low-salinity stress can severely affect the fitness of marine organisms. As desalination has been predicted for many coastal areas with ongoing climate change, it is crucial to gain more insight in mechanisms that constrain salinity acclimation ability. Low-salinity induced depletion of the organic osmolyte pool has been suggested to set a critical boundary in osmoconforming marine invertebrates. Whether inorganic ions also play a persistent role during low-salinity acclimation processes is currently inconclusive. We investigated the salinity tolerance of six marine invertebrate species following a four-week acclimation period around their low-salinity tolerance threshold. To obtain complete osmolyte budgets, we quantified organic and inorganic osmolytes and determined fitness proxies. Our experiments corroborated the importance of the organic osmolyte pool during low-salinity acclimation. Methylamines constituted a large portion of the organic osmolyte pool in molluscs, whereas echinoderms exclusively utilized free amino acids. Inorganic osmolytes were involved in long-term cellular osmoregulation in most species, thus are not just modulated with acute salinity stress. The organic osmolyte pool was not depleted at low salinities, whilst fitness was severely impacted. Instead, organic and inorganic osmolytes often stabilized at low-salinity. These findings suggest that low-salinity acclimation capacity cannot be simply predicted from organic osmolyte pool size. Rather, multiple parameters (i.e. osmolyte pools, net growth, water content and survival) are necessary to establish critical salinity ranges. However, a quantitative knowledge of cellular osmolyte systems is key to understand the evolution of euryhalinity and to characterize targets of selection during rapid adaptation to ongoing desalination.

Description: Artificial upwelling has been proposed as a means of enhancing oceanic CO2 sequestration and/or raising fishery yields through an increase in primary production in unproductive parts of the ocean. However, evidence of its efficacy, applicability and side effects is scarce. Here we present part of the results of a 37-day mesocosm study conducted in oligotrophic waters off the coast of Gran Canaria. The goal was to assess in situ the effects of artificial upwelling on the pelagic community. Upwelling was simulated via two modes: i) a singular deep-water pulse and ii) a recurring supply every 4 days; each mode at four different intensities defined by the total amount of nitrate added: approx. 1.5, 3, 5.7, and 11 µmol L-1. In this study we focus on the phytoplankton response through size-fractionated 14C primary production rates (PP), Chlorophyll a and biomass. We observed increases in PP, accumulated PP, Chlorophyll a and biomass that scaled linearly with upwelling intensity. Upwelling primarily benefitted larger phytoplankton size fractions, causing a shift from pico- and nano- to nano- and microphytoplankton. Recurring deep-water addition produced more biomass under higher upwelling intensities than a single pulse addition. It also reached significantly higher accumulated PP per unit of added nutrients and showed a stronger reduction in percentage extracellular release with increasing upwelling intensity. These results demonstrate that oligotrophic phytoplankton communities can effectively adjust to artificial upwelling regardless of upwelling intensity, but differently depending on the upwelling mode. Recurring supply of upwelled waters generated higher efficiencies in primary production and biomass build-up than a single pulse of the same volume and nutrient load.

Description: Ocean warming is one of the most important factors in shaping the spatial distribution and genetic biodiversity of marine organisms worldwide. The northwest Pacific has been broadly illustrated as an essential seaweed diversity hotspot. However, few studies have yet investigated in this region on whether and how past and ongoing climate warming impacted the distribution and genetic pools of coastal seaweeds. Here, we chose the invasive species Gracilaria vermiculophylla as a model, and identified multiple genetic lineages in the native range through genome-scale microsatellite genotyping. Subsequently, by reconstructing decadal trends of sea surface temperature (SST) change between 1978 and 2018, we found that SST in northern Japan and the East China Sea indeed increased broadly by 0.25-0.4°C/decade. The projections of species distribution models (SDMs) under different future climate change scenarios (RCP 2.6, RCP 4.5, RCP 6.0 and RCP 8.5) indicated that a unique genetic pool of G. vermiculophylla at its current southern range limit (i.e. the South China Sea) is at high risk of disappearance, and that the populations at its current northern range limit (i.e. in Hokkaido region) will undergo poleward expansions, particularly by the year 2100. Such responses, along with this species’ limited dispersal potential, may considerably alter the contemporary distribution and genetic composition of G. vermiculophylla in the northwest Pacific, and ultimately threaten ecological services provided by this habitat-forming species and other associated functional roles.

Description: Two lander-based devices, the Bubble-Box and GasQuant-II, were used to investigate the spatial and temporal variability and total gas flow rates of a seep area offshore Oregon, United States. The Bubble-Box is a stereo camera–equipped lander that records bubbles inside a rising corridor with 80 Hz, allowing for automated image analyses of bubble size distributions and rising speeds. GasQuant is a hydroacoustic lander using a horizontally oriented multibeam swath to record the backscatter intensity of bubble streams passing the swath plain. The experimental set up at the Astoria Canyon site at a water depth of about 500 m aimed at calibrating the hydroacoustic GasQuant data with the visual Bubble-Box data for a spatial and temporal flow rate quantification of the site. For about 90 h in total, both systems were deployed simultaneously and pressure and temperature data were recorded using a CTD as well. Detailed image analyses show a Gaussian-like bubble size distribution of bubbles with a radius of 0.6–6 mm (mean 2.5 mm, std. dev. 0.25 mm); this is very similar to other measurements reported in the literature. Rising speeds ranged from 15 to 37 cm/s between 1- and 5-mm bubble sizes and are thus, in parts, slightly faster than reported elsewhere. Bubble sizes and calculated flow rates are rather constant over time at the two monitored bubble streams. Flow rates of these individual bubble streams are in the range of 544–1,278 mm 3 /s. One Bubble-Box data set was used to calibrate the acoustic backscatter response of the GasQuant data, enabling us to calculate a flow rate of the ensonified seep area (∼1,700 m 2 ) that ranged from 4.98 to 8.33 L/min (5.38 × 10 6 to 9.01 × 10 6 CH 4 mol/year). Such flow rates are common for seep areas of similar size, and as such, this location is classified as a normally active seep area. For deriving these acoustically based flow rates, the detailed data pre-processing considered echogram gridding methods of the swath data and bubble responses at the respective water depth. The described method uses the inverse gas flow quantification approach and gives an in-depth example of the benefits of using acoustic and optical methods in tandem.

Description: During recent years, experimental ecology started to focus on regional to local environmental fluctuations in the context of global climate change. Among these, marine heatwaves can pose significant threats to marine organisms. Yet, experimental studies that include fluctuating thermal stress are rare, and if available often fail to base experimental treatments on available long-term environmental data. We evaluated 22-year high-resolution sea surface temperature data on the occurrence of heatwaves and cold-spells in a temperate coastal marine environment. The absence of a general warming trend in the data may in parts be responsible for a lack of changes in heatwave occurrences (frequency) and their traits (intensity, duration, and rate of change) over time. Yet, the retrieved traits for present-day heatwaves ensured most-natural treatment scenarios, enabling an experimental examination of the impacts of marine heatwaves and phases of recovery on an important temperate predator, the common sea star Asterias rubens. In a 68-days long experiment, we compared a 37- and a 28-days long heatwave with a treatment that consisted of three consecutive 12-days long heatwaves with 4 days of recovery in between. The heatwaves had an intensity of 4.6°C above climatological records, resulting in a maximum temperature of 23.25°C. We demonstrate that heatwaves decrease feeding and activity of A. rubens, with longer heatwaves having a more severe and lasting impact on overall feeding pressure (up to 99.7% decrease in feeding rate) and growth (up to 87% reduction in growth rate). Furthermore, heatwaves of similar overall mean temperature, but interrupted, had a minor impact compared to continuous heatwaves, and the impact diminished with repeated heatwave events. We experimentally demonstrated that mild heatwaves of today’s strength decrease the performance of A. rubens. However, this echinoderm may use naturally occurring short interruptions of thermal stress as recovery to persist in a changing and variable ocean. Thus, our results emphasize the significance of thermal fluctuations and especially, the succession and timing of heat-stress events

Description: Ocean acidification (OA) is a serious consequence of climate change with complex organism-to-ecosystem effects that have been observed through field observations but are mainly derived from experimental studies. Although OA trends and the resulting biological impacts are likely exacerbated in the semi-enclosed and highly populated Mediterranean Sea, some fundamental knowledge gaps still exist. These gaps are at tributed to both the uneven capacity for OA research that exists between Mediterranean countries, as well as to the subtle and long-term biological, physical and chemical interactions that define OA impacts. In this paper, we systematically analyzed the different aspects of OA research in the Mediterranean region based on two sources: the United Nation’s International Atomic Energy Agency’s (IAEA) Ocean Acidification International Coordination Center (OA-ICC) database, and an extensive survey. Our analysis shows that 1) there is an uneven geographic capacity in OA research, and illustrates that both the Algero-Provencal and Ionian sub-basins are currently the least studied Mediterranean areas, 2) the carbonate system is still poorly quantified in coastal zones, and long-term time-series are still sparse across the Mediterranean Sea, which is a challenge for studying its variability and assessing coastal OA trends, 3) the most studied groups of organisms are autotrophs (algae, phanerogams, phytoplankton), mollusks, and corals, while microbes, small mollusks (mainly pteropods), and sponges are among the least studied, 4) there is an overall paucity in socio-economic, paleontological, and modeling studies in the Mediterranean Sea, and 5) in spite of general resource availability and the agreement for improved and coordinated OA governance, there is a lack of consistent OA policies in the Mediterranean Sea. In addition to highlighting the current status, trends and gaps of OA research, this work also provides recommendations, based on both our literature assessment and a survey that targeted the Mediterranean OA scientific community. In light of the ongoing 2021-2030 United Nations Decade of Ocean Science for Sustainable Development, this work might provide a guideline to close gaps of knowledge in the Mediterranean OA research.

Description: In 1964, exploration drilling in the German Sector of the North Sea hit a gas pocket at ∼2900 m depth below the seafloor and triggered a blowout, which formed a 550 m-wide and up to 38 m deep seafloor crater now known as Figge Maar. Although seafloor craters formed by fluid flow are very common structures, little is known about their formation dynamics. Here, we present 2D reflection seismic, sediment echosounder, and multibeam echosounder data from three geoscientific surveys of the Figge Maar blowout crater, which are used to reconstruct its formation. Reflection seismic data support a scenario in which overpressured gas ascended first through the lower part of the borehole and then migrated along steeply inclined strata and faults towards the seafloor. The focused discharge of gas at the seafloor removed up to 4.8 Mt of sediments in the following weeks of vigorous venting. Eyewitness accounts document that the initial phase of crater formation was characterized by the eruptive expulsion of fluids and sediments cutting deep into the substrate. This was followed by a prolonged phase of sediment fluidization and redistribution widening the crater. After fluid discharge ceased, the Figge Maar acted as a sediment trap reducing the crater depth to ∼12 m relative to the surrounding seafloor in 2018, which corresponds to an average sedimentation rate of ∼22,000 m 3 /yr between 1995 and 2018. Hydroacoustic and geochemical data indicate that the Figge Maar nowadays emits primarily biogenic methane, predominantly during low tide. The formation of Figge Maar illustrates hazards related to the formation of secondary fluid pathways, which can bypass safety measures at the wellhead and are thus difficult to control.

Description: High quality pH measurements are required in estuarine and coastal waters to assess the impacts of anthropogenic atmospheric CO 2 emissions on the marine carbonate system, including the resulting decrease in pH. In addition, pH measurements are needed to determine impacts on carbonate chemistry of phytoplankton blooms and their breakdown, following enhanced anthropogenic nutrient inputs. The spectrophotometric pH technique provides high quality pH data in seawater, and is advantageous for long-term deployments as it is not prone to drift and does not require in situ calibration. In this study, a field application of a fully automated submersible spectrophotometric analyzer for high-resolution in situ pH measurements in dynamic estuarine and coastal waters is presented. A Lab-on-Chip (LOC) pH sensor was deployed from a pontoon in the inner Kiel Fjord, southwestern Baltic Sea, for a total period of 6 weeks. We present a time-series of in situ pH T (total pH scale) and ancillary data, with sensor validation using discretely collected samples for pH T and laboratory analysis. The difference between the sensor and laboratory analyses of discrete samples was within ±0.015 pH T unit, with a mean difference of 0.001 (n=65), demonstrating that the LOC sensor can provide stable and accurate pH T measurements over several weeks.

Description: The Youngest Toba Tuff (YTT) supereruption from Toba Caldera in Sumatra at ca. 74,000 years BP is the largest volcanic event recorded in the Pleistocene. Intriguingly, recent radioisotopic dating of the near antipodal Los Chocoyos (LCY) supereruption from the Atitlán caldera in Guatemala finds an identical age within uncertainties to that of YTT. This opens the question of whether these synchronous supereruptions may be a coincidence or could be a consequence of each other? Using the known eruptive record from the past 2 Myr, we find that the likelihood of having two near antipodal supereruptions (〉1,000 km3 tephra volume) within centuries (〈400 years), as suggested by volcanic proxies and annual counting layer chronology in the ice core records, is very small (0.086%), requiring a non-random cause and effect. Considering this analysis, we speculate that one potential physical mechanism that could explain the temporal relationship between these supereruptions is that seismic energy released during YTT eruption focused on the antipodal region, where concentrated stresses ultimately promoted the eruption of the perched LCY magma system (or vice versa). This supereruption “double-whammy” may thus be the more compelling source of the significant environmental impacts often attributed individually to the YTT supereruption. Improving the existing age information of YTT and LCY, and a better understanding of caldera collapse events will enable further testing of the hypothesis that synchronous supereruptions do not result by pure chance.

Description: The subterranean estuary (STE) has been historically defined in terms of the mixing of saline and fresh water, in an analogy to surface estuaries. However, redox gradients are also a defining characteristic of the STE and influence its role as a source or sink for metals in the environment. Approaching the STE from a redox-focused biogeochemical perspective (e.g., considering the role of microbial respiration and availability of organic matter) provides the ability to quantify drivers of metal transport across spatial and temporal scales. This study measured the groundwater composition of a shallow STE over 2 years and used multiple linear regression to characterize the influence of salinity and redox chemistry on the behavior of redox-sensitive metals (RSMs) including Mo, U, V, and Cr. Molybdenum and uranium were both supplied to the STE by surface water, but differed in their removal mechanisms and seasonal behavior. Molybdenum showed non-conservative removal by reaction with sulfide in all seasons. Sulfide concentrations at this site were consistently higher than required for quantitative reaction with Mo (10 µM sulfide), evidently leading to quantitative removal at the same depth regardless of season. In contrast, U appeared to depend directly on microbial activity for removal, and showed more extensive removal at shallower depths in summer. Both V and Cr were elevated in meteoric groundwater (2.5–297 nM and 2.6–236 nM, respectively), with higher endmember concentrations in summer. Both V and Cr also showed non-conservative addition within the STE relative to conservative mixing among the observed endmembers. The mobility of V and Cr in the STE, and therefore their supply to the coastal ocean, was controlled by the availability of dissolved organic matter and Fe, suggesting V and Cr were potentially complexed in the colloidal fraction. Complexation by different organic matter pools led to seasonal variations in V but greater interannual variability of Cr. These results reveal distinct behaviors of RSMs in response to seasonal biogeochemical processes that drive microbial activity, organic matter composition, and complexation by inorganic species.

Description: Effective data management plays a key role in oceanographic research as cruise-based data, collected from different laboratories and expeditions, are commonly compiled to investigate regional to global oceanographic processes. Here we describe new and updated best practice data standards for discrete chemical oceanographic observations, specifically those dealing with column header abbreviations, quality control flags, missing value indicators, and standardized calculation of certain properties. These data standards have been developed with the goals of improving the current practices of the scientific community and promoting their international usage. These guidelines are intended to standardize data files for data sharing and submission into permanent archives. They will facilitate future quality control and synthesis efforts and lead to better data interpretation. In turn, this will promote research in ocean biogeochemistry, such as studies of carbon cycling and ocean acidification, on regional to global scales. These best practice standards are not mandatory. Agencies, institutes, universities, or research vessels can continue using different data standards if it is important for them to maintain historical consistency. However, it is hoped that they will be adopted as widely as possible to facilitate consistency and to achieve the goals stated above.

Description: Considering the reported significant diazotrophic activities in open-ocean regions where primary production is strongly limited by phosphate, we explored the ability of diazotrophs to use other sources of phosphorus to alleviate the phosphate depletion. We tested the actual efficiency of the open-ocean, N2-fixer Crocosphaera watsonii to grow on organic phosphorus as the sole P source, and observed how the P source affects the cellular C, N, and P composition. We obtained equivalent growth efficiencies on AMP and DL-α-glycerophosphate as compared with identical cultures grown on phosphate, and survival of the population on phytic acid. Our results show that Crocosphaera cannot use all phosphomonoesters with the same efficiency, but it can grow without phosphate, provided that usable DOP and sufficient light energy are available. Also, results point out that organic phosphorus uptake is not proportional to alkaline phosphatase activity, demonstrating that the latter is not a suitable proxy to estimate DOP-based growth yields of organisms, whether in culture experiments or in the natural environment. The growth parameters obtained, as a function of the P source, will be critical to improve and calibrate mathematical models of diazotrophic growth and the distribution of nitrogen fixation in the global ocean.

Description: Fast repetition rate fluorometry (FRRf) allows for rapid non-destructive assessment of phytoplankton photophysiology in situ yet has rarely been applied to Trichodesmium. This gap reflects long-standing concerns that Trichodesmium (and other cyanobacteria) contain pigments that are less effective at absorbing blue light which is often used as the sole excitation source in FRR fluorometers—potentially leading to underestimation of key fluorescence parameters. In this study, we use a multi-excitation FRR fluorometer (equipped with blue, green, and orange LEDs) to investigate photophysiological variability in Trichodesmium assemblages from two sites. Using a multi-LED measurement protocol (447+519+634 nm combined), we assessed maximum photochemical efficiency (Fv/Fm), functional absorption cross section of PSII (σPSII), and electron transport rates (ETRs) for Trichodesmium assemblages in both the Northwest Pacific (NWP) and North Indian Ocean in the vicinity of Sri Lanka (NIO-SL). Evaluating fluorometer performance, we showed that use of a multi-LED measuring protocol yields a significant increase of Fv/Fm for Trichodesmium compared to blue-only excitation. We found distinct photophysiological differences for Trichodesmium at both locations with higher average Fv/Fm as well as lower σPSII and non-photochemical quenching (NPQNSV) observed in the NWP compared to the NIO-SL (Kruskal–Wallis t-test df = 1, p 〈 0.05). Fluorescence light response curves (FLCs) further revealed differences in ETR response with a lower initial slope (αETR) and higher maximum electron turnover rate ((Formula presented.)) observed for Trichodesmium in the NWP compared to the NIO-SL, translating to a higher averaged light saturation EK (= (Formula presented.) /αETR) for cells at this location. Spatial variations in physiological parameters were both observed between and within regions, likely linked to nutrient supply and physiological stress. Finally, we applied an algorithm to estimate primary productivity of Trichodesmium using FRRf-derived fluorescence parameters, yielding an estimated carbon-fixation rate ranging from 7.8 to 21.1 mgC mg Chl-a–1 h–1 across this dataset. Overall, our findings demonstrate that capacity of multi-excitation FRRf to advance the application of Chl-a fluorescence techniques in phytoplankton assemblages dominated by cyanobacteria and reveals novel insight into environmental regulation of photoacclimation in natural Trichodesmium populations

Description: Removal of carbon dioxide from the atmosphere will be required over the next decades to achieve the Paris Agreement goal of limiting global warming to well below 2°C aiming at not exceeding 1.5°C. Technological and ecosystem-based options are considered for generating negative emissions through carbon dioxide removal (CDR) and several nations have already included these in their Long-Term Low Greenhouse Gas Emission Development Strategies. However, strategies for development, implementation, and upscaling of CDR options often remain vague. Considering the scale at which CDR deployment is envisioned in emission pathways for limiting global warming to 1.5°C, significant environmental, social, and institutional implications are to be expected and need to be included in national feasibility assessments of CDR options. Following a multi-disciplinary and comprehensive approach, we created a framework that considers the environmental, technological, economic, social, institutional, and systemic implications of upscaling CDR options. We propose the framework as a tool to help guide decision-relevant feasibility assessments of CDR options, as well as identify challenges and opportunities within the national context. As such, the framework can serve as a means to inform and support decision makers and stakeholders in the iterative science-policy process of determining the role of CDR options in national strategies of achieving net-zero carbon emissions.

Description: The effects of climate change (CC) on contaminants and their potential consequences to marine ecosystem services and human wellbeing are of paramount importance, as they pose overlapping risks. Here, we discuss how the interaction between CC and contaminants leads to poorly constrained impacts that affects the sensitivity of organisms to contamination leading to impaired ecosystem function, services and risk assessment evaluations. Climate drivers, such as ocean warming, ocean deoxygenation, changes in circulation, ocean acidification, and extreme events interact with trace metals, organic pollutants, excess nutrients, and radionuclides in a complex manner. Overall, the holistic consideration of the pollutants-climate change nexus has significant knowledge gaps, but will be important in understanding the fate, transport, speciation, bioavailability, toxicity, and inventories of contaminants. Greater focus on these uncertainties would facilitate improved predictions of future changes in the global biogeochemical cycling of contaminants and both human health and marine ecosystems.

Description: The air-sea gas transfer velocity (K-660) is typically assessed as a function of the 10-m neutral wind speed (U-10n), but there remains substantial uncertainty in this relationship. Here K-660 of CO2 derived with the eddy covariance (EC) technique from eight datasets (11 research cruises) are reevaluated with consistent consideration of solubility and Schmidt number and inclusion of the ocean cool skin effect. K-660 shows an approximately linear dependence with the friction velocity (u*) in moderate winds, with an overall relative standard deviation (relative standard error) of about 20% (7%). The largest relative uncertainty in K-660 occurs at low wind speeds, while the largest absolute uncertainty in K-660 occurs at high wind speeds. There is an apparent regional variation in the steepness of the K-660-u* relationships: North Atlantic 〉= Southern Ocean 〉 other regions (Arctic, Tropics). Accounting for sea state helps to collapse some of this regional variability in K-660 using the wave Reynolds number in very large seas and the mean squared slope of the waves in small to moderate seas. The grand average of EC-derived K-660 ( - 1.47 + 76.67 u * + 20.48 u *(2) o r 0.36 + 1.203 U-10n + 0.167 U (2)(10n) ) is similar at moderate to high winds to widely used dual tracer-based K-660 parametrization, but consistently exceeds the dual tracer estimate in low winds, possibly in part due to the chemical enhancement in air-sea CO2 exchange. Combining the grand average of EC-derived K-660 with the global distribution of wind speed yields a global average transfer velocity that is comparable with the global radiocarbon (C-14) disequilibrium, but is similar to 20% higher than what is implied by dual tracer parametrizations. This analysis suggests that CO2 fluxes computed using a U-10n (2) dependence with zero intercept (e.g., dual tracer) are likely underestimated at relatively low wind speeds.

Description: High alkalinity values observed in coastal seas promote the uptake of CO2 from the atmosphere. However, the alkalinity budget of coastal areas and marginal seas is poorly understood, even though some of the recently observed alkalinity enhancement can be ascribed to riverine fluxes and anaerobic processes in shelf sediments. Here, we investigate the alkalinity budget of the Baltic Sea to identify previously unrecognized alkalinity sources. We quantify the generation of alkalinity and dissolved calcium (Ca) in this marginal sea applying simple mass balance calculations. Using this approach, we identify alkalinity and Ca sources of approximately 324 Gmol yr-1 and 122 Gmol yr-1, respectively, that cannot be ascribed to the riverine input. The magnitude of the Ca source suggests that a major fraction of the excess alkalinity (244 Gmol yr-1) is induced by the dissolution of calcium carbonate (CaCO3). A review of available field data shows that carbonate-bearing rocks at the coast and the seabed of the Baltic Sea are rapidly eroded and may provide sufficient CaCO3 to close the Ca budget. Hence, dissolution of eroded CaCO3 is the most likely source for the Ca enrichment observed in Baltic Sea water. This hypothesis is supported by mass accumulation rates of sediments derived from radioisotope data that are evaluated to derive a basin-wide rate of mud to muddy sand accumulation at the bottom of the Baltic Sea. The resulting value (139 Tg yr-1) exceeds current estimates of riverine particle fluxes into the Baltic Sea by more than one order of magnitude and confirms that rates of till erosion are sufficiently high to account for the Ca and most of the alkalinity excess in Baltic Sea water. Finally, we show that deliberate addition of CaCO3 to sediments deposited in the Baltic Sea could neutralize significant amounts of CO2 and help to achieve net-zero greenhouse gas emissions in the Baltic region.

Description: Pollution of the marine environment is an emerging threat. Nowadays, engineered nanoparticles (〈100 nm) such as zinc, copper and silver are widely used as antimicrobial agents, therefore often present in daily-life products. Consequently, the demand and production of nanoparticles are expected to increase. Here, we specifically focus on silver nanoparticles (AgNP). Once released into the environment, AgNPs pose an obvious ecotoxicological risk, potentially affecting ecosystem structure and functioning. For instance, phytoplankton-derived exudates, rich in acidic polysaccharides and amino acids, can abiotically aggregate into microgels such as transparent exopolymer particles (TEP) and Coomassie stainable particles (CSP). Hence, microgels can bridge dissolved and particulate size fractions and facilitate aggregate formation with organic and mineral particles. Both physical and chemical properties make TEP and CSP attractive nutrient hotspots for heterotrophic bacterioplankton. Bacteria, in turn, utilize extracellular enzymes to access these carbon and nitrogen pools. However, knowledge about the mechanisms by which AgNPs might interact with and affect the biogeochemical cycling of TEP and CSP is still insufficient. Therefore, we conducted a mesocosm experiment in the Eastern Mediterranean Sea and investigated the effects of environmentally relevant concentrations of silver ions (Ag+) and AgNP on the properties of TEP and CSP (i.e., area and abundance) along with enzymatic activity measurements. Our results showed that cyanobacteria were likely the primary source of CSP in the ultra-oligotrophic Mediterranean Sea. Also, CSP contributed more to the microgel pool than TEP, as indicated by a strong relationship between CSP and heterotrophic microbial dynamics. While silver (i.e., Ag+ or AgNP) had overall only marginal effects, both species affected the relationships between cell-specific LAPase activity and CSP and cell-specific APase activity and phosphate levels. Thus, Ag+ and AgNP have the potential to regulate microgel dynamics. However, future studies are needed to derive a robust understanding of the effects of silver pollution on the coupling of microgel formation and degradation and the follow-on effect on biogeochemical cycles.

Description: Zooplankton plays a major role in ocean food webs and biogeochemical cycles, and provides major ecosystem services as a main driver of the biological carbon pump and in sustaining fish communities. Zooplankton is also sensitive to its environment and reacts to its changes. To better understand the importance of zooplankton, and to inform prognostic models that try to represent them, spatially-resolved biomass estimates of key plankton taxa are desirable. In this study we predict, for the first time, the global biomass distribution of 19 zooplankton taxa (1-50 mm Equivalent Spherical Diameter) using observations with the Underwater Vision Profiler 5, a quantitative in situ imaging instrument. After classification of 466,872 organisms from more than 3,549 profiles (0-500 m) obtained between 2008 and 2019 throughout the globe, we estimated their individual biovolumes and converted them to biomass using taxa-specific conversion factors. We then associated these biomass estimates with climatologies of environmental variables (temperature, salinity, oxygen, etc.), to build habitat models using boosted regression trees. The results reveal maximal zooplankton biomass values around 60 degrees N and 55 degrees S as well as minimal values around the oceanic gyres. An increased zooplankton biomass is also predicted for the equator. Global integrated biomass (0-500 m) was estimated at 0.403 PgC. It was largely dominated by Copepoda (35.7%, mostly in polar regions), followed by Eumalacostraca (26.6%) Rhizaria (16.4%, mostly in the intertropical convergence zone). The machine learning approach used here is sensitive to the size of the training set and generates reliable predictions for abundant groups such as Copepoda (R2 approximate to 20-66%) but not for rare ones (Ctenophora, Cnidaria, R2 〈 5%). Still, this study offers a first protocol to estimate global, spatially resolved zooplankton biomass and community composition from in situ imaging observations of individual organisms. The underlying dataset covers a period of 10 years while approaches that rely on net samples utilized datasets gathered since the 1960s. Increased use of digital imaging approaches should enable us to obtain zooplankton biomass distribution estimates at basin to global scales in shorter time frames in the future.

Description: Artificial Upwelling (AU) of nutrient-rich Deep Ocean Water (DOW) to the ocean's sunlit surface layer has recently been put forward as a means of increasing marine CO2 sequestration and fish production. AU and its possible benefits have been studied in the context of climate change mitigation as well as food security for a growing human population. However, extensive research still needs to be done into the feasibility, effectiveness and potential risks, and side effects associated with AU to be able to better predict its potential. Fluid dynamic modeling of the AU process and the corresponding inorganic nutrient transport can provide necessary information for a better quantification of the environmental impacts of specific AU devices and represents a valuable tool for their optimization. Yet, appropriate capture of all flow phenomena relevant to the AU process remains a challenging task that only few models are able to accomplish. In this paper, simulation results obtained with a newly developed numerical solution method are presented. The method is based on the open-source modeling environment OpenFOAM. It solves the unsteady Reynolds-Averaged Navier-Stokes (RANS) equations with additional transport equations for energy, salinity, and inorganic nutrients. The method aims to be widely applicable to oceanic flow problems including temperature- and salinity-induced density stratification and passive scalar transport. The studies presented in this paper concentrate on the direct effects of the AU process on nutrient spread and concentration in the ocean's mixed surface layer. Expected flow phenomena are found to be captured well by the new method. While it is a known problem that cold DOW that is upwelled to the surface tends to sink down again due to its high density, the simulations presented in this paper show that the upwelled DOW settles at the lower boundary of the oceans mixed surface layer, thus keeping a considerable portion of the upwelled nutrients available for primary production. Comparative studies of several design variants, with the aim of maximizing the amount of nutrients that is retained inside the mixed surface layer, are also presented and analyzed

Description: When pelagic organisms die and fall onto the deep-sea floor they create food falls, parcels of organic enrichment that subsidize deep benthic scavenging communities. The diversity and quantities of food falls remains unstudied for many ocean regions, since they are stochastically deposited and rapidly scavenged. The Southern Ocean habitat supports large populations of megafauna but few food falls have been documented. To investigate the diversity and quantity of food falls in the northwestern Weddell Sea we analyzed 8476 deep-sea floor images that were captured during the expedition PS118 on RV Polarstern in 2019 by the camera system OFOBS (Ocean Floor Observation and Bathymetry System). OFOBS was towed 1.5 m above the seafloor along five transects (400 to 2200 m seafloor depth) east of the Antarctic Peninsula. We observed the carcasses of one baleen whale, one penguin, and four fish at depths of 647 m, 613 m, 647 m, 2136 m, 2165 m, and 2112 m, respectively, as well as associated scavenging fauna. To the best of our knowledge, we describe here the first in situ observations of deep-sea food falls for penguins and fish in the Southern Ocean. While the whale carcass seemed in an intermediate successional stage, both the penguin and the fish were likely recently deposited and three of the fish potentially resulted from fishery discards. Our relatively small data set suggests that a diverse array of food falls provide nutrients to the slopes of the Powell Basin.

Description: Teleost adaptive immune systems have evolved with more flexibility than previously assumed. A particularly enigmatic system to address immune system modifications in the evolutionary past is represented by the Syngnathids, the family of pipefishes, seahorses and seadragons. These small fishes with their unique male pregnancy have lost the spleen as an important immune organ as well as a functional major histocompatibility class II (MHC II) pathway. How these evolutionary changes have impacted immune cell population dynamics have up to this point remained unexplored. Here, we present the first immune cell repertoire characterization of a syngnathid fish (Syngnathus typhle) using single-cell transcriptomics. Gene expression profiles of individual cells extracted from blood and head-kidney clustered in twelve putative cell populations with eight belonging to those with immune function. Upregulated cell marker genes identified in humans and teleosts were used to define cell clusters. While the suggested loss of CD4+ T-cells accompanied the loss of the MHC II pathway was supported, the upregulation of specific subtype markers within the T-cell cluster indicates subpopulations of regulatory T-cells (il2rb) and cytotoxic T-cells (gzma). Utilizing single-cell RNA sequencing this report is the first to characterize immune cell populations in syngnathids and provides a valuable foundation for future cellular classification and experimental work within the lineage.

Description: The mesophotic domain is a poorly explored part of the oceans, notably in the Mediterranean Sea. Benthic communities in these depths are not well documented and as such are under higher risk from anthropogenic impacts. Hard substrate habitats in this depth window are not common and are a key ecotope. The Malta Plateau in the central Mediterranean, which is characterized by low sedimentation rates, offers a potentially unexplored domain for these features. Bathymetric and backscatter data offshore of the eastern coast of the island of Malta were used to identify 〉 1,700 small structures in mesophotic depths. These structures were verified to be biogenic mounds by dives. The mounds extend from several meters to tens of meters in diameter and occur in two main depth windows −40 to 83 meters below present sea level (mbpsl) and 83–120 mbpsl—each formed probably in a different stage during the last glacial cycle. The mounds are composed of interlocking bioconstruction by encrusting organisms and are colonized by sponges and various cold water corals (most of which are protected; e.g., Madrepora oculata). This unique and important habitat is currently under grave threat by human activity, most immediately by trawling and anchoring activity.

Description: Most studies on the potential impacts of deep-sea mining in the Clarion Clipperton Zone (CCZ) have largely focused on benthic ecosystems but ignore the pelagic environment. To model full-scale impacts, it is important to understand how sediment discharge might affect the pelagic zone as well. This study combines in situ optics, hydrography, and remote sensing to describe particle abundance and size distribution through the entire water column in the CCZ (German sector). CCZ surface waters were characterized as productive over the year. During the winter, we observed the formation of a sharp transition zone in Chla concentration, identifying the area as a productive transitional zone toward a more depleted ocean gyre. In the German sector, median particle size was small (± 77 μm), and large particles (〉300 μm) were rare. By assessing particle flux attenuation, we could show that the presence of a thick oxygen minimum zone (OMZ) plays an essential role in export and transformation of settling aggregates, with strong diel variations. We suggest that the combination of small aggregate size, bottom currents and slow seafloor consolidation may explain the extremely low sedimentation rate in the CCZ. We conclude that sediment incorporations and ballasting effect on settling particulate matter represent the most significant hazard on midwater and benthic ecosystems.

Description: Bacteriophages and their interactions with microbes are not well understood. As a first step toward achieving a better understanding, we isolated and sequenced the Curvibacter phage PCA1 for the purpose of eliminating Curvibacter sp. AEP1.3, the main colonizer of Hydra vulgaris AEP. Our experiments showed that PCA1 phage caused a strong, virulent infection only in sessile Curvibacter sp. AEP1.3 but was unable to infect planktonic and host-associated bacterial cells of the same strain. In an effort to investigate this phenomenon, we compared sessile, planktonic, and host-associated bacteria via RNA sequencing and found that all three states differed significantly in their expression patterns. This finding led us to propose that the adaptive lifestyle of Curvibacter sp. AEP1.3 results in varying degrees of susceptibility to bacteriophage infection. This concept could be relevant for phage research and phage therapy in particular. Finally, we were able to induce phage infection in planktonic cells and pinpoint the infection process to a membrane protein. We further identified potential phage-binding protein candidates based on expression pattern analysis.

Description: Landscape maps based on multivariate cluster analyses provide an objective and comprehensive view on the (marine) environment. They can hence support decision making regarding sustainable ocean resource handling and protection schemes. Across a large number of scales, input parameters and classification methods, numerous studies categorize the ocean into seascapes, hydro-morphological provinces or clusters. Many of them are regional, however, while only a few are on a basin scale. This study presents an automated cluster analysis of the entire Atlantic seafloor environment, based on eight global datasets and their derivatives: Bathymetry, slope, terrain ruggedness index, topographic position index, sediment thickness, POC flux, salinity, dissolved oxygen, temperature, current velocity, and phytoplankton abundance in surface waters along with seasonal variabilities. As a result, we obtained nine seabed areas (SBAs) that portray the Atlantic seafloor. Some SBAs have a clear geological and geomorphological nature, while others are defined by a mixture of terrain and water body characteristics. The majority of the SBAs, especially those covering the deep ocean areas, are coherent and show little seasonal and hydrographic variation, whereas other, nearshore SBAs, are smaller sized and dominated by high seasonal changes. To demonstrate the potential use of the marine landscape map for marine spatial planning purposes, we mapped out local SBA diversity using the patch richness index developed in landscape ecology. It identifies areas of high landscape diversity, and is a practical way of defining potential areas of interest, e.g. for designation as protected areas, or for further research. Clustering probabilities are highest (100%) in the center of SBA patches and decrease towards the edges (〈 98%). On the SBA point cloud which was reduced for probabilities 〈98%, we ran a diversity analysis to identify and highlight regions that have a high number of different SBAs per area, indicating the use of such analyses to automatically find potentially delicate areas. We found that some of the highlights are already within existing EBSAs, but the majority is yet unexplored.

Description: Benthic nitrogen cycling in the Mauritanian upwelling region (NW Africa) was studied in June 2014 from the shelf to the upper slope where minimum bottom water O 2 concentrations of 25 µM were recorded. Benthic incubation chambers were deployed at 9 stations to measure fluxes of O 2 , dissolved inorganic carbon (DIC) and nutrients (NO 3 - , NO 2 - , NH 4 + , PO 4 3- , H 4 SiO 4 ) along with the N and O isotopic composition of nitrate (δ 15 N-NO 3 - and δ 18 O-NO 3 - ) and ammonium (δ 15 N-NH 4 + ). O 2 and DIC fluxes were similar to those measured during a previous campaign in 2011 whereas NH 4 + and PO 4 3- fluxes on the shelf were 2 – 3 times higher and possibly linked to a long-term decline in bottom water O 2 concentrations. The mean isotopic fractionation of NO 3 - uptake on the margin, inferred from the loss of NO 3 - inside the chambers, was 1.5 ± 0.4 ‰ for 15/14 N ( 15 ϵ app ) and 2.0 ± 0.5 ‰ for 18/16 O ( 18 ϵ app ). The mean 18 ϵ app : 15 ϵ app ratio on the shelf (〈 100 m) was 2.1 ± 0.3, and higher than the value of 1 expected for microbial NO 3 - reduction. The 15 ϵ app are similar to previously reported isotope effects for NO 3 - respiration in marine sediments but lower than determined in 2011 at a same site on the shelf. The sediments were also a source of 15 N-enriched NH 4 + (9.0 ± 0.7 ‰). A numerical model tuned to the benthic flux data and that specifically accounts for the efflux of 15 N-enriched NH 4 + from the seafloor, predicted a net benthic isotope effect of N loss ( 15 ϵ sed ) of 3.6 ‰; far above the more widely considered value of ~0‰. This result is further evidence that the assumption of a universally low or negligible benthic N isotope effect is not applicable to oxygen-deficient settings. The model further suggests that 18 ϵ app : 15 ϵ app trajectories > 1 in the benthic chambers are most likely due to aerobic ammonium oxidation and nitrite oxidation in surface sediments rather than anammox, in agreement with published observations in the water column of oxygen deficient regions.

Description: Coastal and ocean island volcanoes are renowned for having unstable flanks. This can lead to flank deformation on a variety of temporal and spatial scales ranging from slow creep to catastrophic sector collapse. A large section of these unstable flanks is often below sea level, where information on the volcano-tectonic structure and ground deformation is limited. Consequently, kinematic models that attempt to explain measured ground deformation onshore associated with flank instability are poorly constrained in the offshore area. Here, we attempt to determine the locations and the morpho-tectonic structures of the boundaries of the submerged unstable southeastern flank of Mount Etna (Italy). The integration of new marine data (bathymetry, microbathymetry, offshore seismicity, reflection seismic lines) and published marine data (bathymetry, seafloor geodesy, reflection seismic lines) allows identifying the lineament north of Catania Canyon as the southern lateral boundary with a high level of confidence. The northern and the distal (seaward) boundaries are less clear because no microbathymetric or seafloor geodetic data are available. Hypotheses for their locations are presented. Geophysical imaging suggests that the offshore Timpe Fault System is a shallow second-order structure that likely results from extensional deformation within the moving flank. Evidence for active uplift and compression upslope of the amphitheater-shaped depression from seismic data along with subsidence of the onshore Giarre Wedge block observed in ground deformation data leads us to propose that this block is a rotational slump, which moves on top of the large-scale instability. The new shoreline-crossing structural assessment may now inform and improve kinematic models.

Description: Barkley Canyon is one of the few known sites worldwide with the occurrence of thermogenic gas seepage and formation of structure-II and structure-H gas hydrate mounds on the seafloor. This site is the location of continuous seafloor monitoring as part of the Ocean Networks Canada (ONC) cabled observatory off the west coast off Vancouver Island, British Columbia, Canada. We combine repeat remotely operated vehicle (ROV) seafloor video observations, mapping with an autonomous underwater vehicle (AUV), ship-, ROV-, and AUV-based identification of gas flares, as well as seismic and Chirp data to investigate the distribution of fluid migration pathways. Geologically, the site with the prominent gas hydrate mounds and associated fluid seepage is covering an area of ∼0.15 km 2 and is situated on a remnant of a rotated fault block that had slipped off the steep flanks of the north-east facing canyon wall. The gas hydrate mounds, nearly constant in dimension over the entire observation period, are associated with gas and oil seepage and surrounded by debris of chemosynthetic communities and authigenic carbonate. The formation of gas hydrate at and near the seafloor requires additional accommodation space created by forming blisters at the seafloor that displace the regular sediments. An additional zone located centrally on the rotated fault block with more diffuse seepage (∼0.02 km 2 in extent) has been identified with no visible mounds, but with bacterial mats, small carbonate concretions, and clam beds. Gas venting is seen acoustically in the water column up to a depth of ∼300 m. However, acoustic water-column imaging during coring and ROV dives showed rising gas bubbles to much shallower depth, even 〈50 m, likely a result of degassing of rising oil droplets, which themselves cannot be seen acoustically. Combining all observations, the location of the gas hydrate mounds is controlled by a combination of fault-focused fluid migration from a deeper reservoir and fluid seepage along more permeable strata within the rotated slope block. Fluids must be provided continuously to allow the sustained presence of the gas hydrate mounds at the seafloor.

Description: Salinity is a major environmental factor shaping the distribution and abundance of marine organisms. Climate change is predicted to alter salinity in many coastal regions due to sea level rise, evaporation, and changes in freshwater input. This exerts significant physiological stress on coastal invertebrates whose body fluid osmolality follows that of seawater ('osmoconformers'). In this study, we conducted a systematic review and meta-analysis of osmolytes (both organic and inorganic) utilized by osmoconforming marine invertebrates during a 〉 14-day acclimation to reduced salinity. Of the 2,389 studies screened, a total of 56 fulfilled the search criteria. Thirty-eight studies reported tissue osmolyte. Following acclimation to reduced salinity, tissue concentrations of six organic compounds and sodium were consistently reduced across phyla. This suggests that intracellular inorganic ions are not only utilized as a rapid response system during acute exposure to low salinity stress but also, in concert with reductions in organic osmolyte concentrations, during longer-term acclimation. Our systematic review demonstrates that only a few studies (n = 13) have quantified salinity-induced long-term changes in intracellular ion concentrations. In addition, no study has compiled a complete intracellular osmolyte budget. Alanine, betaine, glycine, and taurine are the major organic osmolytes that are universally employed across five phyla. The characterization of organic osmolytes was heavily weighted towards free amino acids (FAAs) and derivatives-neglecting methylamines and methylsulfonium compounds, which can be as important as FAAs in modulating intracellular osmolality. As a consequence, we suggest best-practice guidelines to streamline experimental designs and protocols in osmoregulation research in order to better understand the conserved mechanisms that define the limits of salinity acclimation in marine invertebrates. To our best knowledge, this is the first systematic review and meta-analysis on osmolyte concentrations in osmoconformers acclimated to low salinity. It creates a valuable baseline for future research and reveals large research gaps. Our meta-analysis suggests that there are common osmolyte actors employed across phyla but no uniform concept since osmolyte pool composition and proportions were taxon-specific. In light of future salinity changes and their potential consequences, it becomes more important to understand salinity tolerance capacities and limits.

Description: Recent oceanographic observations have identified significant changes of intermediate water masses characterized by increased temperatures, lowered pH and deoxygenation. In order to improve our understanding as to how these changes may impact deep-sea ecosystems one important strategy is to reconstruct past oceanic conditions. Here we examine the applicability of the scleractinian cold-water coral Solenosmilia variabilis as a marine archive for the reconstructions of past intermediate water mass temperatures by using Lithium (Li)/Magnesium (Mg) ratios. In particular, our study addresses 1) the calibration of Li/Mg ratios against in-situ temperature data, 2) the reconstruction of past intermediate water mass temperatures using scleractinian coral fossil samples from the Brazilian continental margin and 3) the identification of intraspecies variability within the coral microstructure. Results showed that Li/Mg ratios measured in the skeletons of S. variabilis fit into existing Li/Mg-T calibrations of other cold-water scleractinian. Furthermore, the coral microstructure exhibits interspecies variability of Li/Ca and Mg/Ca ratios were also similar to what has been observed in other cold-water scleractinian corals, suggesting a similar biomineralization control on the incorporation of Li and Mg into the skeleton. However, the Li/Mg based temperature reconstruction using fossil samples resulted in unexpectedly high variations >10°C, which might not be solely related to temperature variations of the intermediate water mass over the last 160 ka on the Brazilian continental margin. We speculate that such temperature variability may be caused by vertical movements of the aragonite saturation horizon and the associated seawater pH changes, which in turn influence the incorporation of Li and Mg into the coral skeleton. Based on these results it is recommended that future studies investigating past oceanic conditions need to consider the carbonate system parameters and how they might impact the mechanisms of Li and Mg being incorporated into skeletons of cold-water coral species such as S. variabilis.

Description: The modern precipitation balance in southeastern (SE) Brazil is regulated by the South American summer Monsoon and threatened by global climate change. On glacial-interglacial timescales, monsoon intensity was strongly controlled by precession-forced changes in insolation. To date, relatively little is known about the spatiotemporal distribution of tropical precipitation in SE Brazil and the resulting variability of fluvial discharge on glacial-interglacial timescales. Here, we present X-ray diffraction-derived mineralogical data for the 150–70 ka period (marine isotope stage (MIS) 6 to MIS 5) from the Doce River basin. This area was sensitive to changes in monsoonal precipitation intensity due to its proximity to the South Atlantic Convergence Zone. The data, obtained from a marine sediment core (M125-55–7) close to the Doce river mouth (20°S), show pronounced changes in the Doce River suspension load’s mineralogical composition on glacial-interglacial and precessional timescales. While the ratio of silicates to carbonates displays precession-paced changes, the mineralogical composition of the carbonate-free fraction discriminates between two assemblages which strongly vary between glacial and interglacial time scales, with precession-forced variability only visible in MIS 5. The first assemblage, dominated by high contents of kaolinite and gibbsite, indicates intensified lowland erosion of mature tropical soils. The second one, characterized by higher contents of the well-ordered illite, quartz and albite, points to intensified erosion of immature soils in the upper Doce Basin. High kaolinite contents in the silicate fraction prevailed in late MIS 6 and indicate pronounced lowland soil erosion along a steepened topographic gradient. The illite-rich mineral assemblage was more abundant in MIS 5, particularly during times of high austral summer insolation, indicating strong monsoonal rainfall and intense physical erosion in the upper catchment. When the summer monsoon weakened in times of lower insolation, the mineral assemblage was dominated by kaolinite again, indicative of lower precipitation and runoff in the upper catchment and dominant lowland erosion.

Description: In its latest assessment report the IPCC stresses the need for carbon dioxide removal (CDR) to counterbalance residual emissions to achieve net zero carbon dioxide or greenhouse gas emissions. There are currently a wide variety of CDR measures available. Their potential and feasibility, however, depends on context specific conditions, as among others biophysical site characteristics, or availability of infrastructure and resources. In our study, we selected 13 CDR concepts which we present in the form of exemplary CDR units described in dedicated fact sheets. They cover technical CO2 removal (two concepts of direct air carbon capture), hybrid solutions (six bioenergy with carbon capture technologies) and five options for natural sink enhancement. Our estimates for their CO2 removal potentials in 2050 range from 0.06 to 30 million tons of CO2, depending on the option. Ten of the 13 CDR concepts provide technical removal potentials higher than 1 million tons of CO2 per year. To better understand the potential contribution of analyzed CDR options to reaching net-zero CO2 emissions, we compare our results with the current CO2 emissions and potential residual CO2 emissions in 2050 in Germany. To complement the necessary information on technology-based and hybrid options, we also provide an overview on possible solutions for CO2 storage for Germany. Taking biophysical conditions and infrastructure into account, northern Germany seems a preferable area for deployment of many concepts. However, for their successful implementation further socio-economic analysis, clear regulations, and policy incentives are necessary.