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Estuarine intertidal sediments are important centres for organic matter remineralization and nutrients recycling. Nevertheless, there is limited understanding regarding how these processes occur along the salinity gradient and their seasonality. Here, we report on the seasonal biogeochemical cycles from three types of intertidal sedimentary habitats (freshwater, brackish and marine) located in the Western Scheldt estuary (The Netherlands and Belgium). A full year of solute fluxes, porewater nutrient and sediment pigment concentrations at a monthly resolution revealed clear differences in the biogeochemistry of the three sites, indicating that environmental conditions determined the local nutrient dynamics. Temperature controlled sediment oxygen consumption rates and nutrient fluxes, but also affected pore water nutrient concentrations up to 14 cm deep. Fresh and brackish sediments had a net influx of dissolved inorganic nitrogen (DIN) (−1.62 mmol m−2 d−1 and -2.84 mmol m−2 d−1, respectively), while only the freshwater sediments showed a net influx of phosphate (−0.07 mmol m−2 d−1). We estimated that intertidal sediments remineralized a total of 10,000 t C y−1, with 97% of mineralization occurring in the brackish and marine parts. Overall, sediments removed 11% (1500 t N y−1) and 15% (∼200 t P y−1) of the total nitrogen and phosphorus entering the estuary from riverine input. Moreover, observations revealed the historical improvement of water quality resulting from water treatment policies. This spatiotemporal study of OM remineralization and early diagenesis in estuarine systems highlights the importance of intertidal sediments for estuarine systems. Our observations can be used in models to predict estuarine biogeochemistry or assess climate change scenarios.

Coral reefs in the northern Red Sea experience strong seasonality. This affects reef carbon (C) cycling, but ecosystem-wide quantification of C fluxes in such reefs is limited. This study quantified seasonal reef community C fluxes with incubations. Resulting data were then incorporated into seasonal linear inverse models (LIM). For spring, additional sponge incubation results allowed for unique assessment of the contribution of sponges to C cycling. The coral reef ecosystem was heterotrophic throughout all seasons as gross community primary production (GPP; 136–200, range of seasonal means in mmol C m−2 d−1) was less than community respiration (R; 192–279), and balanced by import of organic carbon (52–100), 88‒92% of which being dissolved organic carbon (DOC). Hard coral GPP (74–110) and R (100–137), as well as pelagic bacteria DOC uptake (58–101) and R (42–86), were the largest C fluxes across seasons. The ecosystem was least heterotrophic in spring (highest irradiance) (GPP:R 0.81), but most heterotrophic in summer and fall with higher water temperatures (0.68 and 0.60, respectively). Adding the sponge community to the model increased community R (247 ± 8 without to 353 ± 13 with sponges (mean ± SD)). Sponges balanced this demand primarily with DOC uptake (105 ± 6, 97% by cryptic sponges). This rate is comparable to the uptake of DOC by pelagic bacteria (104 ± 5) placing the cryptic sponges among the dominant C cycling groups in the reef.

Coastal areas are subjected to several anthropogenic stressors with much of the world's intertidal areas receding due to human activities, coastal erosion and sea level rise. The Dutch Eastern Scheldt (ES) has been predicted to lose around 35% of intertidal areas by 2060. This study investigates differences between biogeochemical fluxes of intertidal and subtidal sediments of the ES and assesses how ongoing erosion may modify the sedimentary ecosystem functioning of this coastal bay in the coming decades. Monthly fluxes and porewater concentrations of dissolved inorganicnitrogen (DIN), phosphorous (DIP), silica (DSi), carbon (DIC) and oxygen (O 2) as well as organic matter (OM) characteristics were measured from intertidal and subtidal sediments from June 2016–December 2017. Compared to subtidal stations, OM was significantly more reactive in intertidal samples and exhibited 37% higher O2 fluxes, suggesting a strong influence from microphytobenthos. Subtidal sedimentsexhibited an average efflux of nitrates (0.28 mmol m−2 d −1) and phosphates (0.09 mmol m−2 d−1) into the water column, while intertidal areas displayed average influxes (nitrates = −1.2 mmol m−2 d−1, phosphates = −0.03 mmol m−2 d−1) directed into the sediment. The calculated removal of total DIN and DIP from the water column was 34–38% higher in intertidal compared to subtidal samples suggesting stronger denitrification and phosphorus adsorption to solid particles from intertidal sediments. As an upscaling exercise, we estimate potential erosion induced changes if the ES stations are representative for the system. With this assumption, we estimate 11% and 8% reductions for respective nitrogen and phosphorus removal in the entire ES by 2060. Given the global observations of eroding intertidal areas and rising sea levels, we suggest that the predicted habitat loss may cause significant changes

The Eastern Scheldt estuary in the Netherlands has been anthropogenically changed by the construction of a storm surge barrier about 30 years ago, affecting abiotic conditions as well as phytoplankton and zooplankton (ZP). As ZP communities have not been sampled in the last decades, it is unclear to what extent they have changed during the last 30 years. We analyzed the micro- and mesoZP composition and background parameters at eight stations in the Eastern Scheldt in spring, summer, and autumn 2018. Additionally, we conducted dilution experiments to estimate phytoplankton growth and microZP grazing rates. Seasonal variability of ZP was higher than spatial variability, but there were differences between stations. In agreement with data from the 1980s, we identified salinity and chlorophyll a as the most important factors in explaining the ZP community. Suspended particulate matter and water retention time were additionally identified for mesoZP, as well as dissolved inorganic nitrogen and phosphate concentration for microZP. We observed some changes in ZP composition with respect to the 1980s, most of which could be explained by the changes induced by the barrier construction. However, increased abundances of the cyclopoid copepod Oithona sp. were likely related to changes in phytoplankton composition due to large-scale bivalve cultivation in the estuary. Phytoplankton growth and microZP grazing rates were variable between seasons, which could be explained by changes in the phytoplankton community during the annual plankton succession. Overall, the observed alterations in the ZP community due to changed background parameters and intense bivalve filtration could affect the microbial loop as well as consumers of mesoZP and thus the functioning of the pelagic food web.

The abyssal seafloor (3500–6000m) remains largely unexplored but with deep-sea mining imminent, anthropogenic impacts may soon reach abyssal communities. Thus, there is a growing need for baseline studies of biodiversity, ecosystem functioning, and connectivity in both potential mining and no-mining areas across the Clarion-Clipperton Zone (CCZ), a key target region for polymetallic nodule mining. In this study, in situ pulse-chase lander experiments were conducted for 1.5 days in three no-mining areas (called Areas of Particular Environmental Interest, or APEIs) in the western CCZ, a region with a seafloor particulate organic carbon (POC) flux gradient. A decreasing trend was seen in mean seafloor respiration, macrofaunal abundance, and biomass from the more eutrophic APEI 7 to the more oligotrophic APEI 1, although this trend was not statistically significant (p = 0.18) most likely due to small samples sizes and high variability. In this study, most (96%) of the 13C-labeled processed phytodetritus was respired within 1.5 days. Experimental uptake of phytodetritus by macrofauna and bacteria was detected but was lower than in the previously studied and more eutrophic eastern CCZ over similar time scales (1.5 d). Bacteria dominated the short-term (∼1.5 d) uptake of organic carbon at the seafloor, yet macrofauna processed more organic carbon per unit biomass than previously found in the eastern CCZ (0.003 mg C m −2 d −1 and 0.5 × 10 −5 mg C m −2 d −1 for the western and eastern CCZ, respectively). Our study provides important information on C-uptake and respiration rates in areas set aside from mining in the western CCZ and suggests high variability may occur in the rates of benthic Corg-cycling across the CCZ. We recommend that benthic ecosystem functions be explored across gradients of POC flux which may be a major environmental factor driving ecosystem dynamics in the CCZ.

The deep sea is amongst the most food-limited habitats on Earth, as only a small fraction (<4%) of the surface primary production is exported below 200 m water depth. Here, cold-water coral (CWC) reefs form oases of life: their biodiversity compares with tropical coral reefs, their biomass and metabolic activity exceed other deep-sea ecosystems by far. We critically assess the paradox of thriving CWC reefs in the food-limited deep sea, by reviewing the literature and open-access data on CWC habitats. This review shows firstly that CWCs typically occur in areas where the food supply is not constantly low, but undergoes pronounced temporal variation. High currents, downwelling and/or vertically migrating zooplankton temporally boost the export of surface organic matter to the seabed, creating ‘feast’ conditions, interspersed with ‘famine’ periods during the non-productive season. Secondly, CWCs, particularly the most common reef-builder Desmophyllum pertusum (formerly known as Lophelia pertusa), are well adapted to these fluctuations in food availability. Laboratory and in situ measurements revealed their dietary flexibility, tissue reserves, and temporal variation in growth and energy allocation. Thirdly, the high structural and functional diversity of CWC reefs increases resource retention: acting as giant filters and sustaining complex food webs with diverse recycling pathways, the reefs optimise resource gains over losses. Anthropogenic pressures, including climate change and ocean acidification, threaten this fragile equilibrium through decreased resource supply, increased energy costs, and dissolution of the calcium-carbonate reef framework. Based on this review, we suggest additional criteria to judge the health of CWC reefs and their chance to persist in the future.

Cold-water coral (CWC) reefs are numerous and widespread along the Norwegian continental shelf where oil and gas industry operate. Uncertainties exist regarding their impacts from operational discharges to drilling. Effect thresholds obtained from near-realistic exposure of suspended particle concentrations for use in coral risk modeling areparticularly needed. Here, nubbins of Desmophyllum pertusum ( Lophelia pertusa) were exposed shortly (5 days, 4h repeated pulses) to suspended particles (bentonite BE; barite BA, and drill cuttings DC) in the range of ~ 4 to ~ 60 mg.l-1 (actual concentration). Physiological responses (respiration rate, growth rate, mucus-related particulate organic carbon OC and particulate organic nitrogen ON) and polyp mortality were then measured 2 and 6 weeks post-exposure to assess long-term effects. Respiration and growth rates were not significantly different in any of the treatments tested compared to control. OC production was not affected in any treatment, but a significant increase of OC:ON in mucus produced by BE-exposed (23 and 48 mg.l-1) corals was revealed 2 weeks after exposure. Polyp mortality increased significantly at the two highest DC doses (19 and 49 mg.l-1) 2 and 6 weeks post-exposure but no significant difference was observed in any of the other treatments compared to the control. These findings are adding new knowledge on coral resilience to short realistic exposure of suspended drill particles and indicate overall a risk for long-term effects at a threshold of ~20 mg.l-1.

Cold-water corals (CWCs) are important ecosystem engineers in the deep sea that provide habitat for numerous species and can form large coral mounds. These mounds influence surrounding currents and induce distinct hydrodynamic features, such as internal waves and episodic downwelling events that accelerate transport of organic matter towards the mounds, supplying the corals with food. To date, research on organic matter distribution at coral mounds has focussed either on seasonal timescales or has provided single point snapshots. Data on food distribution at the timescale of a diurnal tidal cycle is currently limited. Here, we integrate physical, biogeochemical, and biological data throughout the water column and along a transect on the south-eastern slope of Rockall Bank, Northeast Atlantic Ocean. This transect consisted of 24-hour sampling stations at four locations: Bank, Upper slope, Lower slope, and the Oreo coral mound. We investigated how the organic matter distribution in the water column along the transect is affected by tidal activity. Repeated CTD casts indicated that the water column above Oreo mound was more dynamic than above other stations in multiple ways. First, the bottom water showed high variability in physical parameters and nutrient concentrations, possibly due to the interaction of the tide with the mound topography. Second, in the surface water a diurnal tidal wave replenished nutrients in the photic zone, supporting new primary production. Third, above the coral mound an internal wave (200 m amplitude) was recorded at 400 m depth after the turning of the barotropic tide. After this wave passed, high quality organic matter was recorded in bottom waters on the mound coinciding with shallow water physical characteristics such as high oxygen concentration and high temperature. Trophic markers in the benthic community suggest feeding on a variety of food sources, including phytodetritus and zooplankton. We suggest that there are three transport m

Nitrogen (N) is often the key nutrient limiting primary production in coastal waters. Quantifying sources and sinks of N is therefore critical to understanding the factors that underpin the productivity of coastal ecosystems. Constraining nitrogen inputs can be difficult for some terms such as N fixation and marine exchange as a consequence of uncertainties associated with scaling and stochasticity. To help overcome these issues, we undertook a N budget incorporating an isotope and mass balance to constrain N sources in a large oligotrophic coastal embayment (Western Port, Australia). The total N input to Western Port was calculated to be 1400 Mg N year−1, which is remarkably consistent with previous estimates of sedimentation rates within the system. Catchment inputs, N fixation, marine sources and atmospheric deposition comprised 44, 28, 28 and 13% of N inputs respectively. Retention of marine-derived N equated to ~3 and ~10% of total N and NOx flushed through the system from the marine end-member. The relatively high contribution of N fixation compared with previous studies was most likely to be due to the high proportion of nutrient-limited intertidal sediments where N is mediated by seagrasses and sediment cyanobacteria.

Cold-water corals rely on currents to transport food towards them and when external conditions are favourable, they can form coral mounds. These structures, which can be over 300 m high, influence the hydrodynamics around the reefs that grow on the mounds, which feeds back to affect coral- and therefore mound-growth. We investigated these feedbacks at the Logachev coral mound province, by running simulations with a 3D hydrodynamic model (Roms-Agrif), using different seafloor bathymetries that represent consecutive stages of mound development. Simulations ranged from a fully smoothened bathymetry without mounds, to a coral mound (Haas mound) at 1.5 times its current size. The effect of mound height on coral growth was investigated by looking at the baroclinic (internal) tide, turbulent energy dissipation, vertical velocities, and horizontal bottom currents. The simulations suggest that with increasing mound height horizontal velocities increase, while turbulent energy dissipation and vertical velocities around the mound foot decrease. This supposedly limits coral growth at the mound foot and hence lateral mound extension in later stages of development. An increase in turbulent energy dissipation and vertical velocities on the mound top and upper flanks, indicates vertical mound growth at all subsequent stages. Our findings of continued vertical mound growth provide an explanation for recently published data on benthic cover from a transect over Haas mound, that show a dominance of live corals on the mound top. We find areas of increased energy conversion rates from the barotropic (surface) to the baroclinic tide on the bathymetry where we artificially eliminated the mounds from (i.e. smoothened bathymetry). Interestingly, these areas overlap with the region where coral mounds are located at present. So, the baroclinic tide is likely an important mechanism in the process of coral mound establishment. Given the relative ease with which the energy conversion rate from t

Both physical and electrical impacts have been linked to North Sea fisheries activity. This study evaluates how these effects can influence marine ecological functioning by assessing their consequences on benthic pelagic coupling . Experiments were conducted on sediment microcosms taken from 9 North Sea and 2 Eastern Scheldt locations. Samples were subjected to physical disturbances by mechanically stirring the sediment surface or electrical stimulation with exposure to high frequency pulsed bipolar or direct currents.Electrical exposure times of 3 and 120-s were used to simulate in situ exposure times related to sole ( Solea solea ) and razor clam (Ensis spp.) electric fisheries respectively. Water column oxygen rapidly declined after sediment resuspension , inducing an immediate uptake ranging from 0.55 to 22 mmol oxygen per m

Deep-sea sponges and their microbial symbionts transform various forms of carbon (C) and nitrogen (N) via several metabolic pathways, which, for a large part, are poorly quantified. Previous flux studies on the common deep-sea sponge Geodia barretti consistently revealed net consumption of dissolved organic carbon (DOC) and oxygen (O2) and net release of nitrate (NO−3NO3-). Here we present a biogeochemical metabolic network model that, for the first time, quantifies C and N fluxes within the sponge holobiont in a consistent manner, including many poorly constrained metabolic conversions. Using two datasets covering a range of individual G. barretti sizes (10–3,500 ml), we found that thevariability in metabolic rates partially resulted from body size as O 2 uptake allometrically scales with sponge volume. Our model analysis confirmed that dissolved organic matter (DOM), with an estimated C:N ratio of 7.7 ± 1.4, is the main energy source of G. barretti. DOM is primarily used for aerobic respiration, then for dissimilatory NO−3NO3- reduction to ammonium (NH+4)NH4+) (DNRA), and, lastly, for denitrification. Dissolved organic carbon (DOC) production efficiencies (production/assimilation) were estimated as 24 ± 8% (larger individuals) and 31 ± 9% (smaller individuals), so most DOC was respired to carbon dioxide (CO2), which was released in a net ratio of 0.77–0.81 to O2 consumption. Internally produced NH+4NH4+ from cellular excretion and DNRA fueled nitrification. Nitrification-associated chemoautotrophic production contributed 5.1–6.7 ± 3.0% to total sponge production. While overall metabolic patterns were rather independent of sponge size, (volume-)specific rates were lower in larger sponges comparedto smaller individuals. Specific biomass production rates were 0.16% day–1 in smaller compared to 0.067% day–1 in larger G. barretti as expected for slow-growi

Arctic coastal ecosystems are rapidly changing due to climate warming. This makes modeling their productivity crucially important to better understand future changes. System primary production in these systems is highest during the pronounced spring bloom, typically dominated by diatoms. Eventually the spring blooms terminate due to silicon or nitrogen limitation. Bacteria can play an important role for extending bloom duration and total CO2 fixation through ammonium regeneration. Current ecosystem models often simplify the effects of nutrient co-limitations on algal physiology and cellular ratios and simplify nutrient regeneration. These simplifications may lead to underestimations of primary production. Detailed biochemistry- and cell-based models can represent these dynamics but are difficult to tune in the environment. We performed a cultivation experiment that showed typical spring bloom dynamics, such as extended algal growth via bacterial ammonium remineralization, reduced algal growth and inhibited chlorophyll synthesis under silicate limitation, and gradually reduced nitrogen assimilation and chlorophyll synthesis under nitrogen limitation. We developed a simplified dynamic model to represent these processes. Overall, model complexity in terms of the number of parameters is comparable to the phytoplankton growth and nutrient biogeochemistry formulations in common ecosystem models used in the Arctic while improving the representation of nutrient-co-limitation-related processes. Such model enhancements that now incorporate increased nutrient inputs and higher mineralization rates in a warmer climate will improve future predictions in this vulnerable system.

Polymetallic nodule fields provide hard substrate for sessile organisms on the abyssal seafloor between 3000 and 6000 m water depth. Deep-seabed mining targets these mineral-rich nodules and will likely modify the consumer-resource (trophic) and substrate-providing (non-trophic) interactions within the abyssal food web. However, the importance of nodules and their associated sessile fauna in supporting food-web integrity remains unclear. Here, we use seafloor imagery and published literature to develop highly-resolved trophic and non-trophic interaction webs for the Clarion-Clipperton Fracture Zone (CCZ, central Pacific Ocean) and the Peru Basin (PB, South-East Pacific Ocean) and to assess how nodule removal may modify these networks. The CCZ interaction web included 1028 compartments connected with 59,793 links and the PB interaction web consisted of 342 compartments and 8044 links. We show that knock-down effects of nodule removal resulted in a 17.9% (CCZ) to 20.8% (PB) loss of all taxa and 22.8% (PB) to 30.6% (CCZ) loss of network links. Subsequent analysis identified stalked glass sponges living attached to the nodules as key structural species that supported a high diversity of associated fauna. We conclude that polymetallic nodules are critical for food-web integrity and that their absence will likely result in reduced local benthic biodiversity.

The feeding biology of deep-sea octocorals remains poorly understood, as attention is more often directed to reef building corals. The present study focused on two common deep-water octocoral species in the Azores Archipelago, Dentomuricea aff. meteor and Viminella flagellum, aiming at determining their ability to exploit different food sources. We adopted an experimental approach, with three different food sources, including live phytoplankton, live zooplankton and dissolved organic matter (DOM), that were artificially enriched with 13C and 15N (C and N tracers). The presence of tracers was subsequently followed in the coral tissue, C respiration and particulate organic C and N (POC and PON) release. In both species, feeding with zooplankton resulted in significantly higher incorporation of tracers in all measured variables, compared to the other food sources, highlighting the importance of zooplankton for major physiological processes. Our results revealed contrasting metabolic strategies between the two species, with D. aff. meteor acquiring higher amounts of prey and allocating higher percentage to respiration and release of POC and PON than V. flagellum. Such metabolic differences can shape species fitness and distributions and have further ecological implications on the ecosystem function of communities formed by different octocoral species.

Cold-water corals (CWC) are known to tolerate a relatively wide range of environmental conditions. However, along the basin margins of the Rockall Trough (NE Atlantic), the habitat of CWC is confined to a narrow range of 525–1200 m water depth, and the reason for that is not fully understood. To investigate the distribution of CWCs in this area, current velocities and water mass distribution in the Logachev Mound Province were measured with two long-term (1 year) moorings equipped with an acoustic doppler current profiler and fluorescence+turbidity sensors. Additional ship-based water column profiles and discrete water samples, covering a full diurnal tidal cycle, were taken for chemical parameters. The results indicate the presence of a boundary current along the eastern flank of the Rockall Bank, that transports a nutrient-rich water mass southwards and governs the direction of particle transport at the depth band of the CWC mounds. Based on literature results, this northern water mass is identified as Wyville Thomson Ridge Overflow Water (WTOW). The density envelope and depth distribution of WTOW match the reported occurrence of CWC mounds in the Rockall Trough. Hence, the presence of WTOW may be a necessary condition for coral growth and therefore mound formation in the Rockall Trough, e.g. by forming a conduit for particles and coral larvae.

The deep sea is the largest biome on Earth but the least explored. Our knowledge of it comes from scattered sources spanning different spatial and temporal scales. Implementation of marine policies like the European Union’s Marine Strategy Framework Directive (MSFD) and support for Blue Growth in the deep sea are therefore hindered by lack of data. Integrated assessments of environmental status require tools to work with different and disaggregated datasets (e.g. density of deep-sea habitat-forming species, body-size distribution of commercial fishes, intensity of bottom trawling) across spatial and temporal scales. A feasibility study was conducted as part of the four-year ATLAS project to assess the effectiveness of the open-access Nested Environmental status Assessment Tool (NEAT) to assess deep-sea environmental status. We worked at nine selected study areas in the North Atlantic focusing on five MSFD descriptors (D1-Biodiversity, D3-Commercial fish and shellfish, D4-Food webs, D6-Seafloor integrity, D10-Marine litter). The objectives of the present study were to i) explore and propose indicators that could be used in the assessment of deep-sea environmental status, ii) evaluate the performance of NEAT in the deep sea, and iii) identify challenges and opportunities for the assessment of deep-sea status. Based on data availability, data quality and expert judgement, in total 24 indicators (one for D1, one for D3, seven for D4, 13 for D6, two for D10) were used in the assessment of the nine study areas, their habitats and ecosystem components. NEAT analyses revealed differences among the study areas for their environmental status ranging from “poor” to “high”. Overall, the NEAT results were in moderate to complete agreement with expert judgement, previous assessments, scientific literature on human-pressure gradients and expected management outcomes. We suggest that the assessment of deep-sea environmental status should take place at habitat and ecosystem level (r

Microphytobenthos forms an important part of the diet of macrofauna (macrozoobenthos) in many intertidal ecosystems. It is unclear, however, whether the dependence of macrofauna on microphytobenthos varies spatially within and among tidal systems. We aim (1) to assess the spatial variability in the importance of microphytobenthos in the diet of macrofauna (i.e., between and within two tidal basins and as function of elevation), (2) to quantify grazing pressure of the macrofaunal community on different potential food sources (microphytobenthos, phytoplankton and terrestrial organic material) for several sites in two tidal basins and (3) to compare microphytobenthic production and summer/autumn grazing of the total macrofaunal community and grazing pressure per feeding type, with potential microphytobenthic production estimated from rates in early spring, when grazing was low. Using a natural stable isotope approach, we identified microphytobenthos as a more important food source for macrofauna than phytoplankton and terrestrial organic material. Microphytobenthos dependency differed between tidal basins for the genera Bathyporeia (sand digger shrimp), Macoma (Baltic tellin), and Peringia (mudsnail) and for sampled individuals of all genera combined, and did not vary as function of elevation. We showed that macrofaunal grazing on microphytobenthos is quantitatively important and, in some cases, approached microphytobenthic production rates in early spring. No positive relation between microphytobenthic production in early spring and macrofaunal grazing in summer/autumn was observed. This suggests that the studied consumer‐resource interactions are coupled on a larger spatial scale (i.e., mesoscale, ≈10 to 100 km), rather than the fine (mm to m) scale.

Cold-water corals build extensive reefs on the seafloor that are oases of biodiversity, biomass, and organic matter processing rates. The reefs baffle sediments, and when coral growth and sedimentation outweigh ambient sedimentation, carbonate mounds of tens to hundreds of meters high and several kilometers wide can form. Because coral mounds form over ten-thousands of years, their development process remains elusive. While several environmental factors influence mound development, the mounds also have a major impact on their environment. This feedback between environment and mounds, and how this drives mound development is the focus of this paper. Based on the similarity of spatial coral mound patterns and patterns in self-organized ecosystems, we provide a new perspective on coral mound development. In accordance with the theory of self-organization through scale-dependent feedbacks, we first elicit the processes that are known to affect mound development, and might cause scale-dependent feedbacks. Then we demonstrate this concept with model output from a study on the Logachev area, SW Rockall Trough margin. Spatial patterns in mound provinces are the result of a complex set of interacting processes. Spatial self-organization provides a framework in which to place and compare these processes, so as to assess if and how they contribute to pattern formation in coral mounds.

Cold-water coral (CWC) reefs are one of the most diverse and productive ecosystems in the deep sea. Especially in periods of seasonally-reduced phytodetritus food supply, their high productivity may depend on the recycling of resources produced on the reef, such as dissolved organic matter (DOM) and bacteria. Here, we demonstrate that abundant suspension feeders Geodia barretti (high-microbial-abundance sponge), Mycale lingua (low-microbial-abundance sponge) and Acesta excavata (bivalve) are able to utilize 13C-enriched (diatom-derived) DOM and bacteria for tissue growth and respiration. While DOM was an important potential resource for all taxa, utilization of bacteria was higher for the sponges as compared to the bivalve, indicating a particle-size differentiation among the investigated suspension feeders. Interestingly, all taxa released 13C-enriched particulate organic carbon, which in turn may feed the detritus pathway on the reef. Especially A. excavata produced abundant (pseudo-)fecal droppings. A second stable-isotope tracer experiment revealed that detritivorous ophiuroids utilized these droppings. The high resource flexibility of dominant reef suspension feeders, and the efficient recycling of their waste products by the detritivore community, may provide important pathways to maintain the high productivity on cold-water coral reefs, especially in periods of low external food supply.

The deep sea plays a critical role in global climate regulation through uptake and storage of heat and carbon dioxide. However, this regulating service causes warming, acidification and deoxygenation of deep waters, leading to decreased food availability at the seafloor. These changes and their projections are likely to affect productivity, biodiversity and distributions of deep‐sea fauna, thereby compromising key ecosystem services. Understanding how climate change can lead to shifts in deep‐sea species distributions is critically important in developing management measures. We used environmental niche modelling along with the best available species occurrence data and environmental parameters to model habitat suitability for key cold‐water coral and commercially important deep‐sea fish species under present‐day (1951–2000) environmental conditions and to project changes under severe, high emissions future (2081–2100) climate projections (RCP8.5 scenario) for the North Atlantic Ocean. Our models projected a decrease of 28%–100% in suitable habitat for cold‐water corals and a shift in suitable habitat for deep‐sea fishes of 2.0°–9.9° towards higher latitudes. The largest reductions in suitable habitat were projected for the scleractinian coral Lophelia pertusa and the octocoral Paragorgia arborea, with declines of at least 79% and 99% respectively. We projected the expansion of suitable habitat by 2100 only for the fishes Helicolenus dactylopterus and Sebastes mentella (20%–30%), mostly through northern latitudinal range expansion. Our results projected limited climate refugia locations in the North Atlantic by 2100 for scleractinian corals (30%–42% of present‐day suitable habitat), even smaller refugia locations for the octocorals Acanella arbuscula and Acanthogorgia armata (6%–14%), and almost no refugia for P. arborea. Our results emphasize the need to understand how anticipated climate change will affect the distr

Many cold-water sponges harbour microorganisms of which the role in the sponge host remains enigmatic. Here, we show a transfer of fixed inorganic carbon by sponge-associated microbes to its host, the cold-water coral encrusting sponge Hymedesmia (Stylopus) coriacea. Sponge were collected at approx. 100 m depth and incubated for 1.5–2.5 days with 13C labelled dissolved inorganic carbon (DIC) as tracer. Total DIC fixation rates ranged from 0.03–0.11 mmol C × mmol C(sub>sponge × d−1. 13C-tracer was recovered in bacterial-specific (i.e. short and branched) and sponge-specific (very long-chained) phospholipid-derived fatty acids (PLFA's), but was not incorporated into archaeal lipids. 13C-incorporation in biomarkers such as C16:1w7c and C18:1w7c indicated that nitrifying and/or sulphur-oxidizing bacteria (chemoautotrophs) were likely active in the sponge. Trophic transfer of microbially-fixed carbon to the sponge host was confirmed by recovery of label in very long chain fatty acids (VLCFA's) including C26:2 and C26:3. Tracer accumulation into several VLCFA's continued after removal of 13C-DIC, while tracer in most bacteria-specific PLFA's declined, indicating a transfer and elongation of bacterial-specific PLFA's to sponge-specific PLFA's. This implies that PLFA precursors released from chemo- as well as heterotrophic microbes in sponges contributed to the synthesis of VLCFA's, identifying sponge-associated bacteria as symbionts of the sponge.

The deep sea is the largest biome on Earth but the least explored. Our knowledge of it comes from scattered sources spanning different spatial and temporal scales. Implementation of marine policies like the European Union’s Marine Strategy Framework Directive (MSFD) and support for Blue Growth in the deep sea are therefore hindered by lack of data. Integrated assessments of environmental status require tools to work with different and disaggregated datasets (e.g. density of deep-sea habitat-forming species, body-size distribution of commercial fishes, intensity of bottom trawling) across spatial and temporal scales. A feasibility study was conducted as part of the four-year ATLAS project to assess the effectiveness of the open-access Nested Environmental status Assessment Tool (NEAT) to assess deep-sea environmental status. We worked at nine selected study areas in the North Atlantic focusing on five MSFD descriptors (D1-Biodiversity, D3-Commercial fish and shellfish, D4-Food webs, D6-Seafloor integrity, D10-Marine litter). The objectives of the present study were to i) explore and propose indicators that could be used in the assessment of deep-sea environmental status, ii) evaluate the performance of NEAT in the deep sea, and iii) identify challenges and opportunities for the assessment of deep-sea status. Based on data availability, data quality and expert judgement, in total 24 indicators (one for D1, one for D3, seven for D4, 13 for D6, two for D10) were used in the assessment of the nine study areas, their habitats and ecosystem components. NEAT analyses revealed differences among the study areas for their environmental status ranging from “poor” to “high”. Overall, the NEAT results were in moderate to complete agreement with expert judgement, previous assessments, scientific literature on human-pressure gradients and expected management outcomes. We suggest that the assessment of deep-sea environmental status should take place at habitat and ecosystem level (r

Benthic suspension feeders have developed a variety of feeding strategies and food availability has often proven to be a key factor explaining their occurrence and distribution. The feeding biology of coral species has been the target of an increasing number of studies, however most of them focus on Scleractinia and Octocorallia, while information for Antipatharia is very scarce. The present study focused on Antipathella wollastoni, a common habitat-forming antipatharian in the Azores Archipelago, forming dense black coral forests between 20 and 150 m. The objective of the study was to investigate the food preferences of the target species upon availability of different isotopically enriched food substrates and determine its ability to capture zooplankton prey under different flow speeds. The species was able to utilize different food sources including live phytoplankton, live zooplankton and dissolved organic matter (DOM), indicating the ability to exploit seasonally available food sources. However, ingestion of zooplankton enhanced carbon (C) and nitrogen (N) incorporation in coral tissue and metabolic activity, highlighting the importance of zooplankton prey for vital physiological processes such as growth and reproduction. Maximum zooplankton capture rates occurred under 4 cm−1, however the species displayed high capacity to capture zooplankton prey over different flow rates highlighting the ability of A. wollastoni to exploit high quantities of shortly available prey.

Benthic fauna refers to all fauna that live in or on the seafloor, which researchers typically divide into size classes meiobenthos (32/64 µm–0.5/1 mm), macrobenthos (250 µm–1 cm), and megabenthos (>1 cm). Benthic fauna play important roles in bioturbation activity, mineralization of organic matter, and in marine food webs. Evaluating their role in these ecosystem functions requires knowledge of their global distribution and biomass. We therefore established the BenBioDen database, the largest open-access database for marine benthic biomass and density data compiled so far. In total, it includes 11,792 georeferenced benthic biomass and 51,559 benthic density records from 384 and 600 studies, respectively. We selected all references following the procedure for systematic reviews and meta-analyses, and report biomass records as grams of wet mass, dry mass, or ash-free dry mass, or carbon per m2 and as abundance records as individuals per m2. This database provides a point of reference for future studies on the distribution and biomass of benthic fauna., European Union (European Union), EMODnet Thematic Lot n° 5 Biology; European Union’ (European Union); Horizon 2020; Research and Innovation Programme; JPI Oceans – Impacts of deep-sea nodule mining project “MiningImpact 2”; Dutch Research Council; Dutch Research Council, Research Programme NWO-Rubicon

Cold-water coral (CWC) reefs are distributed globally and form complex three-dimensional structures on the deep seafloor, providing habitat for numerous species. Here, we measured the community O2 and dissolved inorganic nitrogen (DIN) flux of CWC reef habitats with different coral cover and bare sediment (acting as reference site) in the Logachev mound area (NE Atlantic). Two methodologies were applied: the non-invasive in situaquatic eddy co-variance (AEC) technique, and ex situ whole box core (BC) incubations. The AEC system was deployed twice per coral mound (69 h in total), providing an integral estimate of the O2 flux from a total reef area of up to 500 m2, with mean O2 consumption rates ranging from 11.6 ± 3.9 to 45.3 ± 11.7 mmol Om–2 d–1 (mean ± SE). CWC reef community Ofluxes obtained from the BC incubations ranged from 5.7 ± 0.3 to 28.4 ± 2.4 mmol Om–2 d–1 (mean ± SD) while the Oflux measured by BC incubations on the bare sediment reference site reported 1.9 ± 1.3 mmol Om–2 d–1 (mean ± SD). Overall, Ofluxes measured with AEC and BC showed reasonable agreement, except for one station with high habitat heterogeneity. Our results suggest Ofluxes of CWC reef communities in the North East Atlantic are around five times higher than of sediments from comparable depths and living CWCs are driving the increased metabolism. DIN flux measurements by the BC incubations also revealed around two times higher DIN fluxes at the CWC reef (1.17 ± 0.87 mmol DIN m–2 d–1), compared to the bare sediment reference site (0.49 ± 0.32 mmol DIN m–2 d–1), due to intensified benthic release of NH4+. Our data indicate that the amount of living corals and dead coral framework largely contributes to the observed variability in Ofluxes on CWC reefs. A conservative estimate, based on the

Sediment community oxygen consumption (SCOC) rates provide important information about biogeochemical processes in marine sediments and the activity of benthic microorganisms and fauna. Therefore, several databases of SCOC data have been compiled since the mid-1990s. However, these earlier databases contained much less data records and were not freely available. Additionally, the databases were not transparent in their selection procedure, so that other researchers could not assess the quality of the data. Here, we present the largest, best documented, and freely available database of SCOC data compiled to date. The database is comprised of 3,540 georeferenced SCOC records from 230 studies that were selected following the procedure for systematic reviews and meta-analyses. Each data record states whether the oxygen consumption was measured ex situ or in situ, as total oxygen uptake, diffusive or advective oxygen uptake, and which measurement device was used. The database will be curated and updated annually to secure and maintain an up-to-date global database of SCOC data.

Free-living nematodes, an ancient animal phylum of unsegmented microscopic roundworms, have successfully adapted to nearly every ecosystem on Earth: from marine and freshwater to land, from the polar regions to the tropics, and from the mountains to the ocean depths. They are globally the most abundant animals in sediments and soils. In the present article, we identify the factors that collectively explain the successful ecological proliferation of free-living nematodes and demonstrate the impact they have on vital sediment and soil processes. The ecological success of nematodes is strongly linked to their ability to feed on various food sources that are present in both sediments and soils, and to proliferate rapidly and survive in contrasting environmental conditions. The adaptations, roles, and behaviors of free-living nematodes have important implications for the resilience of sediments and soils, and for emergent animal communities responding to human alterations to ecosystems worldwide.