Search

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.

Highlights • Total modeled carbon cycling at disturbed sites is lower than at reference sites. • Projected microbial loop functioning is reduced 26 years after sediment disturbance. • Estimated faunal respiration has recovered from sediment disturbance. • Estimated microbial respiration has not recovered from the sediment disturbance. Abstract Due to the predicted future demand for critical metals, abyssal plains covered with polymetallic nodules are currently being prospected for deep-seabed mining. Deep-seabed mining will lead to significant sediment disturbance over large spatial scales and for extended periods of time. The environmental impact of a small-scale sediment disturbance was studied during the ‘DISturbance and reCOLonization’ (DISCOL) experiment in the Peru Basin in 1989 when 10.8 km2 of seafloor were ploughed with a plough harrow. Here, we present a detailed description of carbon-based food-web models constructed from various datasets collected in 2015, 26 years after the experiment. Detailed observations of the benthic food web were made at three distinct sites: inside 26-year old plough tracks (IPT, subjected to direct impact from ploughing), outside the plough tracks (OPT, exposed to settling of resuspended sediment), and at reference sites (REF, no impact). The observations were used to develop highly-resolved food-web models for each site that quantified the carbon (C) fluxes between biotic (ranging from prokaryotes to various functional groups in meio-, macro-, and megafauna) and abiotic (e.g. detritus) compartments. The model outputs were used to estimate total system throughput, i.e., the sum of all C flows in the food web (the ‘ecological size’ of the system), and microbial loop functioning, i.e., the C-cycling through the prokaryotic compartment for each site. Both the estimated total system throughput and the microbial loop cycling were significantly reduced (by 16% and 35%, respectively) inside the plough tracks compared to the other two sites. Site differences in modelled faunal respiration varied among the different faunal compartments. Overall, modelled faunal respiration appeared to have recovered to, or exceeded reference values after 26-years. The model results indicate that food-web functioning, and especially the microbial loop, have not recovered from the disturbance that was inflicted on the abyssal site 26 years ago.

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 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 (rather than at species level) and at relatively large spatial scales, in comparison to shallow-water areas. Limited knowledge across space (e.g. distribution of habitat-forming species) and the scarcity of long-term data sets limit our knowledge about natural variability and human impacts in the deep sea preventing a more systematic assessment of habitat and ecosystem components in the deep sea. However, stronger cross-sectoral collaborations, the use of novel technologies and open data-sharing platforms will be critical for establishing environmental baseline indicator values in the deep sea that will contribute to the science base supporting the implementation of marine policies and stimulating Blue Growth., Sí

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 distribution of deep-sea species including commercially important fishes and foundation species, and highlight the importance of identifying and preserving climate refugia for a range of area-based planning and management tools., ATLAS, A Trans-AtLantic Assessment and deep-water ecosystem-based Spatial management plan for Europe, SI

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.

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.

The cycling of carbon (C) by benthic organisms is a key ecosystem function in the deep sea. Pulse-chase experiments are designed to quantify this process, yet few studies have been carried out using abyssal (3500-6000 m) sediments and only a handful of studies have been undertaken in situ. We undertook eight in situ pulse-chase experiments in three abyssal strata (4050-4200 m water depth) separated by tens to hundreds of kilometers in the eastern Clarion-Clipperton Fracture Zone (CCFZ). These experiments demonstrated that benthic bacteria dominated the consumption of phytodetritus over short (similar to 1.5 d) time scales, with metazoan macrofauna playing a minor role. These results contrast with the only other comparable in situ abyssal study, where macrofauna dominated phytodetritus assimilation over short (2.5 d) time scales in the eutrophic NE Atlantic. We also demonstrated that benthic bacteria were capable of converting dissolved inorganic C into biomass and showed that this process can occur at rates that are as high as the bacterial assimilation of algal-derived organic C. This demonstrates the potential importance of inorganic C uptake to abyssal ecosystems in this region. It also alludes to the possibility that some of the C incorporation by bacteria in our algal-addition studies may have resulted from the incorporation of labeled dissolved inorganic carbon initially respired by other unstudied organisms. Our findings reveal the key importance of benthic bacteria in the short-term cycling of C in abyssal habitats in the eastern CCFZ and provide important information on benthic ecosystem functioning in an area targeted for commercial-scale, deep-sea mining activities.

Mining polymetallic nodules on abyssal plains will have adverse impacts on deep‐sea ecosystems, but it is largely unknown whether the impacted ecosystem will recover, and if so at what rate. In 1989 the “DISturbance and reCOLonization” (DISCOL) experiment was conducted in the Peru Basin where the seafloor was disturbed with a plough harrow construction to explore the effect of small‐scale sediment disturbance from deep‐sea mining. Densities of Holothuroidea in the region were last investigated 7 yr post‐disturbance, before 19 yr later, the DISCOL site was re‐visited in 2015. An “ocean floor observatory system” was used to photograph the seabed across ploughed and unploughed seafloor and at reference sites. The images were analyzed to determine the Holothuroidea population density and community composition, which were combined with in situ respiration measurements of individual Holothuroidea to generate a respiration budget of the study area. For the first time since the experimental disturbance, similar Holothuroidea densities were observed at the DISCOL site and at reference sites. The Holothuroidea assemblage was dominated by Amperima sp., Mesothuria sp., and Benthodytes typica, together contributing 46% to the Holothuroidea population density. Biomass and respiration were similar among sites, with a Holothuroidea community respiration of 5.84 10−4 ± 8.74 10−5 mmol C m−2 d−1 at reference sites. Although these results indicate recovery of Holothuroidea, extrapolations regarding recovery from deep‐sea mining activities must be made with caution: results presented here are based on a relatively small‐scale disturbance experiment as compared to industrial‐scale nodule mining, and also only represent one taxonomic class of the megafauna.

The trophic structure of cold-water coral reef communities at two contrasting locations, the 800-m deep Belgica Mounds (Irish margin) and the 300-m deep Traena reefs (Norwegian Shelf), was investigated using stable isotope (δ13C and δ15N) and fatty-acid composition analysis. A broad range of specimens, with emphasis on (commercial) fish specie's, and organic matter sources were sampled using a variety of tools. Irrespective of the environmental and geographical setting, the δ15N values indicated that the food web encompasses roughly 1.5 to 3 trophic levels. Mobile echinoderms, i.e. sea urchins and sea stars, had highest δ15N values, indicative of a high trophic position in the food web. The fraction of bacterial fatty acids in reef fauna was generally low (

Cold-water coral (CWC) reefs are distributed globally and form complex three-dimensional structures on the deep seafloor, providing habitat for numerous species. Biogeochemical cycling and community metabolism of CWC reefs have mostly been studied by targeting sub-sampled organisms (e.g. corals, sponges). 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 situ aquatic 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) for a total of 69 h, 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 O2 m-2 d-1 (mean ± SE). Six box cores from two coral mounds were incubated onboard for ~24h to quantify O2 and DIN fluxes. The CWC reef community O2 fluxes obtained from the BC incubations ranged from 5.7 ± 0.3 to 28.4 ± 2.4 mmol O2 m-2 d-1 (mean ± SD) while the O2 flux measured by BC incubations on the bare sediment reference site reported 1.9 ± 1.3 mmol O2 m-2 d-1 (mean ± SD). Overall, O2 fluxes measured with AEC and BC showed reasonable agreement, except for one station with high habitat heterogeneity. Our results suggest O2 fluxes 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+.Parallel measurements of the DIN flux in the BC incubations also revealed intensified benthic release of NH4+, which gets partly nitrified, 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). Our data indicate that the amount of living corals and dead coral framework largely contributes to the observed variability in O2 fluxes on CWC reefs. A conservative estimate, based on the measured O2 and DIN fluxes, indicates that CWC reefs process 20% to 35% of the total benthic respiration on the southeasterly Rockall Bank area, which demonstrates that CWC reefs are important to carbon and nitrogen mineralization at the habitat scale.

We investigated food supply mechanisms to a cold-water coral (CWC) reef at 260 m depth on the Norwegian continental shelf using data from a cabled ocean observatory equipped with Acoustic Doppler Current Profilers (ADCPs), an echosounder, and sensors for chlorophyll, turbidity and hydrography in the benthic boundary layer (BBL). Tidal currents of up to tens of cm s-1 dominated BBL hydrodynamics while residual currents were weak (~10 cm s-1), emphasizing a supply and high retention of locally produced phytodetritus within the trough. A direct connection between the reefs and surface organic matter (OM) was established by turbulent mixing and passive particle settling, but relative contributions varied seasonally. Fresh OM from a spring-bloom was quickly mixed into the BBL, but temperature stratification in summer reduced the surface-to-bottom connectivity and reduced the phytodetritus supply. A qualitative comparison among acoustic backscatter in the ADCPs (600 kHz, 190 kHz) and echosounder (70 kHz) suggests that vertically migrating zooplankton may present an alternative food source in summer. Nocturnal feeding by zooplankton in the upper water column sustains downward OM transport independent from water column mixing and may dominate as food supply pathway over sedimentation of the phytodetritus, especially during stratified conditions. In addition, it could present a concentrating mechanism for nutritional components as compensation for the deteriorating phytodetritus quality. Overall, the observed patterns suggest seasonal changes in the food supply pathways to the reef communities. The moderating role of temperature stratification in phytodetritus transport suggests stronger dependence of the cold-water corals on zooplankton for their dietary requirements with increased stratification under future climate scenarios. This study demonstrates the added value of permanent ocean observatories to research based on dedicated campaigns and regular monitoring.

Cold‐water coral (CWC) reefs are hotspots of biodiversity and productivity in the deep sea, but their distribution is limited by the availability of food, which undergoes complex local and temporal variability. We studied the resource utilization, metabolism, and tissue storage of CWC Lophelia pertusa during an experimentally simulated 3‐day food pulse, of 13C5N‐enriched phytodetritus, followed by a 4‐week food deprivation. Oxygen consumption (0.145 μmol O2 [mmol organic carbon {OC}]−1 h−1), release of particulate organic matter (0.029 μmol particulate organic carbon [POC] [mmol OC]−1 h−1 and 0.005 μmol particulate organic nitrogen [mmol OC]−1 h−1), ammonium excretion (0.004 μmol NH4+ [mmol OC]−1 h−1), tissue C and N content, and fatty acid (FA) and amino acid composition did not change significantly during the experiment. Metabolization of the labeled phytodetritus, however, underwent distinct temporal dynamics. Initially, L. pertusa preferentially used phytodetritus‐derived C for respiration (2.2 ± 0.36 nmol C [mmol OC]−1 h−1) and mucus production (0.94 ± 0.52 nmol C [mmol OC]−1 h−1), but those tracer fluxes declined exponentially to 13C in individual FAs revealed a mismatch between the FAs incorporated from phytodetritus and the FA requirements of the coral. We suggest that feeding on other resources, such as lipid‐rich zooplankton, could fill this deficiency. A release of 10% of their total OC as respired C and POC during the 4‐week food deprivation underlines the importance of regular food pulses for CWC reefs.

Cold-water corals form prominent reef ecosystems along ocean margins that depend on suspended resources produced in surface waters. In this study, we investigated food processing of 13C and 15N labelled bacteria and algae by the cold-water coral Lophelia pertusa. Coral respiration, tissue incorporation of C and N and metabolically derived C incorporation into the skeleton were traced following the additions of different food concentrations (100, 300, 1300 µg C L−1) and two ratios of suspended bacterial and algal biomass (1 : 1, 3 : 1). Respiration and tissue incorporation by L. pertusa increased markedly following exposure to higher food concentrations. The net growth efficiency of L. pertusa was low (0.08 ± 0.03), which is consistent with its slow growth rate. The contribution of algae and bacteria to total coral assimilation was proportional to the food mixture in the two lowest food concentrations, but algae were preferred over bacteria as a food source at the highest food concentration. Similarly, the stoichiometric uptake of C and N was coupled in the low and medium food treatment, but was uncoupled in the high food treatment and indicated a comparatively higher uptake or retention of bacterial carbon as compared to algal nitrogen. We argue that behavioural responses for these small-sized food particles, such as tentacle behaviour, mucus trapping and physiological processing, are more likely to explain the observed food selectivity as compared to physical–mechanical considerations. A comparison of the experimental food conditions to natural organic carbon concentrations above CWC reefs suggests that L. pertusa is well adapted to exploit temporal pulses of high organic matter concentrations in the bottom water caused by internal waves and downwelling events.

The deep ocean benthic environment plays a role in long-term carbon sequestration. Understanding carbon cycling in the deep ocean floor is critical to evaluate the impact of changing climate on the oceanic systems. Linear inverse modeling was used to quantify carbon transfer between compartments in the benthic food web at a long time-series study site in the abyssal northeastern Pacific (Station M). Linear inverse food web models were constructed for three separate years in the time-series when particulate organic carbon (POC) flux was relatively high (1990: 0.63 mean mmol C m?2 d?1), intermediate (1995: 0.24) and low (1996: 0.12). Carbon cycling in all years was dominated by the flows involved in the microbial loop; dissolved organic carbon uptake by microbes (0.80–0.95 mean mmol C m?2 d?1), microbial respiration (0.52–0.61), microbial biomass dissolution (0.09–0.18) and the dissolution of refractory detritus (0.46–0.65). Moreover, the magnitude of carbon flows involved in the microbial loop changed in relation to POC input, with a decline in contribution during the high POC influxes, such as those recently experienced at Station M. Results indicate that during high POC episodic pulses the role of faunal mediated carbon cycling would increase. Semi-labile detritus dominates benthic faunal diets and the role of labile detritus declined with increased total POC input. Linear inverse modeling represents an effective framework to analyze high-resolution time-series data and demonstrate the impact of climate change on the deep ocean carbon cycle in a coastal upwelling system.

Jellyfish blooms have increased in magnitude in several locations around the world, including in fjords.While the factors that promote jellyfish blooms and the impacts of live blooms on marine ecosystems areoften investigated, the post-bloom effects from the sinking and accumulation of dead jellyfish at the seafloorremain poorly known. Here, we quantified the effect of jellyfish deposition on short-term benthic carboncycling dynamics in benthic cores taken from a cold and deep fjord environment. Respiration was measuredand 13C-labeled algae were used as a tracer to quantify how C-flow through the benthic food web wasaffected over 5 d in the presence and absence of jellyfish carcasses. Benthic respiration rates increased rapidly(within 2 h) in the jellyfish-amended cores, and were significantly higher than cores that were supplied withonly labeled phytodetritus between 17 h and 117 h. In the cores that were supplied with only labeled phytodetritus,macrofauna dominated algal-C uptake over the 5 d study. The addition of jellyfish caused a rapidand significant shift in C-uptake dynamics: macrofaunal C-uptake decreased while bacterial C-uptakeincreased relative to the cores supplied with only phytodetritus. Our results suggest that the addition of jellyfishdetritus to the seafloor can rapidly alter benthic biogeochemical cycling, and substantially modify C-flowthrough benthic communities. If our results are representative for other areas, they suggest that jellyfishblooms may have cascading effects for benthic ecosystem functions and services when blooms senesce, suchas enhanced bacterial metabolism and reduced energy transfer to upper trophic levels.

The potential harvest of polymetallic nodules will heavily impact the abyssal, soft sediment ecosystem by removing sediment, hard substrate, and associated fauna inside mined areas. It is therefore important to know whether the ecosystem can recover from this disturbance and if so at which rate. The first objective of this study was to measure recovery of phytodetritus processing by the benthic food web from a sediment disturbance experiment in 1989. The second objective was to determine the role of holothurians in the uptake of fresh phytodetritus by the benthic food web. To meet both objectives, large benthic incubation chambers (CUBEs; 50 × 50 × 50 cm) were deployed inside plow tracks (with and without holothurian presence) and at a reference site (holothurian presence, only) at 4100 m water depth. Shortly after deployment, 13C- and 15N-labeled phytodetritus was injected in the incubation chambers and during the subsequent 3-day incubation period, water samples were taken five times to measure the production of 13C-dissolved inorganic carbon over time. At the end of the incubation, holothurians and sediment samples were taken to determine biomass, densities and incorporation of 13C and 15N into bacteria, nematodes, macrofauna, and holothurians. For the first objective, the results showed that biomass of bacteria, nematodes and macrofauna did not differ between reference sites and plow track sites when holothurians were present. Additionally, meiofauna and macrofauna taxonomic composition was not significantly different between the sites. In contrast, total 13C uptake by bacteria, nematodes and holothurians was significantly lower at plow track sites compared to reference sites, though the number of replicates was low. This result suggests that important ecosystem functions such as organic matter processing have not fully recovered from the disturbance that occurred 26 years prior to our study. For the second objective, the analysis indicated that holothurians incorporated 2.16 × 10−3 mmol labile phytodetritus C m−2 d−1 into their biomass, which is one order of magnitude less as compared to bacteria, but 1.3 times higher than macrofauna and one order of magnitude higher than nematodes. Additionally, holothurians incorporated more phytodetritus carbon per unit biomass than macrofauna and meiofauna, suggesting a size-dependence in phytodetritus carbon uptake.

Sponges are often major players in the functioning of shallow-water ecosystems through their high biomass and high capacity in filter feeding. In comparison, little is known about the feeding and metabolic strategies of deep-sea sponges, although they can also form dense aggregations with high biomass. This situation hinders our understanding about how some sponge species thrive under the often food-limited conditions of the deep sea. In the present study we examined the feeding and metabolic strategies of 1) the massive demosponge Spongosorites coralliophaga, which was recently described as an important ecosystem engineer in cold-water coral reefs (CWCRs) and 2) the anthozoan Parazoanthus anguicomus and the ophiuroid Ophiura ophiura, i.e. two dominant epibionts on S. coralliophaga. All three benthic species have high density at CWCRs of the North-East Atlantic and knowing their feeding strategies facilitates future studies on carbon (C) and nitrogen (N) cycling at CWCRs. The on-board feeding experiments examined the processing of four isotopically-labelled food sources, namely 15N-ammonium chloride, 13C-glucose, 13C/15N-labelled microalgae, 13C/15N-labelled bacteria by S. coralliophaga and its symbiotic bacteria and the epibionts P. anguicomus and O. ophiura from the Mingulay reef complex and the Logachev mound (North-East Atlantic). There were no significant differences among the three species in terms of biomass-specific C and N assimilation rates; however, there were differences among S. coralliophaga, P. anguicomus and O. ophiura in how they processed the food sources and this is may be linked to interspecific variability in metabolic needs. S. coralliophaga preferentially assimilated particulate organic N (PON) over particulate organic C (POC) while this was not the case for P. anguicomus and O. ophiura. We did not detect the 15N tracer in the bacterial biomarker D-Alanine suggesting that the preferential assimilation of N over C in S. coralliophaga was mediated by sponge cells instead of the bacterial symbionts. S. coralliophaga assimilated C and N from all four food sources and this versatile feeding strategy was accompanied by an ability for de novo synthesis of essential and non-essential hydrolysable amino acids (HAAs). We suggest that the recorded feeding and metabolic flexibility of S. coralliophaga plays an important role in the survival of this massive sponge under food-limited conditions in the deep sea.

The extraction of minerals from land-based mines necessitates the disposal of large amounts of mine tailings. Dumping and storage of tailings into the marine environment, such as fjords, is currently being performed without knowing the potential ecological consequences. This study investigated the effect of short-term exposure to different deposition depths of inert iron ore tailings (0.1, 0.5, and 3 cm) and dead subsurface sediment (0.5 and 3 cm) on a deep water (200 m) fjord benthic assemblage in a microcosm experiment. Biotic and abiotic variables were measured to determine structural and functional changes of the benthic community following an 11 and 16 day exposure with tailings and dead sediment, respectively. Structural changes of macrofauna, meiofauna, and bacteria were measured in terms of biomass, density, community composition and mortality while measures of oxygen penetration depth, sediment community oxygen consumption and 13C-uptake and processing by biota revealed changes in the functioning of the system. Burial with mine tailings and natural sediments modified the structure and functioning of the benthic community albeit in a different way. Mine tailings deposition of 0.1 cm and more resulted in a reduced capacity of the benthic community to remineralize fresh 13C-labeled algal material, as evidenced by the reduced sediment community oxygen consumption and uptake rates in all biological compartments. At 3 cm of tailings deposition, it was evident that nematode mortality was higher inside the tailings layer, likely caused by reduced food availability. In contrast, dead sediment addition led to an increase in oxygen consumption and bacterial carbon uptake comparable to control conditions, thereby leaving deeper sediment layers anoxic and in turn causing nematode mortality at 3 cm deposition. This study clearly shows that even small levels (0.1 cm) of instantaneous burial by mine tailings may significantly reduce benthic ecosystem functioning on the short term. Furthermore, it reveals the importance of substrate characteristics and origin when studying the effects of substrate addition on marine benthic fauna. Our findings should alert decision makers when considering approval of new deep-sea tailings placement sites as this technique will have major negative impacts on benthic ecosystem functioning over large areas.

With increasing demand for mineral resources, extraction of polymetallic sulphides at hydrothermal vents, cobalt-rich ferromanganese crusts at seamounts, and polymetallic nodules on abyssal plains may be imminent. Here, we shortly introduce ecosystem characteristics of mining areas, report on recent mining developments, and identify potential stress and disturbances created by mining. We analyze species' potential resistance to future mining and perform meta-analyses on population density and diversity recovery after disturbances most similar to mining: volcanic eruptions at vents, fisheries on seamounts, and experiments that mimic nodule mining on abyssal plains. We report wide variation in recovery rates among taxa, size, and mobility of fauna. While densities and diversities of some taxa can recover to or even exceed pre-disturbance levels, community composition remains affected after decades. The loss of hard substrata or alteration of substrata composition may cause substantial community shifts that persist over geological timescales at mined sites. (C) 2017 Elsevier Ltd. All rights reserved. European Union Seventh Framework Programme (FP7) under the MIDAS project; FCT [IF/00029/2014/CP1230/CT0002, SFRH/ BPD/110278/2015]; Spanish RTD project NUREIEV [CTM2013-44598-R]; Ministry of Economy and Competitiveness [SGR 1068]; Generalitat de Catalunya autonomous government; European Union Horizon research and innovation programme [689518]; Fundacao para a Ciencia e a Tecnologia [UID/MAR/04292/2013]; German Ministry of Research (BMBF) [03F0707A-G]; Program Investigador FCT [IF/01194/2013/CP1199/CT0002] info:eu-repo/semantics/publishedVersion

In order to investigate survival of diatoms and long-term pigment dynamics in dark sediment, we incubated samples of homogenized, sieved, tidal-flat sediment for 1 yr in darkness. Microscopic observations revealed that some diatoms survived the full year in darkness and retained their pigments. Concentrations of the diatomspecific phospholipid-derived fatty acid (PLFA) 20:5v3 indicate that these survivors represented only a small fraction of the original diatom community, probably no more than a few percent. Consequently, the effect of pigment retention in living diatoms on the overall pigment dynamics was negligible. The photosynthetic potential of the surviving diatoms was further investigated by re-exposing the sediment to light in the presence of 13Cbicarbonate. Rapid 13C fixation into algal PLFAs at a level similar to that in sediment freshly collected from the field indicates that surviving diatoms fully retained their photosynthetic capacity. Concentration dynamics of photosynthetic pigments generally reflected degradation of two different pools with clearly different loss rates. Losses during the first 1–2 months of the experiment (half lives of 6–22 d) reflected degradation of fresh algal material, while losses during the rest of the experiment reflected much slower degradation of more refractory pigment pools (half lives of $ 1 yr). Individual pigments showed distinctly different labilities, with losses ranging from 0% to 86% over the full year. The order of overall loss was fucoxanthin . chlorophyll c . chlorophyll a . diadinoxanthin . pheophorbide . pheophytin . diatoxanthin. A major fraction of the organic matter that enters marine sediments is derived from algae. For sediments in shallow waters, this organic matter includes material from

A stable isotope (13C)-labeling experiment was performed to quantify the importance of bacterial carbon as a food source for an Arctic deep-sea nematode community. Bacterial functional groups were isotopically enriched with 13C-glucose, 13C-acetate, 13C-bicarbonate, and 13C-amino acids injected into sediments collected from 1280 m depth at 79°N, 6°E, west of Svalbard. Incorporation of the 13C label into bacterial phospholipid-derived fatty acids (PLFAs) and nematodes in the top 5 cm of the sediment was monitored over a 7-d period. The 13C dynamics of nematodes was fitted with a simple isotope turnover model to derive the importance of the different bacterial functional groups as carbon sources for the nematodes. The different substrates clearly labeled different bacterial groups as evidenced by differential labeling of the PLFA patterns. The deep-sea nematode community incorporated a very limited amount of the label, and the isotope turnover model showed that the dynamics of the isotope transfer could not be attributed to bacterivory. The low enrichment of nematodes suggests a limited passive uptake of injected 13C-labeled substrates. The lack of accumulation suggests that the injected 13C-labeled dissolved organic carbon compounds are not important as carbon sources for deep-sea nematodes. Since earlier studies with isotopically enriched algae also found limited uptake by nematodes, the food sources of deep-sea nematodes remain unclear.

Ongoing climate warming is causing a dramatic loss of sea ice in the Arctic Ocean, and it is projected that the Arctic Ocean will become seasonally ice-free by 2040. Many studies of local Arctic food webs now exist, and with this review paper we aim to synthesize these into a large-scale assessment of the current status of knowledge on the structure of various Arctic marine food webs and their response to climate change, and to sea-ice retreat in particular. Key drivers of ecosystem change and potential consequences for ecosystem functioning and Arctic marine food webs are identified along the sea-ice gradient, with special emphasis on the following regions: seasonally ice-free Barents and Chukchi seas, loose ice pack zone of the Polar Front and Marginal Ice Zone, and permanently sea-ice covered High Arctic. Finally, we identify knowledge gaps in different Arctic marine food webs and provide recommendations for future studies. Keywords: Arctic; food web; climate change; sea-ice retreat; trophic transfer; pelagic–benthic coupling. (Published: 20 May 2015) Citation: Polar Research 2015, 34 , 23775,http://dx.doi.org/10.3402/polar.v34.23775

Sediments sampled from the Galicia Bank seamount and the adjacent slope (northeast Atlantic), and from a western Mediterranean slope site, were injected onboard with 13C-enriched dissolved organic matter (DOM) to evaluate nematode feeding strategies and the fate of DOM carbon in different benthic environments. We hypothesized that nematode 13C label assimilation resulted from either direct DOM uptake or feeding on 13C labeled bacteria. Slope sediments were injected with glucose (“simple” DOM) or “complex” diatom-derived DOM to investigate the influence of DOM composition on carbon assimilation.The time-series (1, 7 and 14 days) experiment at the seamount site was the first study to reveal a higher 13C enrichment of nematodes than bacteria and sediments after 7 days. Although isotope dynamics indicated that both DOM and bacteria were plausible candidate food sources, the contribution to nematode secondary production and metabolic requirements (estimated from biomass-dependent respiration rates) was higher for bacteria than for DOM at all sites. The seamount nematode community showed higher carbon assimilation rates than the slope assemblages, which may reflect an adaptation to the food-poor environment. Our results suggested that the trophic importance of bacteria did not depend on the amount of labile sedimentary organic matter. Furthermore, there was a discrepancy between carbon assimilation rates observed in the experiments and the feeding type classification, based on buccal morphology. Sites with a similar feeding type composition (i.e. the northeast Atlantic sites) showed large differences in uptake, whilst the nematode assemblages at the two slope sites, which had a differing trophic structure, took up similar amounts of the DOM associated carbon.Our results did not indicate substantial differences in carbon processing related to the complexity of the DOM substrate. The quantity of processed carbon (5–42% of added DOM) was determined by the bacteria, and was primarily respired. The bulk of the added 13C-DOM was not ingested by the benthic biota under study, and a considerable fraction was possibly adsorbed onto the sediment grains.

The cold-water coral Lophelia pertusa was exposed to suspended particles (

Carbon flows in the benthic food web of the Porcupine Abyssal Plain (4850 m, northeast Atlantic) were reconstructed using linear inverse modeling based on mass balances of the various compartments, biomass data, carbon flux measurements, physiological constraints, and data from a previously published C-13-phytodetritus pulse-chase experiment. Food web compartments consisted of bacteria, Foraminifera, three feeding types of nematodes and of macrofauna, and four feeding types of megafauna. Three lability classes of detritus were defined to clarify especially the possible role of labile detritus in this abyssal plain food web. Total detritus inputs to the food web were 0.56 mmol C m(-2) d(-1), with only a minor (3.8%) contribution of labile detritus. The dominant pathway in the food web was dissolution of more refractory detritus, uptake by bacteria, and subsequent respiration (0.36 mmol C m(-2) d(-1), 80% of the community respiration). Lower contributions to community respiration were found for surface-deposit-feeding macrobenthos (5.6%), Foraminifera (4.9%), predatory macrobenthos (3.8%), and other deposit-feeding macrobenthos (3.1%). Contributions by other benthic compartments were < 1%. C-13-phytodetritus transfer data strongly constrained the carbon source for bacteria, Foraminifera, and the nematode and macrobenthic compartments. Surprisingly, labile detritus contributed < 5% of the total carbon requirements for all these compartments. Carbon sources were much less constrained for the megabenthic compartments because for these compartments no C-13 pulse-chase data were available. The carbon flow model evidently shows a limited role of labile detritus in the food web.

The fate of microbial carbon, nitrogen, hydrolysable amino acids (HAAs), monosaccharides, and fatty acids in sediment was investigated experimentally. The microbial community of a tidal flat sediment was labeled with 13C-enriched glucose and 15N-enriched ammonium, and sediment was incubated for up to 371 days. Analysis of total concentrations and 13C- and 15N content of bulk sediment, hydrolysable amino acids (including d-alanine), monosaccharides, total fatty acids (TFAs), and phospholipid-derived fatty acids (PLFAs) allowed us to trace the fate of microbial biomass and -detritus and the major biochemical groups therein (proteins, carbohydrates, and lipids) over intermediate time scales (weeks–months). Moreover, the unidentified fraction of the labeled material (i.e. not analyzed as HAA, FA, or carbohydrate) provided information on the formation and fate of molecularly uncharacterizable organic matter. Loss of 13C and 15N from the sediment was slow (half live of 433 days) which may have been due to the permanently anoxic conditions in the experiment. Loss rates for the different biochemical groups were also low with the following order of loss rate constants: PLFA > TFA > HAA > monosaccharides. The unidentified 13C-pool was rapidly formed (within days) and then decreased relatively slowly, resulting in a gradual relative accumulation of this pool over time. Degradation and microbial reworking of the labeled material resulted in subtle, yet consistent, diagenetic changes within the different biochemical groups. In the HAA pool, glycine, lysine, and proline were lost relatively slowly (i.e. best preserved) while there was no accumulation of d-alanine relative to l-alanine, indicating no relative accumulation of bacterial macromolecules rich in d-alanine. In the fatty acid pool, there was very little difference between PLFAs and TFAs, indicating a very similar lability of these pools. Differences between individual fatty acids included a relatively slow loss of i15:0, ai15:0 and 18:1ω7c which likely resulted from production of these bacterial fatty acids during bacterial reworking of the organic matter. Differences between loss rate constants for individual monosaccharides were not significant. An exception was ribose that was produced and lost relatively rapidly, which may be related to ribose being an important component of RNA. Losses of bulk 13C and 15N were closely coupled despite partly being present in different biochemicals and partly being derived from different microbial sources, indicating no selective preservation of either C or N during organic matter diagenesis.

The HAUSGARTEN observatory is located in the eastern Fram Strait (Arctic Ocean) and used as long-term monitoring site to follow changes in the Arctic benthic ecosystem. Linear inverse modelling was applied to decipher carbon flows among the compartments of the benthic food web at the central HAUSGARTEN station (2500 m) based on an empirical data set consisting of data on biomass, prokaryote production, total carbon deposition and community respiration. The model resolved 99 carbon flows among 4 abiotic and 10 biotic compartments, ranging from prokaryotes up to megafauna. Total carbon input was 3.78±0.31 mmol C m −2 d −1 , which is a comparatively small fraction of total primary production in the area. The community respiration of 3.26±0.20 mmol C m −2 d −1 is dominated by prokaryotes (93%) and has lower contributions from surface-deposit feeding macro- (1.7%) and suspension feeding megafauna (1.9%), whereas contributions from nematode and other macro- and megabenthic compartments were limited to

Biomass and respiration rates of bacteria, nematodes and macrobenthos were estimated in relation to the deposition of the spring phytoplankton bloom at two contrasting sites in the Southern North Sea: one with fine-grained sediment close to the coastline and another with highly permeable sediments. Sediment community oxygen consumption (SCOC) was also measured. Bacterial biomass was relatively similar at both stations, whereas nematode and macrobenthic biomass were higher in fine-grained sediment. In fine sediments, bacterial biomass increased quickly after deposition of the phytoplankton bloom, whereas the response of nematodes and macrobenthos was delayed. In coarser sediments, nematodes and macrobenthos also showed a fast response in terms of density and biomass. Respiration in permeable sediments was mainly dominated by bacteria at all periods of the year. Hence, nematode and macrobenthic respiration did not contribute strongly to SCOC. This is in contrast to the patterns observed in finer sediments, where both macrofauna and nematodes were important oxygen consumers as well. Macrobenthos contributed more to total SCOC than did nematodes in winter. However, shortly after the arrival of phytodetritus at the sea floor, nematodes and macrobenthos contributed equally to the total SCOC, indicating that all benthic size classes should be taken into account when investigating marine benthic respiration rates.

Cold-water corals (CWCs) form large mounds on the seafloor that are hotspots of biodiversity in the deep sea, but it remains enigmatic how CWCs can thrive in this food-limited environment. Here, we infer from model simulations that the interaction between tidal currents and CWC-formed mounds induces downwelling events of surface water that brings organic matter to 600-m deep CWCs. This positive feedback between CWC growth on carbonate mounds and enhanced food supply is essential for their sustenance in the deep sea and represents an example of ecosystem engineering of unparalleled magnitude. This ’topographically-enhanced carbon pump’ leaks organic matter that settles at greater depths. The ubiquitous presence of biogenic and geological topographies along ocean margins suggests that carbon sequestration through this pump is of global importance. These results indicate that enhanced stratification and lower surface productivity, both expected consequences of climate change, may negatively impact the energy balance of CWCs.