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Coccolithophores are globally important marine calcifying phytoplankton that utilize a haplo-diplontic life cycle. The haplo-diplontic life cycle allows coccolithophores to divide in both life cycle phases and potentially expands coccolithophore niche volume. Research has, however, to date largely overlooked the life cycle of coccolithophores and has instead focused on the diploid life cycle phase of coccolithophores. Through the synthesis and analysis of global scanning electron microscopy (SEM) coccolithophore abundance data (n=2534), we find that calcified haploid coccolithophores generally constitute a minor component of the total coccolithophore abundance (≈ 2 %–15 % depending on season). However, using case studies in the Atlantic Ocean and Mediterranean Sea, we show that, depending on environmental conditions, calcifying haploid coccolithophores can be significant contributors to the coccolithophore standing stock (up to ≈30 %). Furthermore, using hypervolumes to quantify the niche of coccolithophores, we illustrate that the haploid and diploid life cycle phases inhabit contrasting niches and that on average this allows coccolithophores to expand their niche by ≈18.8 %, with a range of 3 %–76 % for individual species. Our results highlight that future coccolithophore research should consider both life cycle stages, as omission of the haploid life cycle phase in current research limits our understanding of coccolithophore ecology. Our results furthermore suggest a different response to nutrient limitation and stratification, which may be of relevance for further climate scenarios. Our compilation highlights the spatial and temporal sparsity of SEM measurements and the need for new molecular techniques to identify uncalcified haploid coccolithophores. Our work also emphasizes the need for further work on the carbonate chemistry niche of the coccolithophore life cycle.

In this paper we provide an overview of new knowledge on oxygen depletion (hypoxia) and related phenomena in aquatic systems resulting from the EU-FP7 project HYPOX ("In situ monitoring of oxygen depletion in hypoxic ecosystems of coastal and open seas, and landlocked water bodies", http://www.hypox.net). In view of the anticipated oxygen loss in aquatic systems due to eutrophication and climate change, HYPOX was set up to improve capacities to monitor hypoxia as well as to understand its causes and consequences. Temporal dynamics and spatial patterns of hypoxia were analyzed in field studies in various aquatic environments, including the Baltic Sea, the Black Sea, Scottish and Scandinavian fjords, Ionian Sea lagoons and embayments, and Swiss lakes. Examples of episodic and rapid (hours) occurrences of hypoxia, as well as seasonal changes in bottom-water oxygenation in stratified systems, are discussed. Geologically driven hypoxia caused by gas seepage is demonstrated. Using novel technologies, temporal and spatial patterns of water-column oxygenation, from basin-scale seasonal patterns to meter-scale sub-micromolar oxygen distributions, were resolved. Existing multidecadal monitoring data were used to demonstrate the imprint of climate change and eutrophication on long-term oxygen distributions. Organic and inorganic proxies were used to extend investigations on past oxygen conditions to centennial and even longer timescales that cannot be resolved by monitoring. The effects of hypoxia on faunal communities and biogeochemical processes were also addressed in the project. An investigation of benthic fauna is presented as an example of hypoxia-devastated benthic communities that slowly recover upon a reduction in eutrophication in a system where naturally occurring hypoxia overlaps with anthropogenic hypoxia. Biogeochemical investigations reveal that oxygen intrusions have a strong effect on the microbially mediated redox cycling of elements. Observations and modeling studies of the sediments demonstrate the effect of seasonally changing oxygen conditions on benthic mineralization pathways and fluxes. Data quality and access are crucial in hypoxia research. Technical issues are therefore also addressed, including the availability of suitable sensor technology to resolve the gradual changes in bottom-water oxygen in marine systems that can be expected as a result of climate change. Using cabled observatories as examples, we show how the benefit of continuous oxygen monitoring can be maximized by adopting proper quality control. Finally, we discuss strategies for state-of-the-art data archiving and dissemination in compliance with global standards, and how ocean observations can contribute to global earth observation attempts.

Changes in marine net primary productivity (PP) and export of particulate organic carbon (EP) are projected over the 21st century with four global coupled carbon cycle-climate models. These include representations of marine ecosystems and the carbon cycle of different structure and complexity. All four models show a decrease in global mean PP and EP between 2 and 20% by 2100 relative to preindustrial conditions, for the SRES A2 emission scenario. Two different regimes for productivity changes are consistently identified in all models. The first chain of mechanisms is dominant in the low- and mid-latitude ocean and in the North Atlantic: reduced input of macro-nutrients into the euphotic zone related to enhanced stratification, reduced mixed layer depth, and slowed circulation causes a decrease in macro-nutrient concentrations and in PP and EP. The second regime is projected for parts of the Southern Ocean: an alleviation of light and/or temperature limitation leads to an increase in PP and EP as productivity is fueled by a sustained nutrient input. A region of disagreement among the models is the Arctic, where three models project an increase in PP while one model projects a decrease. Projected changes in seasonal and interannual variability are modest in most regions. Regional model skill metrics are proposed to generate multi-model mean fields that show an improved skill in representing observation-based estimates compared to a simple multi-model average. Model results are compared to recent productivity projections with three different algorithms, usually applied to infer net primary production from satellite observations.

Macrophytes growing in shallow coastal zones characterised by intense metabolic activity have the capacity to modify pH within their canopy and beyond. We observed diel pH changes in shallow (5–12 m) seagrass (Posidonia oceanica) meadows spanning 0.06 pH units in September to 0.24 units in June. The carbonate system (pH, DIC, and aragonite saturation state (ΩAr)) and O2 within the meadows displayed strong diel variability driven by primary productivity, and changes in chemistry were related to structural parameters of the meadow, in particular, the leaf surface area available for photosynthesis (LAI). LAI was positively correlated to mean, max and range pHNBS and max and range ΩAr. In June, vertical mixing (as Turbulent Kinetic Energy) influenced max and min ΩAr, while in September there was no effect of hydrodynamics on the carbonate system within the canopy. Max and range ΩAr within the meadow showed a positive trend with the calcium carbonate load of the leaves, pointing to a possible link between structural parameters, ΩAr and carbonate deposition. Calcifying organisms, e.g. epiphytes with carbonate skeletons, may benefit from the modification of the carbonate system by the meadow. There is, however, concern for the ability of seagrasses to provide modifications of similar importance in the future. The predicted decline of seagrass meadows may alter the scope for alteration of pH within a seagrass meadow and in the water column above the meadow, particularly if shoot density and biomass decline, on which LAI is based. Organisms associated with seagrass communities may therefore suffer from the loss of pH buffering capacity in degraded meadows.

The Black Sea northwestern shelf (NWS) is a shallow eutrophic area in which the seasonal stratification of the water column isolates the bottom waters from the atmosphere. This prevents ventilation from counterbalancing the large consumption of oxygen due to respiration in the bottom waters and in the sediments, and sets the stage for the development of seasonal hypoxia. A three-dimensional (3-D) coupled physical–biogeochemical model is used to investigate the dynamics of bottom hypoxia in the Black Sea NWS, first at seasonal and then at interannual scales (1981–2009), and to differentiate its driving factors (climatic versus eutrophication). Model skills are evaluated by a quantitative comparison of the model results to 14 123 in situ oxygen measurements available in the NOAA World Ocean and the Black Sea Commission databases, using different error metrics. This validation exercise shows that the model is able to represent the seasonal and interannual variability of the oxygen concentration and of the occurrence of hypoxia, as well as the spatial distribution of oxygen-depleted waters. During the period 1981–2009, each year exhibits seasonal bottom hypoxia at the end of summer. This phenomenon essentially covers the northern part of the NWS – which receives large inputs of nutrients from the Danube, Dniester and Dnieper rivers – and extends, during the years of severe hypoxia, towards the Romanian bay of Constanta. An index H which merges the aspects of the spatial and temporal extension of the hypoxic event is proposed to quantify, for each year, the intensity of hypoxia as an environmental stressor. In order to explain the interannual variability of H and to disentangle its drivers, we analyze the long time series of model results by means of a stepwise multiple linear regression. This statistical model gives a general relationship that links the intensity of hypoxia to eutrophication and climate-related variables. A total of 82% of the interannual variability of H is explained by the combination of four predictors: the annual riverine nitrate load (N), the sea surface temperature in the month preceding stratification (Ts), the amount of semi-labile organic matter accumulated in the sediments (C) and the sea surface temperature during late summer (Tf). Partial regression indicates that the climatic impact on hypoxia is almost as important as that of eutrophication. Accumulation of organic matter in the sediments introduces an important inertia in the recovery process after eutrophication, with a typical timescale of 9.3 yr. Seasonal fluctuations and the heterogeneous spatial distribution complicate the monitoring of bottom hypoxia, leading to contradictory conclusions when the interpretation is done from different sets of data. In particular, it appears that the recovery reported in the literature after 1995 was overestimated due to the use of observations concentrated in areas and months not typically affected by hypoxia. This stresses the urgent need for a dedicated monitoring effort in the Black Sea NWS focused on the areas and months concerned by recurrent hypoxic events.

The present study provides new knowledge about the so far largely unexplored Coral Patch seamount which is located in the NE Atlantic Ocean half-way between the Iberian Peninsula and Madeira. For the first time a detailed hydroacoustic mapping (MBES) in conjunction with video surveys (ROV, camera sled) were performed to describe the sedimentological and biological characteristics of this sub-elliptical ENE-WSW elongated seamount. Video observations were restricted to the southwestern summit area of Coral Patch seamount (water depth: 560–760 m) and revealed that this part of the summit is dominated by exposed hard substrate, whereas soft sediment is just a minor substrate component. Although exposed hardgrounds are dominant for this summit area and, thus, offer suitable habitat for settlement by benthic organisms, the benthic megafauna shows rather scarce occurrence. In particular, scleractinian framework-building cold-water corals are apparently rare with very few isolated and small-sized live occurrences of the species Lophelia pertusa and Madrepora oculata. In contrast, dead coral framework and coral rubble are more frequent pointing to a higher abundance of cold-water corals on Coral Patch during the recent past. This is even supported by the observation of fishing lines that got entangled with rather fresh-looking coral frameworks. Overall, long lines and various species of commercially important fish were frequently observed emphasising the potential of Coral Patch as an important target for fisheries that may have impacted the entire benthic community. Hydroacoustic seabed classification covered the entire summit of Coral Patch and its northern and southern flanks (water depth: 560–2660 m) and revealed extended areas dominated by mixed and soft sediments at the northern flank and to a minor degree at its easternmost summit and southern flank. Nevertheless, these data also predict most of the summit area to be dominated by exposed bedrock which would offer suitable habitat for benthic organisms. By comparing the locally restricted video observations and the broad-scale monitoring of a much larger and deeper seafloor area as derived by hydroacoustic seabed classification, it becomes obvious that habitat information obtained by in situ sampling may provide a rather scattered pattern about the entire seamount ecosystem. Solely with a combination of both methods, a satisfactory approach to describe the diverse characteristics of a seamount ecosystem can be derived which is in turn indispensable for future scientific monitoring campaigns as well as management and conservation purposes.

Submarine canyons have often been identified as hotspots of secondary production with the potential to house distinct faunal assemblages and idiosyncratic ecosystems. Within these deep-sea habitats, assemblages of scavenging fauna play a vital role in reintroducing organic matter from large food falls into the wider deep-sea food chain. Free-fall baited traps were set at different depths within three submarine canyons on the Iberian Margin. Amphipods from the traps were identified to species level and counted. Scavenging amphipod assemblages were compared at different depths within each canyon and between individual canyon systems. Using data from literature, abyssal plain assemblages were compared to submarine canyon assemblages. Samples from canyons were found to contain common abyssal plain species but in greater than expected abundances. It is proposed that this is a result of the high organic carbon input into canyon systems owing to their interception of sediment from the continental shelf and input from associated estuarine systems. Community composition differed significantly between the submarine canyons and abyssal plains. The cause of this difference cannot be attributed to one environmental variable due to the numerous inherent differences between canyons and abyssal plains.

Bivalve shells can provide excellent archives of past environmental change but have not been used to interpret ocean acidification events. We investigated carbon, oxygen and trace element records from different shell layers in the mussels Mytilus galloprovincialis combined with detailed investigations of the shell ultrastructure. Mussels from the harbour of Ischia (Mediterranean, Italy) were transplanted and grown in water with mean pHT 7.3 and mean pHT 8.1 near CO2 vents on the east coast of the island. Most prominently, the shells recorded the shock of transplantation, both in their shell ultrastructure, textural and geochemical record. Shell calcite, precipitated subsequently under acidified seawater responded to the pH gradient by an in part disturbed ultrastructure. Geochemical data from all test sites show a strong metabolic effect that exceeds the influence of the low-pH environment. These field experiments showed that care is needed when interpreting potential ocean acidification signals because various parameters affect shell chemistry and ultrastructure. Besides metabolic processes, seawater pH, factors such as salinity, water temperature, food availability and population density all affect the biogenic carbonate shell archive.

Interest in seamount research has gathered momentum over the past five years in an effort to understand the physical, geochemical and biological characteristics as well as the interconnectedness of seamount ecosystems. The majority of biological seamount research has concentrated upon the rich and diverse suspension feeding organisms that dominate the megafauna, such as gorgonians and antipatharian corals; by comparison there have been few studies that have investigated the no less enigmatic, but possibly just as important infauna. To help fill this knowledge gap, the macrofaunal community was sampled from a total of five stations along a northerly transect (capturing water depths from ∼130 m to ∼3300 m), on Senghor Seamount (NE Atlantic). The focus of this study is on the polychaete communities. Polychaete abundance peaked at the summit and a mid-slope station (∼1500 m), a pattern mirrored by the biomass values. The polychaete community along the transect appeared to be particularly diverse, with 135 species nominally identified to putative species from a total of 954 individuals. A diversity maximum was identified on the upper slope at ∼800 m depth, with species diversity, richness and evenness also all peaking at this station. Depth is likely to be a significant factor in determining levels of similarity between stations.

Climate change and changing nutrient loadings are the two main aspects of global change that are linked to the increase in the prevalence of coastal hypoxia – the depletion of oxygen in the bottom waters of coastal areas. However, it remains uncertain how strongly these two drivers will each increase the risk of hypoxia over the next decades. Through model simulations we have investigated the relative influence of climate change and nutrient run-off on the bottom water oxygen dynamics in the Oyster Grounds, an area in the central North Sea experiencing summer stratification. Simulations were performed with a one-dimensional ecosystem model that couples hydrodynamics, pelagic biogeochemistry and sediment diagenesis. Climatological conditions for the North Sea over the next 100 yr were derived from a global-scale climate model. Our results indicate that changing climatological conditions will increase the risk of hypoxia. The bottom water oxygen concentration in late summer is predicted to decrease by 24 μM or 11.5% in the year 2100. More intense stratification is the dominant factor responsible for this decrease (58%), followed by the reduced solubility of oxygen at higher water temperature (27%), while the remaining part could be attributed to enhanced metabolic rates in warmer bottom waters (15%). Relative to these climate change effects, changes in nutrient runoff are also important and may even have a stronger impact on the bottom water oxygenation. Decreased nutrient loadings strongly decrease the probability of hypoxic events. This stresses the importance of continued eutrophication management in coastal areas, which could function as a mitigation tool to counteract the effects of rising temperatures.