ALBERT

Description: The Atlantic meridional overturning circulation (AMOC) carries large amounts of heat into the North Atlantic influencing climate regionally as well as globally. Palaeo-records and simulations with comprehensive climate models suggest that the positive salt-advection feedback may yield a threshold behaviour of the system. That is to say that beyond a certain amount of freshwater flux into the North Atlantic, no meridional overturning circulation can be sustained. Concepts of monitoring the AMOC and identifying its vicinity to the threshold rely on the fact that the volume flux defining the AMOC will be reduced when approaching the threshold. Here we advance conceptual models that have been used in a paradigmatic way to understand the AMOC, by introducing a density-dependent parameterization for the Southern Ocean eddies. This additional degree of freedom uncovers a mechanism by which the AMOC can increase with additional freshwater flux into the North Atlantic, before it reaches the threshold and collapses: an AMOC that is mainly wind-driven will have a constant upwelling as long as the Southern Ocean winds do not change significantly. The downward transport of tracers occurs either in the northern sinking regions or through Southern Ocean eddies. If freshwater is transported, either atmospherically or via horizontal gyres, from the low to high latitudes, this would reduce the eddy transport and by continuity increase the northern sinking which defines the AMOC until a threshold is reached at which the AMOC cannot be sustained. If dominant in the real ocean this mechanism would have significant consequences for monitoring the AMOC.

Description: In this study we present a comparative quantification of CaCO3 production rates by rhodolith-forming coralline red algal communities situated in high polar latitudes and assess which environmental parameters control these production rates. The present rhodoliths act as ecosystem engineers, and their carbonate skeletons provide an important ecological niche to a variety of benthic organisms. The settings are distributed along the coasts of the Svalbard archipelago, being Floskjeret (78◦180N) in Isfjorden, Krossfjorden (79◦080N) at the eastern coast of Haakon VII Land, Mosselbukta (79◦530N) at the eastern coast of Mosselhalvøya, and Nordkappbukta (80◦310N) at the northern coast of Nordaustlandet. All sites feature Arctic climate and strong seasonality.

Description: Anthropogenic litter is present in all marine habitats, from beaches to the most remote points in the oceans. On the seafloor, marine litter, particularly plastic, can accumulate in high densities with deleterious consequences for its inhabitants. Yet, because of the high cost involved with sampling the seafloor, no large-scale assessment of distribution patterns was available to date. Here, we present data on litter distribution and density collected during 588 video and trawl surveys across 32 sites in European waters. We found litter to be present in the deepest areas and at locations as remote from land as the Charlie-Gibbs Fracture Zone across the Mid-Atlantic Ridge. The highest litter density occurs in submarine canyons, whilst the lowest density can be found on continental shelves and on ocean ridges. Plastic was the most prevalent litter item found on the seafloor. Litter from fishing activities (derelict fishing lines and nets) was particularly common on seamounts, banks, mounds and ocean ridges. Our results highlight the extent of the problem and the need for action to prevent increasing accumulation of litter in marine environments.

Description: Linking lower and higher trophic levels requires special focus on the essential role played by mid-trophic levels, i.e., the zooplankton. One of the most relevant pieces of information regarding zooplankton in terms of flux of energy lies in its size structure. In this study, an extensive data set of size measurements is presented, covering parts of the western European continental shelf and slope, from the Galician coast to the Ushant front, during the springs from 2005 to 2012. Zooplankton size spectra were estimated using measurements carried out in situ with the Laser Optical Plankton Counter (LOPC) and with an image analysis of WP2 net samples (200 μm mesh size) performed following the ZooScan methodology. The LOPC counts and sizes particles within 100–2000 μm of spherical equivalent diameter (ESD), whereas the WP2/ZooScan allows for counting, sizing and identification of zooplankton from ~ 400 μm ESD. The difference between the LOPC (all particles) and the WP2/ZooScan (zooplankton only) was assumed to provide the size distribution of non-living particles, whose descriptors were related to a set of explanatory variables (including physical, biological and geographic descriptors). A statistical correction based on these explanatory variables was further applied to the LOPC size distribution in order to remove the non-living particles part, and therefore estimate the size distribution of zooplankton. This extensive data set provides relevant information about the zooplankton size distribution variability, productivity and trophic transfer efficiency in the pelagic ecosystem of the Bay of Biscay at a regional and interannual scale.

Description: Knowledge of calcium phosphate (Ca-P) solubility is crucial for understanding temporal and spatial variations of phosphorus (P) concentrations in water bodies and sedimentary reservoirs. In situ relationships between liquid-and solid-phase levels cannot be fully explained by dissolved analytes alone and need to be verified by determining particular sediment P species. Lack of quantification methods for these species limits the knowledge of the P cycle. To address this issue, we (i) optimized a specifically developed conversion-extraction (CONVEX) method for P species quantification using standard additions, and (ii) simultaneously determined solubilities of Ca-P standards by measuring their pH-dependent contents in the sediment matrix. Ca-P minerals including various carbonate fluorapatite (CFAP) specimens from different localities, fluorapatite (FAP), fish bone apatite, synthetic hydroxylapatite (HAP) and octacalcium phosphate (OCP) were characterized by XRD, Raman, FTIR and elemental analysis. Sediment samples were incubated with and without these reference minerals and then sequentially extracted to quantify Ca-P species by their differential dissolution at pH values between 3 and 8. The quantification of solid-phase phosphates at varying pH revealed solubilities in the following order: OCP〉 HAP〉 CFAP (4.5% CO3)〉 CFAP (3.4% CO3)〉 CFAP (2.2% CO3)〉 FAP. Thus, CFAP was less soluble in sediment than HAP, and CFAP solubility increased with carbonate content. Unspiked sediment analyses together with standard addition analyses indicated consistent differential dissolution of natural sediment species vs. added reference species and therefore verified the applicability of the CONVEX method in separately determining the most prevalent Ca-P minerals. We found surprisingly high OCP contents in the coastal sediments analyzed, which supports the hypothesis of apatite formation by an OCP precursor mechanism.

Description: Dedicated to the memory of our colleague Klaus Hochheim, who tragically lost his life in the Arctic expedition in September 2013. A distinct, subsurface density front along the eastern St. Anna Trough in the northern Kara Sea is inferred from hydrographic observations in 1996 and 2008–2010. Direct velocity measurements show a persistent northward subsurface current (~ 18 cm s−1) along the St. Anna Trough eastern flank. This sheared flow, carrying the outflow from the Barents and Kara seas to the Arctic Ocean, is also evident from shipboard observations as well as from geostrophic velocities and numerical model simulations. Although we cannot substantiate our conclusions by direct observation-based estimates of mixing rates in the area, we hypothesize that the enhanced vertical mixing along the St. Anna Trough eastern flank favors the upward heat loss from the intermediate warm Atlantic water layer. Modeling results support this hypothesis. The upward heat flux inferred from hydrographic data and model simulations is of O(30–100) W m−2. The region of lowered sea ice thickness and concentration seen both in sea ice remote sensing observations and model simulations marks the Atlantic water pathway in the St. Anna Trough and adjacent Nansen Basin continental margin. In fact, the sea ice shows a delayed freeze-up onset during fall and a reduction in the sea ice thickness during winter. This is consistent with our results on the enhanced Atlantic water heat loss along the Atlantic water pathway in the St. Anna Trough.

Description: Following the launch of ESA's Soil Moisture and Ocean Salinity (SMOS) mission, it has been shown that brightness temperatures at a low microwave frequency of 1.4 GHz (L-band) are sensitive to sea ice properties. In the first demonstration study, sea ice thickness up to 50 cm has been derived using a semi-empirical algorithm with constant tie-points. Here, we introduce a novel iterative retrieval algorithm that is based on a thermodynamic sea ice model and a three-layer radiative transfer model, which explicitly takes variations of ice temperature and ice salinity into account. In addition, ice thickness variations within the SMOS spatial resolution are considered through a statistical thickness distribution function derived from high-resolution ice thickness measurements from NASA's Operation IceBridge campaign. This new algorithm has been used for the continuous operational production of a SMOS-based sea ice thickness data set from 2010 on. The data set is compared to and validated with estimates from assimilation systems, remote sensing data, and airborne electromagnetic sounding data. The comparisons show that the new retrieval algorithm has a considerably better agreement with the validation data and delivers a more realistic Arctic-wide ice thickness distribution than the algorithm used in the previous study (Kaleschke et al., 2012).

Description: The first long-term aerosol sampling and chemical characterization results from measurements at the Cape Verde Atmospheric Observatory (CVAO) on the island of São Vicente are presented and are discussed with respect to air mass origin and seasonal trends. In total 671 samples were collected using a high-volume PM10 sampler on quartz fiber filters from January 2007 to December 2011. The samples were analyzed for their aerosol chemical composition, including their ionic and organic constituents. Back trajectory analyses showed that the aerosol at CVAO was strongly influenced by emissions from Europe and Africa, with the latter often responsible for high mineral dust loading. Sea salt and mineral dust dominated the aerosol mass and made up in total about 80% of the aerosol mass. The 5-year PM10 mean was 47.1 ± 55.5 μg m−2, while the mineral dust and sea salt means were 27.9 ± 48.7 and 11.1 ± 5.5 μg m−2, respectively. Non-sea-salt (nss) sulfate made up 62% of the total sulfate and originated from both long-range transport from Africa or Europe and marine sources. Strong seasonal variation was observed for the aerosol components. While nitrate showed no clear seasonal variation with an annual mean of 1.1 ± 0.6 μg m−3, the aerosol mass, OC (organic carbon) and EC (elemental carbon), showed strong winter maxima due to strong influence of African air mass inflow. Additionally during summer, elevated concentrations of OM were observed originating from marine emissions. A summer maximum was observed for non-sea-salt sulfate and was connected to periods when air mass inflow was predominantly of marine origin, indicating that marine biogenic emissions were a significant source. Ammonium showed a distinct maximum in spring and coincided with ocean surface water chlorophyll a concentrations. Good correlations were also observed between nss-sulfate and oxalate during the summer and winter seasons, indicating a likely photochemical in-cloud processing of the marine and anthropogenic precursors of these species. High temporal variability was observed in both chloride and bromide depletion, differing significantly within the seasons, air mass history and Saharan dust concentration. Chloride (bromide) depletion varied from 8.8 ± 8.5% (62 ± 42%) in Saharan-dust-dominated air mass to 30 \textpm 12% (87 ± 11%) in polluted Europe air masses. During summer, bromide depletion often reached 100% in marine as well as in polluted continental samples. In addition to the influence of the aerosol acidic components, photochemistry was one of the main drivers of halogenide depletion during the summer; while during dust events, displacement reaction with nitric acid was found to be the dominant mechanism. Positive matrix factorization (PMF) analysis identified three major aerosol sources: sea salt, aged sea salt and long-range transport. The ionic budget was dominated by the first two of these factors, while the long-range transport factor could only account for about 14% of the total observed ionic mass.

Description: An earth system model of intermediate complexity (CLIMate and BiosphERe – CLIMBER-2) and a land surface model (JSBACH), which dynamically represent vegetation, are used to simulate natural fire dynamics through the last 8000 yr. Output variables of the fire model (burned area and fire carbon emissions) are used to compare model results with sediment-based charcoal reconstructions. Several approaches for processing model output are also tested. Charcoal data are reported in Z-scores with a base period of 8000–200 BP in order to exclude the strong anthropogenic forcing of fire during the last two centuries. The model–data comparison reveals a robust correspondence in fire activity for most regions considered, while for a few regions, such as Europe, simulated and observed fire histories show different trends. The difference between modelled and observed fire activity may be due to the absence of anthropogenic forcing (e.g. human ignitions and suppression) in the model simulations, and also due to limitations inherent to modelling fire dynamics. The use of spatial averaging (or Z-score processing) of model output did not change the directions of the trends. However, Z-score-transformed model output resulted in higher rank correlations with the charcoal Z-scores in most regions. Therefore, while both metrics are useful, processing model output as Z-scores is preferable to areal averaging when comparing model results to transformed charcoal records.

Description: Satellite observations of microwave brightness temperatures between 19 GHz and 85 GHz are the main data sources for operational sea-ice monitoring and retrieval of ice concentrations. However, microwave brightness temperatures depend on the emissivity of snow and ice, which is subject to pronounced seasonal variations and shows significant hemispheric contrasts. These mainly arise from differences in the rate and strength of snow metamorphism and melt. We here use the thermodynamic snow model SNTHERM forced by European Re-Analysis (ERA) interim data and the Microwave Emission Model of Layered Snowpacks (MEMLS), to calculate the sea-ice surface emissivity and to identify the contribution of regional patterns in atmospheric conditions to its variability in the Arctic and Antarctic. The computed emissivities reveal a pronounced seasonal cycle with large regional variability. The emissivity variability increases from winter to early summer and is more pronounced in the Antarctic. In the pre-melt period (January–May, July–November) the standard deviations in surface microwave emissivity due to diurnal, regional and inter-annual variability of atmospheric forcing reach up to Δε = 0.034, 0.043, and 0.097 for 19 GHz, 37 GHz and 85 GHz channels, respectively. Between 2000 and 2009, small but significant positive emissivity trends were observed in the Weddell Sea during November and December as well as in Fram Strait during February, potentially related to earlier melt onset in these regions. The obtained results contribute to a better understanding of the uncertainty and variability of sea-ice concentration and snow-depth retrievals in regions of high sea-ice concentrations.

Description: Sea-ice diatoms are known to accumulate in large aggregates in and under sea ice and in melt ponds. There is recent evidence from the Arctic that such aggregates can contribute substantially to particle export when sinking from the ice. The role and regulation of microbial aggregation in the highly seasonal, nutrient- and light-limited Arctic sea-ice ecosystem is not well understood. To elucidate the mechanisms controlling the formation and export of algal aggregates from sea ice, we investigated samples taken in late summer 2011 and 2012, during two cruises to the Eurasian Basin of the Central Arctic Ocean. Spherical aggregates densely packed with pennate diatoms, as well as filamentous aggregates formed by Melosira arctica showed sign of different stages of degradation and physiological stoichiometries, with carbon to chlorophyll a ratios ranging from 110 to 66700, and carbon to nitrogen molar ratios of 8–35 and 9–40, respectively. Sub-ice algal aggregate densities ranged between 1 and 17 aggregates m−2, maintaining an estimated net primary production of 0.4–40 mg C m−2 d−1, and accounted for 3–80% of total phototrophic biomass and up to 94% of local net primary production. A potential factor controlling the buoyancy of the aggregates was light intensity, regulating photosynthetic oxygen production and the amount of gas bubbles trapped within the mucous matrix, even at low ambient nutrient concentrations. Our data-set was used to evaluate the distribution and importance of Arctic algal aggregates as carbon source for pelagic and benthic communities.

Description: The candidate phylum Poribacteria is one of the most dominant and widespread members of the microbial communities residing within marine sponges. Cell compartmentalization had been postulated along with their discovery about a decade ago and their phylogenetic association to the Planctomycetes, Verrucomicrobia, Chlamydiae superphylum was proposed soon thereafter. In the present study we revised these features based on genomic data obtained from six poribacterial single cells. We propose that Poribacteria form a distinct monophyletic phylum contiguous to the PVC superphylum together with other candidate phyla. Our genomic analyses supported the possibility of cell compartmentalization in form of bacterial microcompartments. Further analyses of eukaryote-like protein domains stressed the importance of such proteins with features including tetratricopeptide repeats, leucin rich repeats as well as low density lipoproteins receptor repeats, the latter of which are reported here for the first time from a sponge symbiont. Finally, examining the most abundant protein domain family on poribacterial genomes revealed diverse phyH family proteins, some of which may be related to dissolved organic posphorus uptake.

Description: Epibenthos communities play an important role in the marine ecosystems of the Weddell Sea. Information on the factors controlling their structure and distribution are, however, still rare. In particular, the interactions between environmental factors and biotic assemblages are not fully understood. Nachtigaller Hill, a newly discovered seabed structure on the over-deepened shelf of the northwest Weddell Sea (Southern Ocean), offers a unique site to study these interactions in a high-latitude Antarctic setting. Based on high-resolution bathymetry and georeferenced biological data, the effect of the terrain and related environmental parameters on the epibenthos was assessed. At Nachtigaller Hill, both geomorphological and biological data showed complex distribution patterns, reflecting local processes such as iceberg scouring and locally amplified bottom currents. This variability was also generally reflected in the variable epibenthos distribution patterns although statistical analyses did not show strong correlations between the selected environmental parameters and species abundances. By analysing the interactions between environmental and biological patterns, this study provides crucial information towards a better understanding of the factors and processes that drive epibenthos communities on the shelves of the Weddell Sea and probably also on other Antarctic shelves.

Description: The response of coccolithophore calcification to ocean acidification has been studied in culture experiments as well as in present and past oceans. The response, however, is different between species and strains, and for the relatively small carbonate chemistry changes observed in natural environments, a uniform response of the entire coccolithophore community has not been documented so far. Moreover, previous palaeo-studies basically focus on changes in coccolith weight due to increasing CO2 and the resulting changes in the carbonate system, and only few studies focus on the influence of other environmental factors. In order to untangle changes in coccolithophore calcification due to environmental factors such as temperature and/or productivity from changes caused by increasing pCO2 and decreasing carbonate ion concentration, we here present a study on coccolith calcification from the Holocene North Atlantic Ocean. The pre-industrial Holocene, with its predominantly stable atmospheric CO2, provides the conditions for such a comprehensive analysis. For an analysis on changes in major components of Holocene coccolithophores under natural conditions, the family Noelaerhabdaceae was selected, which constitutes the main part of the assemblage in the North Atlantic. Records of average coccolith weights from three Holocene sediment cores along a north–south transect in the North Atlantic were analysed. During the Holocene, mean weight (and therefore calcification) of Noelaerhabdaceae (Emiliania huxleyi and Gephyrocapsa) coccoliths decreased at the Azores (Geofar KF 16) from around 7 to 6 pg, but increased at the Rockall Plateau (ODP site 980) from around 6 to 8 pg, and at the Vøring Plateau (MD08-3192) from 7 to 10 pg. The amplitude of average weight variability is within the range of glacial–interglacial changes that were interpreted to be an effect of decreasing carbonate ion concentration. By comparison with SEM assemblage counts, we show that weight changes are not only partly due to variations in the coccolithophore assemblage but also an effect of a change in calcification and/or morphotype variability within single species. Our results indicate that there is no single key factor responsible for the observed changes in coccolith weight. A major increase in coccolith weight occurs during a slight decrease in carbonate ion concentration in the late Holocene at the Rockall Plateau and Vøring Plateau. Here, more favourable productivity conditions apparently lead to an increase in coccolith weight, either due to the capability of coccolithophore species, especially E. huxleyi, to adapt to decreasing carbonate ion concentration or due to a shift towards heavier calcifying morphotypes.

Description: Four sediment cores from the central and northern Greenland Sea basin, a crucial area for the renewal of North Atlantic deep water, were analyzed for planktic foraminiferal fauna, planktic and benthic stable oxygen and carbon isotopes as well as ice-rafted debris to reconstruct the environmental variability in the last 23 kyr. During the Last Glacial Maximum, the Greenland Sea was dominated by cold and sea-ice bearing surface water masses. Meltwater discharges from the surrounding ice sheets affected the area during the deglaciation, influencing the water mass circulation. During the Younger Dryas interval the last major freshwater event occurred in the region. The onset of the Holocene interglacial was marked by an increase in the advection of Atlantic Water and a rise in sea surface temperatures (SST). Although the thermal maximum was not reached simultaneously across the basin, benthic isotope data indicate that the rate of overturning circulation reached a maximum in the central Greenland Sea around 7 ka. After 6–5 ka a SST cooling and increasing sea-ice cover is noted. Conditions during this so-called "Neoglacial" cooling, however, changed after 3 ka, probably due to enhanced sea-ice expansion, which limited the deep convection. As a result, a well stratified upper water column amplified the warming of the subsurface waters in the central Greenland Sea, which were fed by increased inflow of Atlantic Water from the eastern Nordic Seas. Our data reveal that the Holocene oceanographic conditions in the Greenland Sea did not develop uniformly. These variations were a response to a complex interplay between the Atlantic and Polar water masses, the rate of sea-ice formation and melting and its effect on vertical convection intensity during times of Northern Hemisphere insolation changes.

Description: We investigate the respective role of variations in subpolar deep water formation and Nordic Seas overflows for the decadal to multidecadal variability of the Atlantic meridional overturning circulation (AMOC). This is partly done by analysing long (order of 1000 years) control simulations with five coupled climate models. For all models, the maximum influence of variations in subpolar deep water formation is found at about 45° N, while the maximum influence of variations in Nordic Seas overflows is rather found at 55 to 60° N. Regarding the two overflow branches, the influence of variations in the Denmark Strait overflow is, for all models, substantially larger than that of variations in the overflow across the Iceland–Scotland Ridge. The latter might, however, be underestimated, as the models in general do not realistically simulate the flow path of the Iceland–Scotland overflow water south of the Iceland–Scotland Ridge. The influence of variations in subpolar deep water formation is, on multimodel average, larger than that of variations in the Denmark Strait overflow. This is true both at 45° N, where the maximum standard deviation of decadal to multidecadal AMOC variability is located for all but one model, and at the more classical latitude of 30° N. At 30° N, variations in subpolar deep water formation and Denmark Strait overflow explain, on multimodel average, about half and one-third respectively of the decadal to multidecadal AMOC variance. Apart from analysing multimodel control simulations, we have performed sensitivity experiments with one of the models, in which we suppress the variability of either subpolar deep water formation or Nordic Seas overflows. The sensitivity experiments indicate that variations in subpolar deep water formation and Nordic Seas overflows are not completely independent. We further conclude from these experiments that the decadal to multidecadal AMOC variability north of about 50° N is mainly related to variations in Nordic Seas overflows. At 45° N and south of this latitude, variations in both subpolar deep water formation and Nordic Seas overflows contribute to the AMOC variability, with neither of the processes being very dominant compared to the other.

Description: Methyl iodide (CH3I}, bromoform (CHBr3) and dibromomethane (CH2Br2), which are produced naturally in the oceans, take part in ozone chemistry both in the troposphere and the stratosphere. The significance of oceanic upwelling regions for emissions of these trace gases in the global context is still uncertain although they have been identified as important source regions. To better quantify the role of upwelling areas in current and future climate, this paper analyzes major factors that influenced halocarbon emissions from the tropical North East Atlantic including the Mauritanian upwelling during the DRIVE expedition. Diel and regional variability of oceanic and atmospheric CH3I, CHBr3 and CH2Br2 was determined along with biological and meteorological parameters at six 24 h-stations. Low oceanic concentrations of CH3I from 0.1–5.4 pmol L-1 were equally distributed throughout the investigation area. CHBr3 of 1.0–42.4 pmol L-1 and CH2Br2 of 1.0–9.4 pmol L-1 were measured with maximum concentrations close to the Mauritanian coast. Atmospheric mixing rations of CH3I of up to 3.3, CHBr3 to 8.9 and CH2Br2 to 3.1 ppt above the upwelling and 1.8, 12.8, respectively 2.2 ppt at a Cape Verdean coast were detected during the campaign. While diel variability in CH3I emissions could be mainly ascribed to oceanic non-biological production, no main driver was identified for its emissions in the entire study region. In contrast, oceanic bromocarbons resulted from biogenic sources which were identified as regional drivers of their sea-to-air fluxes. The diel impact of wind speed on bromocarbon emissions increased with decreasing distance to the coast. The height of the marine atmospheric boundary layer (MABL) was determined as an additional factor influencing halocarbon emissions. Oceanic and atmospheric halocarbons correlated well in the study region and in combination with high oceanic CH3I, CHBr3 and CH2Br2 concentrations, local hot spots of atmospheric halocarbons could solely be explained by marine sources. This conclusion is in contrast with previous studies that hypothesized the occurrence of elevated atmospheric halocarbons over the eastern tropical Atlantic mainly originating from the West-African continent.

Description: Marine ecosystem models used to investigate how global change affects ocean ecosystems and their functioning typically omit pelagic plankton diversity. Diversity, however, may affect functions such as primary production and their sensitivity to environmental changes. Here we use a global ocean ecosystem model that explicitly resolves phytoplankton diversity by defining subtypes within four phytoplankton functional types (PFTs). We investigate the model's ability to capture diversity effects on primary production under environmental change. An idealized scenario with a sudden reduction in vertical mixing causes diversity and primary-production changes that turn out to be largely independent of the number of coexisting phytoplankton subtypes. The way diversity is represented in the model provides a small number of niches with respect to nutrient use in accordance with the PFTs defined in the model. Increasing the number of phytoplankton subtypes increases the resolution within the niches. Diversity effects such as niche complementarity operate between, but not within PFTs, and are constrained by the variety of traits and trade-offs resolved in the model. The number and nature of the niches formulated in the model, for example via trade-offs or different PFTs, thus determines the diversity effects on ecosystem functioning captured in ocean ecosystem models.

Description: Two consecutive summer upwelling events, each lasting for less than 24 h, were surveyed in high temporal and vertical resolution close to the Boknis Eck time-series station (BE) in the western Belt Sea (Baltic Sea) in summer 2010 with an autonomous glider. Driven only by moderate offshore winds both events resulted in more than 5 K cooling of surface waters, while only for the second event were significant irreversible changes in the vertical stratification observed. Generalizing the glider survey observations with hourly wind data from nearby meteorological stations, it is found that upwelling in the BE area occurs for wind directions between 190 to 260° and wind speed exceeding 4 m s−1. Based on these thresholds the wind-induced summer (June to September) upwelling conditions in the BE area for the period 1982 to 2012 are reconstructed. On average about 18 days of upwelling favourable wind conditions are found for the four summer months, with significant interannual variability ranging from 7.7 days (2006) to more than 28 days (1985). By aligning upwelling favourable wind conditions with the monthly BE surveys it is found that extreme anomalies in BE surveys follow extended periods of upwelling favourable winds.

Description: The Surface Ocean CO2 Atlas (SOCAT), an activity of the international marine carbon research community, provides access to synthesis and gridded fCO2 (fugacity of carbon dioxide) products for the surface oceans. Version 2 of SOCAT is an update of the previous release (version 1) with more data (increased from 6.3 million to 10.1 million surface water fCO2 values) and extended data coverage (from 1968–2007 to 1968–2011). The quality control criteria, while identical in both versions, have been applied more strictly in version 2 than in version 1. The SOCAT website (http://www.socat.info/) has links to quality control comments, metadata, individual data set files, and synthesis and gridded data products. Interactive online tools allow visitors to explore the richness of the data. Applications of SOCAT include process studies, quantification of the ocean carbon sink and its spatial, seasonal, year-to-year and longerterm variation, as well as initialisation or validation of ocean carbon models and coupled climate-carbon models.

Description: Production pathways of the prominent volatile organic halogen compound methyl iodide (CH3I) are not fully understood. Based on observations, production of CH3I via photochemical degradation of organic material or via phytoplankton production has been proposed. Additional insights could not be gained from correlations between observed biological and environmental variables or from biogeochemical modeling to identify unambiguously the source of methyl iodide. In this study, we aim to address this question of source mechanisms with a three-dimensional global ocean general circulation model including biogeochemistry (MPIOM-HAMOCC (MPIOM - Max Planck Institute Ocean Model HAMOCC - HAMburg Ocean Carbon Cycle model)) by carrying out a series of sensitivity experiments. The simulated fields are compared with a newly available global data set. Simulated distribution patterns and emissions of CH3I differ largely for the two different production pathways. The evaluation of our model results with observations shows that, on the global scale, observed surface concentrations of CH3I can be best explained by the photochemical production pathway. Our results further emphasize that correlations between CH3I and abiotic or biotic factors do not necessarily provide meaningful insights concerning the source of origin. Overall, we find a net global annual CH3I air-sea flux that ranges between 70 and 260 Gg yr(-1). On the global scale, the ocean acts as a net source of methyl iodide for the atmosphere, though in some regions in boreal winter, fluxes are of the opposite direction (from the atmosphere to the ocean).

Description: 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", 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.

Description: Late Miocene tectonic changes in Mediterranean–Atlantic connectivity and climatic changes caused Mediterranean salinity to fluctuate dramatically, including a ten-fold increase and near-freshening. Recent proxy- and model-based evidence suggests that at times during this Messinian Salinity Crisis (MSC, 5.96–5.33 Ma), highly saline and highly fresh Mediterranean water flowed into the North Atlantic Ocean, whilst at others, no Mediterranean Outflow Water (MOW) reached the Atlantic. By running extreme, sensitivity-type experiments with a fully coupled ocean–atmosphere general circulation model, we investigate the potential of these various MSC MOW scenarios to impact global-scale climate. The simulations suggest that although the effect remains relatively small, MOW had a greater influence on North Atlantic Ocean circulation and climate than it does today. We also find that depending on the presence, strength and salinity of MOW, the MSC could have been capable of cooling mid–high northern latitudes by a few degrees, with the greatest cooling taking place in the Labrador, Greenland–Iceland–Norwegian and Barents seas. With hypersaline MOW, a component of North Atlantic Deep Water formation shifts to the Mediterranean, strengthening the Atlantic Meridional Overturning Circulation (AMOC) south of 35° N by 1.5–6 Sv. With hyposaline MOW, AMOC completely shuts down, inducing a bipolar climate anomaly with strong cooling in the north (mainly −1 to −3 °C, but up to −8 °C) and weaker warming in the south (up to +0.5 to +2.7 °C). These simulations identify key target regions and climate variables for future proxy reconstructions to provide the best and most robust test cases for (a) assessing Messinian model performance, (b) evaluating Mediterranean–Atlantic connectivity during the MSC and (c) establishing whether or not the MSC could ever have affected global-scale climate.

Description: Most of the short-lived biogenic and anthropogenic chemical species that are emitted into the atmosphere break down efficiently by reaction with OH and do not reach the stratosphere. Here we show the existence of a pronounced minimum in the tropospheric column of ozone over the West Pacific, the main source region for stratospheric air, and suggest a corresponding minimum of the tropospheric column of OH. This has the potential to amplify the impact of surface emissions on the stratospheric composition compared to the impact when assuming globally uniform OH conditions. Specifically, the role of emissions of biogenic halogenated species for the stratospheric halogen budget and the role of increasing emissions of SO2 in Southeast Asia or from minor volcanic eruptions for the increasing stratospheric aerosol loading need to be reassessed in light of these findings. This is also important since climate change will further modify OH abundances and emissions of halogenated species. Our study is based on ozone sonde measurements carried out during the TransBrom cruise with the RV Sonne roughly along 140-150 degrees E in October 2009 and corroborating ozone and OH measurements from satellites, aircraft campaigns and FTIR instruments. Model calculations with the GEOS-Chem Chemistry and Transport Model (CTM) and the ATLAS CTM are used to simulate the tropospheric OH distribution over the West Pacific and the transport pathways to the stratosphere. The potential effect of the OH minimum on species transported into the stratosphere is shown via modeling the transport and chemistry of CH2Br2 and SO2.

Description: Despite its huge ecological importance, microbial oxygen respiration in pelagic waters is little studied, primarily due to methodological difficulties. Respiration measurements are challenging because of the required high resolution of oxygen concentration measurements. Recent improvements in oxygen sensing techniques bear great potential to overcome these limitations. Here we compare 3 different methods to measure oxygen consumption rates at low oxygen concentrations, utilizing amperometric Clark type sensors (STOX), optical sensors (optodes), and mass spectrometry in combination with 18-18O2 labeling. Oxygen concentrations and consumption rates agreed well between the different methods when applied in the same experimental setting. Oxygen consumption rates between 30 and 400 nmol L−1 h−1 were measured with high precision and relative standard errors of less than 3%. Rate detection limits in the range of 1 nmol L−1 h−1 were suitable for rate determinations in open ocean water and were lowest at the lowest applied O2 concentration.

Description: Natural compounds from marine fungi are an excellent source for the discovery and development of new drug leads. The distinct activity profiles of the two cyclodepsipeptides scopularide A and B against cancer cell lines set their marine producer strain Scopulariopsis brevicaulis LF580 into the focus of the EU project MARINE FUNGI. One of the main goals was the development of a sustainable biotechnological production process for these compounds. The secondary metabolite production of strain LF580 was optimized by random mutagenesis employing UV radiation. For a fast and reliable detection of the intracellular secondary metabolite production level, a miniaturized bioactivity-independent screening method was developed, as the random mutagenesis yielded a large number of mutants to be analysed quantitatively and none of the existing hyphenated bioassay-dependent screening systems could be applied. The method includes decreased cultivation volume, a fast extraction procedure as well as an optimized LC-MS analysis. We show that deviation could be specifically reduced at each step of the process: The measuring deviation during the analysis could be minimized to 5% and technical deviation occurring in the downstream part to 10–15%. Biological variation during the cultivation process still has the major influence on the overall variation. However, the approach led to a 10-fold reduction of time and similar effects on costs and effort compared to standard reference screening methods. The method was applied to screen the UV-mutants library of Scopulariopsis brevicaulis LF580. For validation purposes, the occurring variations in the miniaturized scale were compared to those in the classical Erlenmeyer flask scale. This proof of concept was performed using the wild type strain and 23 randomly selected mutant strains. One specific mutant strain with an enhanced production behavior could be obtained.

Description: It has been proposed that increasing levels of pCO(2) in the surface ocean will lead to more partitioning of the organic carbon fixed by marine primary production into the dissolved rather than the particulate fraction. This process may result in enhanced accumulation of dissolved organic carbon (DOC) in the surface ocean and/or concurrent accumulation of transparent exopolymer particles (TEPs), with important implications for the functioning of the marine carbon cycle. We investigated this in shipboard bioassay experiments that considered the effect of four different pCO(2) scenarios (ambient, 550, 750 and 1000 mu atm) on unamended natural phytoplankton communities from a range of locations in the northwest European shelf seas. The environmental settings, in terms of nutrient availability, phytoplankton community structure and growth conditions, varied considerably between locations. We did not observe any strong or consistent effect of pCO(2) on DOC production. There was a significant but highly variable effect of pCO(2) on the production of TEPs. In three of the five experiments, variation of TEP production between pCO(2) treatments was caused by the effect of pCO(2) on phytoplankton growth rather than a direct effect on TEP production. In one of the five experiments, there was evidence of enhanced TEP production at high pCO(2) (twice as much production over the 96 h incubation period in the 750 mu atm treatment compared with the ambient treatment) independent of indirect effects, as hypothesised by previous studies. Our results suggest that the environmental setting of experiments (community structure, nutrient availability and occurrence of phytoplankton growth) is a key factor determining the TEP response to pCO(2) perturbations.

Description: Four carbonate system variables were measured in surface waters during a cruise aimed at investigating ocean acidification impacts traversing northwestern European shelf seas in the summer of 2011. High-resolution surface water data were collected for partial pressure of carbon dioxide (pCO2; using two independent instruments) and pH using the total pH scale (pHT), in addition to discrete measurements of total alkalinity and dissolved inorganic carbon. We thus overdetermined the carbonate system (four measured variables, two degrees of freedom), which allowed us to evaluate the level of agreement between the variables on a cruise whose main aim was not intercomparison, and thus where conditions were more representative of normal working conditions. Calculations of carbonate system variables from other measurements generally compared well with direct observations of the same variables (Pearson's correlation coefficient always greater than or equal to 0.94; mean residuals were similar to the respective accuracies of the measurements). We therefore conclude that four of the independent data sets of carbonate chemistry variables were of high quality. A diurnal cycle with a maximum amplitude of 41 μatm was observed in the difference between the pCO2 values obtained by the two independent analytical pCO2 systems, and this was partly attributed to irregular seawater flows to the equilibrator and partly to biological activity inside the seawater supply and one of the equilibrators. We discuss how these issues can be addressed to improve carbonate chemistry data quality on future research cruises.

Description: Large scale patterns of ecologically relevant traits may help identify drivers of their variability and conditions beneficial or adverse to the expression of these traits. Antimicrofouling defenses in scleractinian corals regulate the establishment of the associated biofilm as well as the risks of infection. The Saudi Arabian Red Sea coast features a pronounced thermal and nutritional gradient including regions and seasons with potentially stressful conditions to corals. Assessing the patterns of antimicrofouling defenses across the Red Sea may hint at the susceptibility of corals to global change. We investigated microfouling pressure as well as the relative strength of 2 alternative antimicrofouling defenses (chemical antisettlement activity, mucus release) along the pronounced environmental gradient along the Saudi Arabian Red Sea coast in 2 successive years. Microfouling pressure was exceptionally low along most of the coast but sharply increased at the southernmost sites. Mucus release correlated with temperature. Chemical defense tended to anti-correlate with mucus release. As a result, the combined action of mucus release and chemical antimicrofouling defense seemed to warrant sufficient defense against microbes along the entire coast. In the future, however, we expect enhanced energetic strain on corals when warming and/or eutrophication lead to higher bacterial fouling pressure and a shift towards putatively more costly defense by mucus release

Description: Nitrate (NO3-) is the major nutrient responsible for coastal eutrophication worldwide and its production is related to intensive food production and fossil-fuel combustion. In the Baltic Sea NO3- inputs have increased 4-fold over recent decades and now remain constantly high. NO3- source identification is therefore an important consideration in environmental management strategies. In this study focusing on the Baltic Sea, we used a method to estimate the proportional contributions of NO3- from atmospheric deposition, N-2 fixation, and runoff from pristine soils as well as from agricultural land. Our approach combines data on the dual isotopes of NO3- (delta N-15-NO3- and delta O-18-NO3-) in winter surface waters with a Bayesian isotope mixing model (Stable Isotope Analysis in R, SIAR). Based on data gathered from 47 sampling locations over the entire Baltic Sea, the majority of the NO3- in the southern Baltic was shown to derive from runoff from agricultural land (33-100 %), whereas in the northern Baltic, i.e. the Gulf of Bothnia, NO3- originates from nitrification in pristine soils (34-100 %). Atmospheric deposition accounts for only a small percentage of NO3- levels in the Baltic Sea, except for contributions from northern rivers, where the levels of atmospheric NO3- are higher. An additional important source in the central Baltic Sea is N-2 fixation by diazotrophs, which contributes 49-65% of the overall NO3- pool at this site. The results obtained with this method are in good agreement with source estimates based upon delta N-15 values in sediments and a three-dimensional ecosystem model, ERGOM. We suggest that this approach can be easily modified to determine NO3- sources in other marginal seas or larger near-coastal areas where NO3- is abundant in winter surface waters when fractionation processes are minor.

Description: The goals of this study were (1) to investigate whether Fucus vesiculosus regulates the production of its antifouling defence chemicals against epibacteria in response to light limitation and temperature shifts and (2) to investigate if different surface concentrations of defence compounds shape epibacterial communities. F. vesiculosus was incubated in indoor mesocosms at five different temperature conditions (5 to 25°C) and in outdoor mesocosms under six differently reduced sunlight conditions (0 to 100%), respectively. Algal surface concentrations of previously identified antifouling compounds - dimethylsulphopropionate (DMSP), fucoxanthin and proline – were determined and the bacterial community composition was characterized by in-depth sequencing of the 16S-rRNA gene. Altogether, the effect of different treatment levels upon defence compound concentrations was limited. Under all conditions DMSP alone appeared to be sufficiently concentrated to warrant for at least a partial inhibitory action against epibiotic bacteria of F. vesiculosus. In contrast, proline and fucoxanthin rarely reached the necessary concentration ranges for self-contained inhibition. Nonetheless, in both experiments along with the direct influence of temperature and light, all three compounds apparently affected the overall bacterial community composition associated with F. vesiculosus since tendencies for insensitivity towards all three compounds were observed among bacterial taxa that typically dominate those communities. Given that the concentrations of at least one of the compounds (in most cases DMSP) were always high enough to inhibit bacterial settlement, we conclude that the capacity of F. vesiculosus for such defence will hardly be compromised by shading or warming to temperatures up to 25°C.

Description: Quantifying the scale and importance of individual dispersion between populations and life stages is a key challenge in marine ecology. The common sole (Solea solea), an important commercial flatfish in the North Sea, Atlantic Ocean and the Mediterranean Sea, has a marine pelagic larval stage, a benthic juvenile stage in coastal nurseries (lagoons, estuaries or shallow marine areas) and a benthic adult stage in deeper marine waters on the continental shelf. To date, the ecological connectivity among these life stages has been little assessed in the Mediterranean. Here, such an assessment is provided for the first time for the Gulf of Lions, NW Mediterranean, based on a dataset on otolith microchemistry and stable isotopic composition as indicators of the water masses inhabited by individual fish. Specifically, otolith Ba/Ca and Sr/Ca profiles, and delta C-13 and delta O-18 values of adults collected in four areas of the Gulf of Lions were compared with those of young-of-the-year collected in different coastal nurseries. Results showed that a high proportion of adults (〉46%) were influenced by river inputs during their larval stage. Furthermore Sr/Ca ratios and the otolith length at one year of age revealed that most adults (similar to 70%) spent their juvenile stage in nurseries with high salinity, whereas the remainder used brackish environments. In total, data were consistent with the use of six nursery types, three with high salinity (marine areas and two types of highly saline lagoons) and three brackish (coastal areas near river mouths, and two types of brackish environments), all of which contributed to the replenishment of adult populations. These finding implicated panmixia in sole population in the Gulf of Lions and claimed for a habitat integrated management of fisheries

Description: Marine calcareous sediments provide a fundamental basis for paleoceanographic studies aiming to reconstruct past oceanic conditions and understand key biogeochemical element cycles. Calcifying unicellular phytoplankton (coccolithophores) are a major contributor to both carbon and calcium cycling by photosynthesis and the production of calcite (coccoliths) in the euphotic zone and the subsequent long-term deposition and burial into marine sediments. Here we present data from controlled laboratory experiments on four coccolithophore species and elucidate the relation between the divalent cation (Sr, Mg and Ca) partitioning in coccoliths and cellular physiology (growth, calcification and photosynthesis). Coccolithophores were cultured under different seawater temperature and carbonate chemistry conditions. The partition coefficient of strontium (DSr) was positively correlated with both carbon dioxide (pCO2) and temperature but displayed no coherent relation to particulate organic and inorganic carbon production rates. Furthermore, DSr correlated positively with cellular growth rates when driven by temperature but no correlation was present when changes in growth rates were pCO2-induced. The results demonstrate the complex interaction between environmental forcing and physiological control on the strontium partitioning in coccolithophore calcite. The partition coefficient of magnesium (DMg) displayed species-specific differences and elevated values under nutrient limitation. No conclusive correlation between coccolith DMg and temperature was observed but pCO2 induced a rising trend in coccolith DMg. Interestingly, the best correlation was found between coccolith DMg and chlorophyll a production suggesting that chlorophyll a and calcite associated Mg originate from the same intracellular pool. These results give an extended insight into the driving factors that lead to variations in the coccolith Mg / Ca ratio and can be used for Sr / Ca and Mg / Ca paleoproxy calibration.

Description: In a pilot study conducted in October and November 2011, air–sea gas transfer velocities of the two sparingly soluble trace gases hexafluorobenzene and 1,4-difluorobenzene were measured in the unique High-Speed Wind-Wave Tank at Kyoto University, Japan. This air–sea interaction facility is capable of producing hurricane strength wind speeds of up to u10=67 m s−1. This constitutes the first lab study of gas transfer at such high wind speeds. The measured transfer velocities k600 spanned two orders of magnitude, lying between 11 cm h−1 and 1180 cm h−1 with the latter being the highest ever measured wind induced gas transfer velocity. The measured gas transfer velocities are in agreement with the only available dataset at hurricane wind speeds (McNeil and D'Asaro, 2007). The disproportionately large increase of the transfer velocities found at highest wind speeds indicates a new regime of air–sea gas transfer, which is characterized by strong wave breaking, enhanced turbulence and bubble cloud entrainment. It was found that tracers spanning a wide range of solubilities and diffusivities are needed to separate the effects of enhanced surface area and turbulence due to breaking waves from the effects of bubble and spray mediated gas transfer.

Description: Mass deposition fluxes of mineral dust to the tropical northeast Atlantic Ocean were determined within this study. In the framework of SOPRAN (Surface Ocean Processes in the Anthropocene), the interaction between the atmosphere and the ocean in terms of material exchange were investigated at the Cape Verde atmospheric observatory (CVAO) on the island Sao Vicente for January 2009. Five different methods were applied to estimate the deposition flux, using different meteorological and physical measurements, remote sensing, and regional dust transport simulations. The set of observations comprises micrometeorological measurements with an ultra-sonic anemometer and profile measurements using 2-D anemometers at two different heights, and microphysical measurements of the size-resolved mass concentrations of mineral dust. In addition, the total mass concentration of mineral dust was derived from absorption photometer observations and passive sampling. The regional dust model COSMO-MUSCAT was used for simulations of dust emission and transport, including dry and wet deposition processes. This model was used as it describes the AOD's and mass concentrations realistic compared to the measurements and because it was run for the time period of the measurements. The four observation-based methods yield a monthly average deposition flux of mineral dust of 12–29 ng m−2 s−1. The simulation results come close to the upper range of the measurements with an average value of 47 ng m−2 s−1. It is shown that the mass deposition flux of mineral dust obtained by the combination of micrometeorological (ultra-sonic anemometer) and microphysical measurements (particle mass size distribution of mineral dust) is difficult to compare to modeled mass deposition fluxes when the mineral dust is inhomogeneously distributed over the investigated area.

Description: Pro- and eukaryotic microbes associated with multi-cellular organisms are receiving increasing attention as a driving factor in ecosystems. Endophytes in plants can change host performance by altering nutrient uptake, secondary metabolite production or defense mechanisms. Recent studies detected widespread prevalence of Labyrinthula zosterae in European Zostera marina meadows, a protist that allegedly caused a massive amphi-Atlantic seagrass die-off event in the 1930's, while showing only limited virulence today. As a limiting factor for pathogenicity, we investigated genotype×genotype interactions of host and pathogen from different regions (10–100 km-scale) through reciprocal infection. Although the endophyte rapidly infected Z. marina, we found little evidence that Z. marina was negatively impacted by L. zosterae. Instead Z. marina showed enhanced leaf growth and kept endophyte abundance low. Moreover, we found almost no interaction of protist×eelgrass-origin on different parameters of L. zosterae virulence/Z. marina performance, and also no increase in mortality after experimental infection. In a target gene approach, we identified a significant down-regulation in the expression of 6/11 genes from the defense cascade of Z. marina after real-time quantitative PCR, revealing strong immune modulation of the host's defense by a potential parasite for the first time in a marine plant. Nevertheless, one gene involved in phenol synthesis was strongly up-regulated, indicating that Z. marina plants were probably able to control the level of infection. There was no change in expression in a general stress indicator gene (HSP70). Mean L. zosterae abundances decreased below 10% after 16 days of experimental runtime. We conclude that under non-stress conditions L. zosterae infection in the study region is not associated with substantial virulence.

Description: Diatoms can occur as single cells or as chain-forming aggregates. These two strategies affect buoyancy, predator evasion, light absorption and nutrient uptake. Adjacent cells in chains establish connections through various processes that determine strength and flexibility of the bonds, and at distinct cellular locations defining colony structure. Chain length has been found to vary with temperature and nutrient availability as well as being positively correlated with growth rate. However, the potential effect of enhanced carbon dioxide (CO2) concentrations and consequent changes in seawater carbonate chemistry on chain formation is virtually unknown. Here we report on experiments with semi-continuous cultures of the freshly isolated diatom Asterionellopsis glacialis grown under increasing CO2 levels ranging from 320 to 3400 mu atm. We show that the number of cells comprising a chain, and therefore chain length, increases with rising CO2 concentrations. We also demonstrate that while cell division rate changes with CO2 concentrations, carbon, nitrogen and phosphorus cellular quotas vary proportionally, evident by unchanged organic matter ratios. Finally, beyond the optimum CO2 concentration for growth, carbon allocation changes from cellular storage to increased exudation of dissolved organic carbon. The observed structural adjustment in colony size could enable growth at high CO2 levels, since longer, spiral-shaped chains are likely to create microclimates with higher pH during the light period. Moreover increased chain length of Asterionellopsis glacialis may influence buoyancy and, consequently, affect competitive fitness as well as sinking rates. This would potentially impact the delicate balance between the microbial loop and export of organic matter, with consequences for atmospheric carbon dioxide.

Description: Enhanced permafrost warming and increased arctic river discharges have heightened concern about the input of terrigeneous matter into Arctic coastal waters. We used optical operational satellite data from the Ocean Colour sensor MERIS onboard the ENVISAT satellite mission for synoptic monitoring of the pathways of terrigeneous matter in the southern Laptev Sea. MERIS satellite data from 2006 on to 2011 were processed using the Case2Regional Processor, C2R, installed in the open-source software ESA BEAM-VISAT. Since optical remote sensing using Ocean Colour satellite data has seen little application in Siberian Arctic coastal and shelf waters, we assess the applicability of the calculated MERIS parameters with surface water sampling data from the Russian-German ship expeditions LENA2010 and TRANSDRIFT-XVII taking place in August and September 2010 in the southern Laptev Sea. The surface waters of the southern Laptev Sea are characterized by low transparencies, due to turbid river water input, terrestrial input by coastal erosion, resuspension events and, therefore, high background concentrations of Suspended Particulate Matter, SPM, and coloured Dissolved Organic Matter, cDOM. The mapped calculated optical water parameters, such as the first attenuation depth, Z90, the attenuation coefficient, k, and Suspended Particulate Matter, SPM, visualize resuspension events that occur in shallow coastal and shelf waters indicating vertical mixing events. The mapped optical water parameters also visualize that the hydrography of the Laptev Sea is dominated by frontal meanders with amplitudes up to 30 km and eddies and filaments with diameters up to 100 km that prevail throughout the ice-free season. The meander crests, filaments and eddy-like structures that become visible through the mapped MERIS C2R parameters indicate enhanced vertical and horizontal transport energy for the transport of terrigenous and living biological matter in the surface waters during the ice-free season.

Description: Sea ice thickness information is important for sea ice modelling and ship operations. Here a method to detect the thickness of sea ice up to 50 cm during the freeze-up season based on high incidence angle observations of the Soil Moisture and Ocean Salinity (SMOS) satellite working at 1.4 GHz is suggested. By comparison of thermodynamic ice growth data with SMOS brightness temperatures, a high correlation to intensity and an anticorrelation to the difference between vertically and horizontally polarised brightness temperatures at incidence angles between 40 and 50° are found and used to develop an empirical retrieval algorithm sensitive to thin sea ice up to 50 cm thickness. The algorithm shows high correlation with ice thickness data from airborne measurements and reasonable ice thickness patterns for the Arctic freeze-up period.

Description: Accurate assessment of anthropogenic carbon dioxide (CO2) emissions and their redistribution among the atmosphere, ocean, and terrestrial biosphere is important to better understand the global carbon cycle, support the development of climate policies, and project future climate change. Here we describe data sets and a methodology to quantify all major components of the global carbon budget, including their uncertainties, based on the combination of a range of data, algorithms, statistics and model estimates and their interpretation by a broad scientific community. We discuss changes compared to previous estimates, consistency within and among components, alongside methodology and data limitations. CO2 emissions from fossil-fuel combustion and cement production (EFF) are based on energy statistics, while emissions from land-use change (ELUC), mainly deforestation, are based on combined evidence from land-cover change data, fire activity associated with deforestation, and models. The global atmospheric CO2 concentration is measured directly and its rate of growth (GATM) is computed from the annual changes in concentration. The mean ocean CO2 sink (SOCEAN) is based on observations from the 1990s, while the annual anomalies and trends are estimated with ocean models. The variability in SOCEAN is evaluated for the first time in this budget with data products based on surveys of ocean CO2 measurements. The global residual terrestrial CO2 sink (SLAND) is estimated by the difference of the other terms of the global carbon budget and compared to results of independent dynamic global vegetation models forced by observed climate, CO2 and land cover change (some including nitrogen–carbon interactions). All uncertainties are reported as ±1σ, reflecting the current capacity to characterise the annual estimates of each component of the global carbon budget. For the last decade available (2003–2012), EFF was 8.6 ± 0.4 GtC yr−1, ELUC 0.9 ± 0.5 GtC yr−1, GATM 4.3 ± 0.1 GtC yr−1, SOCEAN 2.5 ± 0.5 GtC yr−1, and SLAND 2.8 ± 0.8 GtC yr−1. For year 2012 alone, EFF grew to 9.7 ± 0.5 GtC yr−1, 2.2% above 2011, reflecting a continued growing trend in these emissions, GATM was 5.1 ± 0.2 GtC yr−1, SOCEAN was 2.9 ± 0.5 GtC yr−1, and assuming an ELUC of 1.0 ± 0.5 GtC yr−1 (based on the 2001–2010 average), SLAND was 2.7 ± 0.9 GtC yr−1. GATM was high in 2012 compared to the 2003–2012 average, almost entirely reflecting the high EFF. The global atmospheric CO2 concentration reached 392.52 ± 0.10 ppm averaged over 2012. We estimate that EFF will increase by 2.1% (1.1–3.1%) to 9.9 ± 0.5 GtC in 2013, 61% above emissions in 1990, based on projections of world gross domestic product and recent changes in the carbon intensity of the economy. With this projection, cumulative emissions of CO2 will reach about 535 ± 55 GtC for 1870–2013, about 70% from EFF (390 ± 20 GtC) and 30% from ELUC (145 ± 50 GtC). This paper also documents any changes in the methods and data sets used in this new carbon budget from previous budgets (Le Quéré et al., 2013). All observations presented here can be downloaded from the Carbon Dioxide Information Analysis Center (doi:10.3334/CDIAC/GCP_2013_V2.3).

Description: Marine bacteria are the main consumers of freshly produced organic matter. Many enzymatic processes involved in the bacterial digestion of organic compounds were shown to be pH sensitive in previous studies. Due to the continuous rise in atmospheric CO2 concentration, seawater pH is presently decreasing at a rate unprecedented during the last 300 million years but the consequences for microbial physiology, organic matter cycling and marine biogeochemistry are still unresolved. We studied the effects of elevated seawater pCO2 on a natural plankton community during a large-scale mesocosm study in a Norwegian fjord. Nine Kiel Off-Shore Mesocosms for Future Ocean Simulations (KOSMOS) were adjusted to different pCO2 levels ranging initially from ca. 280 to 3000 µatm and sampled every second day for 34 days. The first phytoplankton bloom developed around day 5. On day 14, inorganic nutrients were added to the enclosed, nutrient-poor waters to stimulate a second phytoplankton bloom, which occurred around day 20. Our results indicate that marine bacteria benefit directly and indirectly from decreasing seawater pH. During the first phytoplankton bloom, 5–10% more transparent exopolymer particles were formed in the high pCO2 mesocosms. Simultaneously, the efficiency of the protein-degrading enzyme leucine aminopeptidase increased with decreasing pH resulting in up to three times higher values in the highest pCO2/lowest pH mesocosm compared to the controls. In general, total and cell-specific aminopeptidase activities were elevated under low pH conditions. The combination of enhanced enzymatic hydrolysis of organic matter and increased availability of gel particles as substrate supported up to 28% higher bacterial abundance in the high pCO2 treatments. We conclude that ocean acidification has the potential to stimulate the bacterial community and facilitate the microbial recycling of freshly produced organic matter, thus strengthening the role of the microbial loop in the surface ocean.

Description: We provide a time series of Agulhas leakage anomalies over the last 20-years from satellite altimetry. Until now, measuring the interannual variability of Indo-Atlantic exchange has been the major barrier in the investigation of the dynamics and large scale impact of Agulhas leakage. We compute the difference of transport between the Agulhas Current and Agulhas Return Current, which allows us to deduce Agulhas leakage. The main difficulty is to separate the Agulhas Return Current from the southern limb of the subtropical "supergyre" south of Africa. For this purpose, an algorithm that uses absolute dynamic topography data is developed. The algorithm is applied to a state-of-the-art ocean model. The comparison with a Lagrangian method to measure the leakage allows us to validate the new method. An important result is that it is possible to measure Agulhas leakage in this model using the velocity field along a section that crosses both the Agulhas Current and the Agulhas Return Current. In the model a good correlation is found between measuring leakage using the full depth velocities and using only the surface geostrophic velocities. This allows us to extend the method to along-track absolute dynamic topography from satellites. It is shown that the accuracy of the mean dynamic topography does not allow us to determine the mean leakage but that leakage anomalies can be accurately computed.

Description: Due to the unprecedented rate at which our climate is changing, the ultimate consequence for many species is likely to be either extinction or migration to an alternate habitat. Certain species might, however, evolve at a rate that could make them resilient to the effects of a rapidly changing environment. This scenario is most likely to apply to species that have large population sizes and rapid generation times, such that the genetic variation required for adaptive evolution can be readily supplied. Emiliania huxleyi (Lohm.) Hay and Mohler (Prymnesiophyceae) is likely to be such a species, as it is the most conspicuous extant calcareous phytoplankton species in our oceans with growth rates of 1 day−1. Here we report on a validated set of microsatellites, in conjunction with the coccolithophore morphology motif genetic marker, to genotype 93 clonal isolates collected from across the world. Of these, 52 came from a single bloom event in the North Sea collected on the D366 United Kingdom Ocean Acidification cruise in June–July 2011. There were 26 multilocus genotypes (MLGs) encountered only once in the North Sea bloom and 8 MLGs encountered twice or up to six times. Each of these repeated MLGs exhibited Psex values of less than 0.05, indicating each repeated MLG was the product of asexual reproduction and not separate meiotic events. In addition, we show that the two most polymorphic microsatellite loci, EHMS37 and P01E05, are reporting on regions likely undergoing rapid genetic drift during asexual reproduction. Despite the small sample size, there were many more repeated genotypes than previously reported for other bloom-forming phytoplankton species, including a previously genotyped E. huxleyi bloom event. This study challenges the current assumption that sexual reproduction predominates during bloom events. Whilst genetic diversity is high amongst extant populations of E. huxleyi, the root cause for this diversity and ultimate fate of these populations still requires further examination. Nonetheless, we show that certain CMM genotypes are found everywhere, while others appear to have a regional bias.

Description: In this study we report diapycnal diffusive fluxes of dissolved iron (dFe), dissolved aluminium (dAl) and the major macronutrients to the surface waters of the North Atlantic subpolar gyre. Turbulent diffusivities at the base of the summer mixed layer ranged from 0.01 to 0.5 (median 0.07) cm2 s−1 and daily macronutrient fluxes into the surface mixed layer typically represented 〈 0.5% of integrated mixed layer inventories, although fluxes were highly variable. Elevated nutrient fluxes of up to 4% of mixed layer inventories were identified on the Greenland Shelf, where integrated nutrient pools were lowest due to localised shoaling of the mixed layer. Diffusive fluxes of dFe and dAl were typically 〈0.1% of mixed layer inventories but were also highly variable between stations. Approximations of daily phytoplankton nutrient and Fe uptake indicate that the diffusive flux may at best represent 〈10% of phytoplankton macronutrient uptake, and only 1% of daily phytoplankton Fe uptake. The daily turbulent diffusive flux of dFe was comparable in magnitude to coincident estimates of aeolian Fe supply but despite shallower than normal convective mixing in winter 2010 the diffusive supply was 22 and 59 times smaller than the annual convective supply of Fe to the Irminger and Iceland basins respectively. The general picture obtained from this study is one of small magnitude diffusive nutrient and Fe fluxes to the subpolar North Atlantic during the period of annual nutrient minima and indicates that the diffusive supply mechanism is unlikely to alleviate the recently identified presence of seasonal iron limitation within the North Atlantic subpolar gyre; a condition exacerbated by low dFe:NO3− ratios in subsurface source waters.

Description: Calcifying foraminifera are expected to be endangered by ocean acidification, However, the response of a complete community kept in natural sediment and over multiple generations under controlled laboratory conditions has not been constrained to date. During 5 six month incubation, foraminiferal assemblages were treated with pCO2 enriched seawater of 430, 907, 1865 and 3247 μatm pCO2. The fauna was dominated by Ammonia aomoriensis and Elphidium species, whereas agglutinated species were rare. After 6 months incubation, pore water alkalinity was much higher in comparison to the overlying seawater. Consequently, the saturation state of Òcalc was much higher in the sedi10 ment than in the water column in all pCO2 treatments and remained close to saturation. As a result, the life cycle of living assemblages was largely unaffected by the tested pCO2 treatments. Growth rates, reproduction and mortality, and therefore population densities and size-frequency distribution of Ammonia aomoriensis varied markedly during the experimental period. Growth rates varied between 25 and 50 μm per month, 15 which corresponds to an addition of 1 or 2 new chambers per month. According to the size-frequency distribution, foraminifera start reproduction at a diameter of 250 μm. Mortality of large foraminifera was recognized, commencing at a test size of 285 μm at a pCO2 ranging from 430 to 1865 μatm, and of 258 μm at 3247 μatm. The total organic content of living Ammonia aomoriensis has been determined to be 4.3% of dry 20 weight. Living individuals had a calcium carbonate production rate of 0.47 gm−2 yr−1, whereas dead empty tests accumulated at a rate of 0.27 gm−2a−1. Although Òcalc was close to 1, some empty tests of Ammonia aomoriensis showed dissolution features at the end of incubation. In contrast, tests of the subdominant species, Elphidium incertum, stayed intact. This species specific response could be explained by differences in 25 the elemental test composition, in particular the higher Mg-concentrations in Ammonia aomoriensis tests. Our results emphasize that the sensitivity to ocean acidification of endobenthic foraminifera in their natural sediment habitat is much lower compared to the experimental response of specimens isolated from the sediment.

Description: A general pattern in water mass distribution and potential shelf–basin exchange is revealed at the Laptev Sea continental slope based on hydrochemical and stable oxygen isotope data from the summers 2005–2009. Despite considerable interannual variations, a frontal system can be inferred between shelf, continental slope and central Eurasian Basin waters in the upper 100 m of the water column along the continental slope. Net sea-ice melt is consistently found at the continental slope. However, the sea-ice meltwater signal is independent from the local retreat of the ice cover and appears to be advected from upwind locations. In addition to the along-slope frontal system at the continental shelf break, a strong gradient is identified on the Laptev Sea shelf between 122° E and 126° E with an eastward increase of riverine and sea-ice related brine water contents. These waters cross the shelf break at ~ 140° E and feed the low-salinity halocline water (LSHW, salinity S 〈 33) in the upper 50 m of the water column. High silicate concentrations in Laptev Sea bottom waters may lead to speculation about a link to the local silicate maximum found within the salinity range of ~ 33 to 34.5, typical for the Lower Halocline Water (LHW) at the continental slope. However brine signatures and nutrient ratios from the central Laptev Sea differ from those observed at the continental slope. Thus a significant contribution of Laptev Sea bottom waters to the LHW at the continental slope can be excluded. The silicate maximum within the LHW at the continental slope may be formed locally or at the outer Laptev Sea shelf. Similar to the advection of the sea-ice melt signal along the Laptev Sea continental slope, the nutrient signal at 50–70 m water depth within the LHW might also be fed by advection parallel to the slope. Thus, our analyses suggest that advective processes from upstream locations play a significant role in the halocline formation in the northern Laptev Sea.

Description: Ash layers from explosive volcanic eruptions (i.e. tephra) represent isochronous surfaces independent from the environment in which they are deposited and the distance from their source. In comparison to eastern Beringia (non-glaciated Yukon and Alaska), few Plio-Pleistocene distal tephra are known from western Beringia (non-glaciated arctic and subarctic eastern Russia), hindering the dating and correlation of sediments beyond the limit of radiocarbon and luminescence methods. The identification of eight visible tephra layers (T0–T7) in sediment cores extracted from Lake El'gygytgyn, in the Far East Russian Arctic, indicates the feasibility of developing a tephrostratigraphic framework for this region. These tephra range in age from ca. 45 ka to 2.2 Ma, and each is described and characterized by its major-, minor-, trace-element and Pb isotope composition. These data show that subduction zone related volcanism from the Kurile–Kamchatka–Aleutian–Arc and Alaska Peninsula is the most likely source, with Pb isotope data indicating a Kamchatkan volcanic source for tephra layers T0–T5 and T7, while a source in the Aleutian Arc is possible probable for Tephra T6. The location of Lake El'gygytgyn relative to potential source volcanoes (〉1000 km) suggests these tephra are distributed over a vast area. These deposits provide a unique opportunity to correlate the high-resolution paleoenvironmental records of Lake El'gygytgyn to other terrestrial paleoenvironmental archives from western Beringia and marine records from the northwest Pacific and Bering Sea. This is an important first step towards the development of a robust integrated framework between the continuous paleoclimatic records of Lake El'gygytgyn and other terrestrial and marine records in NE Eurasia.

Description: Under modern conditions only North Pacific Intermediate Water is formed in the Northwest Pacific Ocean. This situation might have changed in the past. Recent studies with General Circulation Models indicate a switch to deep-water formation in the Northwest Pacific during Heinrich Stadial 1 (17.5–15.0 kyr) of the last glacial termination. Reconstructions of past ventilation changes based on paleoceanographic proxy records are still insufficient to test whether a deglacial mode of deep-water formation in the North Pacific Ocean existed. Here we present deglacial ventilation records based on radiocarbon-derived ventilation ages in combination with epibenthic stable carbon isotopes from the Northwest Pacific including the Okhotsk Sea and Bering Sea, the two potential source regions for past North Pacific ventilation changes. Evidence for most rigorous ventilation of the mid-depth North Pacific occurred during Heinrich Stadial 1 and the Younger Dryas, simultaneous to significant reductions in Atlantic Meridional Overturning Circulation. Concurrent changes in δ13C and ventilation ages point to the Okhotsk Sea as driver of millennial-scale changes in North Pacific Intermediate Water ventilation during the last deglaciation. Our records additionally indicate that changes in the δ13C intermediate water (700–1750 m water depth) signature and radiocarbon-derived ventilation ages are in antiphase to those of the deep North Pacific Ocean (〉2100 m water depth) during the last glacial termination. Thus, intermediate and deep-water masses of the Northwest Pacific have a differing ventilation history during the last deglaciation.

Description: The global ocean–climate system has been highly sensitive to the formation and advection of deep overflow water from the Nordic Seas as integral part of the Atlantic Meridional Overturning Circulation (AMOC) but its evolution over the Pliocene–Pleistocene global cooling is not fully understood. In particular, changes in the sources and mixing of prevailing deep waters that were involved in driving overturning throughout the Pliocene–Pleistocene climate transitions are not well constrained. Here we investigate the evolution of a substantial deep southward return overflow of the AMOC over the last 4 million years. We present new records of the bottom-water radiogenic neodymium isotope (ϵNd) variability obtained from three sediment cores (DSDP site 610 and ODP sites 980/981 and 900) at water depths between 2170 and 5050 m in the northeast Atlantic. We find that prior to the onset of major Northern Hemisphere glaciation (NHG) ∼3 million years ago (Ma), ϵNd values primarily oscillated between −9 and −11 at all sites, consistent with enhanced vertical mixing and weak stratification of the water masses during the warmer-than-today Pliocene period. From 2.7 Ma to ∼2.0 Ma, the ϵNd signatures of the water masses gradually became more distinct, which documents a significant advection of Nordic Seas overflow deep water coincident with the intensification of NHG. Most markedly, however, at ∼1.6 Ma the interglacial ϵNd signatures at sites 610 (2420 m water depth (w.d.)) and 980/981 (2170 m w.d.) synchronously and permanently shifted by 2 to 3 ϵNd units to less radiogenic values, respectively. Since then the difference between glacial and interglacial ϵNd values has been similar to the Late Quaternary at each site. A decrease of ∼2ϵNd units at 1.6 Ma was also recorded for the deepest water masses by site 900 (∼5050 m w.d.), which thereafter, however, evolved to more radiogenic values again until the present. This major ϵNd change across the 1.6 Ma transition reflects a significant reorganization of the overturning circulation in the northeast Atlantic paving the way for the more stratified water column with distinct water masses prevailing thereafter.

Description: X-ray absorption near edge structure (XANES) studies on calcium phosphate species (Ca-P) deal with marginal differences among subtle spectral features despite a hitherto missing systematic breakdown of these differences. Related fingerprinting approaches depend therefore on spectral libraries that are not validated against each other, incomplete and scattered among publications. This study compiled a comprehensive spectral library from published reference compound libraries in order to establish more clear-cut criteria for Ca-P determination by distinctive phosphorus K-edge XANES features. A specifically developed normalization method identified diagnostic spectral features within the compiled library, e.g. by uniform calculation of ratios between white-line and secondary peak heights. Post-processing of the spectra (n = 81) verified distinguishability among most but not all phases, which included hydroxylapatite (HAP), poorly crystalline HAP, amorphous HAP, fluorapatite, carbonate fluorapatite (CFAP), carbonate hydroxylapatite, β-tricalcium phosphate, octacalcium phosphate (OCP), brushite, monetite, monocalcium phosphate, amorphous calcium phosphate (ACP), anapaite, herderite, scholzite, messelite, whiteite and P on CaCO3. Particularly, peak height ratios significantly improved analyte specificity, e.g. by supplementary breakdown into OCP and ACP. The spectral analysis also revealed Ca-P standards that were rarely investigated or inappropriately synthesized, and thus provides a basis for standard selection and synthesis. The developed method and resulting breakdown by species were subsequently tested on Ca-P spectra from studies on bone and sediment. The test indicated that bone material likely comprises only poorly crystalline apatite, which implies direct nucleation of apatite in bone. This biological apatite formation is likely opposed to that of sedimentary apatite, which apparently forms by successive crystallization. Application of the method to μXANES spectra of sediment particles indicated authigenic apatite formation by an OCP precursor.

Description: Subduction of the oceanic Cocos plate offshore Costa Rica causes strong advection of methane-charged fluids. Presented here are the first direct measurements of microbial anaerobic oxidation of methane (AOM) and sulfate reduction (SR) rates in sediments from the two mounds, applying radiotracer techniques in combination with numerical modeling. In addition, analysis of carbonate δ18O, δ13C, and 87Sr / 86Sr signatures constrain the origin of the carbonate-precipitating fluid. Average rates of microbial activities showed differences with a factor of 4.8 to 6.3 between Mound 11 [AOM 140.71 (±40.84 SD); SR 117.25 (±82.06 SD) mmol m−2 d−1, respectively] and Mound 12 [AOM 22.37 (±0.85 SD); SR 23.99 (±5.79 SD) mmol m−2 d−1, respectively]. Modeling results yielded flow velocities of 50 cm a−1 at Mound 11 and 8–15 cm a−1 at Mound 12. Analysis of oxygen and carbon isotope variations of authigenic carbonates from the two locations revealed higher values for Mound 11 (δ18O: 4.7 to 5.9‰, δ13C: −21.0 to −29.6‰), compared to Mound 12 (δ18O: 4.1 to 4.5‰, δ13C: −45.7 to −48.9‰). Analysis of carbonates 87Sr / 86Sr indicated temporal changes of deep-source fluid admixture at Mound 12. The present study is in accordance with previous work supporting considerable differences of methane flux between the two Mounds. It also strengthens the hypothesis of a predominantly deep fluid source for Mound 11 versus a rather shallow source of biogenic methane for Mound 12. The results demonstrate that methane-driven microbial activity is a valid ground truthing tool for geophysical measurements of fluid advection and constraining of recent methane fluxes in the study area. The study further shows that the combination of microbial rate measurements, numerical modeling, and authigenic carbonate analysis provide a suitable approach to constrain temporal and spatial variations of methane charged fluid flow at the Pacific Costa Rican margin.

Description: Bacteria of the genus Vibrio occur at a continuum from free-living to symbiotic life forms, including opportunists and pathogens, that can contribute to severe diseases, for instance summer mortality events of Pacific oysters Crassostrea gigas. While most studies focused on Vibrio isolated from moribund oysters during mortality outbreaks, investigations of the Vibrio community in healthy oysters are rare. Therefore, we characterized the persistence, diversity, seasonal dynamics, and pathogenicity of the Vibrio community isolated from healthy Pacific oysters. In a reciprocal transplant experiment we repeatedly sampled hemolymph from adult Pacific oysters to differentiate population from site-specific effects during six months of in situ incubation in the field. We characterized virulence phenotypes and genomic diversity based on multilocus sequence typing in a total of 70 Vibrio strains. Based on controlled infection experiments we could show that strains with the ability to colonize healthy adult oysters can also have the potential to induce high mortality rates on larvae. Diversity and abundance of Vibrio varied significantly over time with highest values during and after spawning season. Vibrio communities from transplanted and stationary oysters converged over time, indicating that communities were not population specific, but rather assemble from the surrounding environment forming communities, some of which can persist over longer periods

Description: With an extension of 〉40 km2 the recently discovered Campeche cold-water coral province located at the northeastern rim of the Campeche Bank in the southern Gulf of Mexico belongs to the largest coherent cold-water coral areas discovered so far. The Campeche province consists of numerous 20 to 40 m high coral ridges that are developed in intermediate water depths of 500 to 600 m. The ridges are colonized by a vivid cold-water coral ecosystem that covers the upper flanks and summits. The rich coral community is dominated by the framework-building scleractinia Enallopsammia profunda and Lophelia pertusa while the associated benthic megafauna shows a rather scarce occurrence. The recent environmental setting is characterized by a high surface water production caused by a local upwelling center and a dynamic bottom water regime comprising vigorous bottom currents, internal waves and strong density contrasts, which all together provide optimal conditions for the growth of cold-water corals. The strong hydrodynamics – potentially supported by the diel vertical migration of zooplankton in the Campeche area – drive the delivering of food particles to the corals. The Campeche cold-water coral province is, thus, an excellent example highlighting the importance of the hydrographic setting in securing the food supply for the development of large and vivid cold-water coral ecosystems.

Description: Ventilation is the primary pathway for atmosphere–ocean boundary perturbations, such as temperature anomalies, to be relayed to the ocean interior. It is also a conduit for gas exchange between the interface of atmosphere and ocean. Thus it is a mechanism whereby, for instance, the ocean interior is oxygenated and enriched in anthropogenic carbon. The ventilation of the Mediterranean Sea is fast in comparison to the world ocean and has large temporal variability. Here we present transient tracer data from a field campaign in April 2011 that sampled a unique suite of transient tracers (SF6, CFC-12, 3H and 3He) in all major basins of the Mediterranean. We apply the transit time distribution (TTD) model to the data in order to constrain the mean age, the ratio of the advective / diffusive transport and the number of water masses significant for ventilation. We found that the eastern part of the eastern Mediterranean can be reasonably described with a one-dimensional inverse Gaussian TTD (IG-TTD), and thus constrained with two independent tracers. The ventilation of the Ionian Sea and the western Mediterranean can only be constrained by a linear combination of IG-TTDs. We approximate the ventilation with a one-dimensional, two inverse Gaussian TTD (2IG-TTD) for these areas and demonstrate a possibility of constraining a 2IG-TTD from the available transient tracer data. The deep water in the Ionian Sea has a mean age between 120 and 160 years and is therefore substantially older than the mean age of the Levantine Basin deep water (60–80 years). These results are in contrast to those expected by the higher transient tracer concentrations in the Ionian Sea deep water. This is partly due to deep water of Adriatic origin having more diffusive properties in transport and formation (i.e., a high ratio of diffusion over advection), compared to the deep water of Aegean Sea origin that still dominates the deep Levantine Basin deep water after the Eastern Mediterranean Transient (EMT) in the early 1990s. The tracer minimum zone (TMZ) in the intermediate of the Levantine Basin is the oldest water mass with a mean age up to 290 years. We also show that the deep western Mediterranean has contributed approximately 40% of recently ventilated deep water from the Western Mediterranean Transition (WMT) event of the mid-2000s. The deep water has higher transient tracer concentrations than the mid-depth water, but the mean age is similar with values between 180 and 220 years.

Description: Coastal hypoxia and anoxia have become a global key stressor to marine ecosystems, with almost 500 dead zones recorded worldwide. By triggering cascading effects from the individual organism to the community-and ecosystem level, oxygen depletions threaten marine biodiversity and can alter ecosystem structure and function. By integrating both physiological function and ecological processes, animal behaviour is ideal for assessing the stress state of benthic macrofauna to low dissolved oxygen. The initial response of organisms can serve as an early warning signal, while the successive behavioural reactions of key species indicate hypoxia levels and help assess community degradation. Here we document the behavioural responses of a representative spectrum of benthic macrofauna in the natural setting in the Northern Adriatic Sea (Mediterranean). We experimentally induced small-scale anoxia with a benthic chamber in 24m depth to overcome the difficulties in predicting the onset of hypoxia, which often hinders full documentation in the field. The behavioural reactions were documented with a time-lapse camera. Oxygen depletion elicited significant and repeatable changes in general (visibility, locomotion, body movement and posture, location) and species-specific reactions in virtually all organisms (302 individuals from 32 species and 2 species groups). Most atypical (stress) behaviours were associated with specific oxygen thresholds: arm-tipping in the ophiuroid Ophiothrix quinquemaculata, for example, with the onset of mild hypoxia (〈 2mLO(2) L-1), the emergence of polychaetes on the sediment surface with moderate hypoxia (〈 1mLO(2) L-1), the emergence of the infaunal sea urchin Schizaster canaliferus on the sediment with severe hypoxia (〈 0.5mLO(2) L-1) and heavy body rotations in sea anemones with anoxia. Other species changed their activity patterns, for example the circadian rhythm in the hermit crab Paguristes eremita or the bioherm-associated crab Pisidia longimana. Intra-and interspecific reactions were weakened or changed: decapods ceased defensive and territorial behaviour, and predator-prey interactions and relationships shifted. This nuanced scale of resolution is a useful tool to interpret present benthic community status (behaviour) and past mortalities (community composition, e.g. survival of tolerant species). This information on the sensitivity (onset of stress response), tolerance (mortality, survival), and characteristics (i. e. life habit, functional role) of key species also helps predict potential future changes in benthic structure and ecosystem functioning. This integrated approach can transport complex ecological processes to the public and decision-makers and help define specific monitoring, assessment and conservation plans

Description: The Mediterranean Sea (MedSea) is considered a "laboratory basin" being an ocean in miniature, suffering dramatic changes in its oceanographic and biogeochemical conditions derived from natural and anthropogenic forces. Moreover, the MedSea is prone to absorb and store anthropogenic carbon due to the particular CO2 chemistry and the active overturning circulation. Despite this, water column CO2 measurements covering the whole basin are scarce. This work aims to be a base-line for future studies about the CO2 system space-time variability in the MedSea combining historic and modern CO2 cruises in the whole area. Here we provide an extensive vertical and longitudinal description of the CO2 system variables (total alkalinity – TA, dissolved inorganic carbon – DIC and pH) along an East-West transect and across the Sardinia-Sicily passage in the MedSea from two oceanographic cruises conducted in 2011 measuring CO2 variables in a coordinated fashion, the RV Meteor M84/3 and the RV Urania EuroFleets 11, respectively. In this sense, we provide full-depth and length CO2 distributions across the MedSea, and property-property plots showing in each sub-basin post-Eastern Mediterranean Transient (EMT) situation with regard to TA, DIC and pH. The over-determined CO2 system in 2011 allowed performing the first internal consistency analysis for the particularly warm, high salinity and alkalinity MedSea waters. The CO2 constants by Mehrbach et al. (1973) refitted by Dickson and Millero (1987) are recommended. The sensitivity of the CO2 system to the atmospheric CO2 increase, DIC and/or TA changes is evaluated by means of the Revelle and buffer factors.

Description: The Baltic Sea is a seasonally ice-covered marginal sea located in a densely populated area in northern Europe. Severe sea ice conditions have the potential to hinder the intense ship traffic considerably. Thus, sea ice fore- and nowcasts are regularly provided by the national weather services. Typically, the forecast comprises several ice properties that are distributed as prognostic variables, but their actual usefulness is difficult to measure, and the ship captains must determine their relative importance and relevance for optimal ship speed and safety ad hoc. The present study provides a more objective approach by comparing the ship speeds, obtained by the automatic identification system (AIS), with the respective forecasted ice conditions. We find that, despite an unavoidable random component, this information is useful to constrain and rate fore- and nowcasts. More precisely, 62–67% of ship speed variations can be explained by the forecasted ice properties when fitting a mixed-effect model. This statistical fit is based on a test region in the Bothnian Sea during the severe winter 2011 and employs 15 to 25 min averages of ship speed.

Description: Coccolithophores are a key functional group in terms of the pelagic production of calcium carbonate (calcite), although their contribution to shelf sea biogeochemistry, and how this relates to environmental conditions, is poorly constrained. Measurements of calcite production (CP) and coccolithophore abundance were made on the north-west European shelf to examine trends in coccolithophore calcification along natural gradients of carbonate chemistry, macronutrient availability and plankton composition. Similar measurements were also made in three bioassay experiments where nutrient (nitrate, phosphate) and pCO2 levels were manipulated. Nanoflagellates (〈 10 μm) dominated chlorophyll biomass and primary production (PP) at all but one sampling site, with CP ranging from 0.6 to 9.6 mmol C m−2 d−1. High CP and coccolithophore abundance occurred in a diatom bloom in fully mixed waters off Heligoland, but not in two distinct coccolithophore blooms in the central North Sea and Western English Channel. Coccolithophore abundance and CP showed no correlation with nutrient concentrations or ratios, while significant (p 〈 0.01) correlations between CP, cell-specific calcification (cell-CF) and irradiance in the water column highlighted how light availability exerts a strong control on pelagic CP. In the experimental bioassays, Emiliania-huxleyi-dominated coccolithophore communities in shelf waters (northern North Sea, Norwegian Trench) showed a strong response in terms of CP to combined nitrate and phosphate addition, mediated by changes in cell-CF and growth rates. In contrast, an offshore diverse coccolithophore community (Bay of Biscay) showed no response to nutrient addition, while light availability or mortality may have been more important in controlling this community. Sharp decreases in pH and a rough halving of calcite saturation states in the bioassay experiments led to decreased CP in the Bay of Biscay and northern North Sea, but not the Norwegian Trench. These decreases in CP were related to slowed growth rates in the bioassays at elevated pCO2 (750 μatm) relative to those in the ambient treatments. The combined results from our study highlight the variable coccolithophore responses to irradiance, nutrients and carbonate chemistry in north-west European shelf waters, which are mediated by changes in growth rates, cell-CF and species composition.

Description: The ongoing oceanic uptake of anthropogenic carbon dioxide (CO2) is significantly altering the carbonate chemistry of seawater, a phenomenon referred to as ocean acidification. Experimental manipulations have been increasingly used to gauge how continued ocean acidification will potentially impact marine ecosystems and their associated biogeochemical cycles in the future; however, results amongst studies, particularly when performed on natural communities, are highly variable, which may reflect community/environment-specific responses or inconsistencies in experimental approach. To investigate the potential for identification of more generic responses and greater experimentally reproducibility, we devised and implemented a series (n = 8) of short-term (2–4 days) multi-level (≥4 conditions) carbonate chemistry/nutrient manipulation experiments on a range of natural microbial communities sampled in Northwest European shelf seas. Carbonate chemistry manipulations and resulting biological responses were found to be highly reproducible within individual experiments and to a lesser extent between geographically separated experiments. Statistically robust reproducible physiological responses of phytoplankton to increasing pCO2, characterised by a suppression of net growth for small-sized cells (〈10 μm), were observed in the majority of the experiments, irrespective of natural or manipulated nutrient status. Remaining between-experiment variability was potentially linked to initial community structure and/or other site-specific environmental factors. Analysis of carbon cycling within the experiments revealed the expected increased sensitivity of carbonate chemistry to biological processes at higher pCO2 and hence lower buffer capacity. The results thus emphasise how biogeochemical feedbacks may be altered in the future ocean.

Description: Algal symbionts (zooxanthellae, genus Symbiodinium) of scleractinian corals respond strongly to temperature, nutrient and light changes. These factors vary greatly along the north-south gradient in the Red Sea and include conditions, which are outside of those typically considered optimal for coral growth. Nevertheless, coral communities thrive throughout the Red Sea, suggesting that zooxanthellae have successfully acclimatized or adapted to the harsh conditions they experience particularly in the south (high temperatures and high nutrient supply). As such, the Red Sea is a region, which may help to better understand how zooxanthellae and their coral hosts successfully acclimatize or adapt to environmental change (e.g. increased temperatures and localized eutrophication). To gain further insight into the physiology of coral symbionts in the Red Sea, we examined the abundance of dominant Symbiodinium types associated with the coral Pocillopora verrucosa, and measured Symbiodinium physiological characteristics (i.e. photosynthetic processes, cell density, pigmentation, and protein composition) along the latitudinal gradient of the Red Sea in summer and winter. Despite the strong environmental gradients from north to south, our results demonstrate that Symbiodinium microadriaticum (type A1) was the predominant species in P. verrucosa along the latitudinal gradient. Furthermore, measured physiological characteristics were found to vary more with prevailing seasonal environmental conditions than with region-specific differences, although the measured environmental parameters displayed much higher spatial than temporal variability. We conclude that our findings might present the result of long-term acclimatization or adaptation of S. microadriaticum to regionally specific conditions within the Red Sea. Of additional note, high nutrients in the South correlated with high zooxanthellae density indicating a compensation for a temperature-driven loss of photosynthetic performance, which may prove promising for the resilience of these corals under increase of temperature increase and eutrophication.

Description: The response of the phytoplanktonic community (primary production and algal biomass) to contrasted Saharan dust events (wet and dry deposition) was studied in the framework of the DUNE ("a DUst experiment in a low-Nutrient, low-chlorophyll Ecosystem") project. We simulated realistic dust deposition events (10 gm(-2)) into large mesocosms (52m(3)). Three distinct dust addition experiments were conducted in June 2008 (DUNE-1-P: simulation of a wet deposition; DUNE-1-Q: simulation of a dry deposition) and 2010 (DUNE-2-R1 and DUNE-2-R2: simulation of two successive wet depositions) in the northwestern oligotrophic Mediterranean Sea. No changes in primary production (PP) and chlorophyll a concentrations (Chl a) were observed after a dry deposition event, while a wet deposition event resulted in a rapid (24 h after dust addition), strong (up to 2.4-fold) and long (at least a week in duration) increase in PP and Chl a. We show that, in addition to being a source of dissolved inorganic phosphorus (DIP), simulated wet deposition events were also a significant source of nitrate (NO3-) (net increases up to +9.8 mu M NO3- at 0.1m in depth) to the nutrient-depleted surface waters, due to cloud processes and mixing with anthropogenic species such as HNO3. The dry deposition event was shown to be a negligible source of NO3-. By transiently increasing DIP and NO3- concentrations in N-P starved surface waters, wet deposition of Saharan dust was able to relieve the potential N or NP co-limitation of the phytoplanktonic activity. Due to the higher input of NO3- relative to DIP, and taking into account the stimulation of the biological activity, a wet deposition event resulted in a strong increase in the NO3-/DIP ratio, from initially less than 6, to over 150 at the end of the DUNE-2-R1 experiment, suggesting a switch from an initial N or NP co-limitation towards a severe P limitation. We also show that the contribution of new production to PP strongly increased after wet dust deposition events, from initially 15% to 60-70% 24 h after seeding, indicating a switch from a regenerated-production based system to a new-production based system. DUNE experiments show that wet and dry dust deposition events induce contrasting responses of the phytoplanktonic community due to differences in the atmospheric supply of bioavailable new nutrients. Our results from original mesocosm experiments demonstrate that atmospheric dust wet deposition greatly influences primary productivity and algal biomass in LNLC environments through changes in the nutrient stocks, and alters the NO3-/DIP ratio, leading to a switch in the nutrient limitation of the phytoplanktonic activity.

Description: The marine CaCO3 cycle is an important component of the oceanic carbon system and directly affects the cycling of natural and the uptake of anthropogenic carbon. In numerical models of the marine carbon cycle, the CaCO3 cycle component is often evaluated against the observed distribution of alkalinity. Alkalinity varies in response to the formation and remineralization of CaCO3 and organic matter. However, it also has a large conservative component, which may strongly be affected by a deficient representation of ocean physics (circulation, evaporation, and precipitation) in models. Here we apply a global ocean biogeochemical model run into preindustrial steady state featuring a number of idealized tracers, explicitly capturing the model's CaCO3 dissolution, organic matter remineralization, and various preformed properties (alkalinity, oxygen, phosphate). We compare the suitability of a variety of measures related to the CaCO3 cycle, including alkalinity (TA), potential alkalinity and TA*, the latter being a measure of the time-integrated imprint of CaCO3 dissolution in the ocean. TA* can be diagnosed from any data set of TA, temperature, salinity, oxygen and phosphate. We demonstrate the sensitivity of total and potential alkalinity to the differences in model and ocean physics, which disqualifies them as accurate measures of biogeochemical processes. We show that an explicit treatment of preformed alkalinity (TA0) is necessary and possible. In our model simulations we implement explicit model tracers of TA0 and TA*. We find that the difference between modelled true TA* and diagnosed TA* was below 10% (25%) in 73% (81%) of the ocean's volume. In the Pacific (and Indian) Oceans the RMSE of A* is below 3 (4) mmol TA m−3, even when using a global rather than regional algorithms to estimate preformed alkalinity. Errors in the Atlantic Ocean are significantly larger and potential improvements of TA0 estimation are discussed. Applying the TA* approach to the output of three state-of-the-art ocean carbon cycle models, we demonstrate the advantage of explicitly taking preformed alkalinity into account for separating the effects of biogeochemical processes and circulation on the distribution of alkalinity. In particular, we suggest to use the TA* approach for CaCO3 cycle model evaluation.

Description: Climate forcing in complex ecosystems can have profound implications for ecosystem sustainability and may thus challenge a precautionary ecosystem management. Climatic influences documented to affect various ecological functions on a global scale, may themselves be observed on quantitative or qualitative scales including regime shifts in complex marine ecosystems. This study investigates the potential climatic impact on the reproduction success of spring-spawning herring (Clupea harengus) in the Western Baltic Sea (WBSS herring). To test for climate effects on reproduction success, the regionally determined and scientifically well-documented spawning grounds of WBSS herring represent an ideal model system. Climate effects on herring reproduction were investigated using two global indices of atmospheric variability and sea surface temperature, represented by the North Atlantic Oscillation (NAO) and the Atlantic Multi-decadal Oscillation (AMO), respectively, and the Baltic Sea Index (BSI) which is a regional-scale atmospheric index for the Baltic Sea. Moreover, we combined a traditional approach with modern time series analysis based on a recruitment model connecting parental population components with reproduction success. Generalized transfer functions (ARIMAX models) allowed evaluating the dynamic nature of exogenous climate processes interacting with the endogenous recruitment process. Using different model selection criteria our results reveal that in contrast to NAO and AMO, the BSI shows a significant positive but delayed signal on the annual dynamics of herring recruitment. The westward influence of the Siberian high is considered strongly suppressing the influence of the NAO in this area leading to a higher explanatory power of the BSI reflecting the atmospheric pressure regime on a North-South transect between Oslo, Norway and Szczecin, Poland. We suggest incorporating climate-induced effects into stock and risk assessments and management strategies as part of the EU ecosystem approach to support sustainable herring fisheries in the Western Baltic Sea.

Description: Increasing atmospheric CO2 concentrations are expected to impact pelagic ecosystem functioning in the near future by driving ocean warming and acidification. While numerous studies have investigated impacts of rising temperature and seawater acidification on planktonic organisms separately, little is presently known on their combined effects. To test for possible synergistic effects we exposed two coccolithophore species, Emiliania huxleyi and Gephyrocapsa oceanica, to a CO2 gradient ranging from ,0.5–250 mmol kg21 (i.e. ,20–6000 matm pCO2) at three different temperatures (i.e. 10, 15, 20uC for E. huxleyi and 15, 20, 25uC for G. oceanica). Both species showed CO2-dependent optimum-curve responses for growth, photosynthesis and calcification rates at all temperatures. Increased temperature generally enhanced growth and production rates and modified sensitivities of metabolic processes to increasing CO2. CO2 optimum concentrations for growth, calcification, and organic carbon fixation rates were only marginally influenced from low to intermediate temperatures. However, there was a clear optimum shift towards higher CO2 concentrations from intermediate to high temperatures in both species. Our results demonstrate that the CO2 concentration where optimum growth, calcification and carbon fixation rates occur is modulated by temperature. Thus, the response of a coccolithophore strain to ocean acidification at a given temperature can be negative, neutral or positive depending on that strain’s temperature optimum. This emphasizes that the cellular responses of coccolithophores to ocean acidification can only be judged accurately when interpreted in the proper eco-physiological context of a given strain or species. Addressing the synergistic effects of changing carbonate chemistry and temperature is an essential step when assessing the success of coccolithophores in the future ocean.

Description: The Baltic Sea is a seasonally ice-covered, marginal sea in central northern Europe. It is an essential waterway connecting highly industrialised countries. Because ship traffic is intermittently hindered by sea ice, the local weather services have been monitoring sea ice conditions for decades. In the present study we revisit a historical monitoring data set, covering the winters 1960/1961 to 1978/1979. This data set, dubbed Data Bank for Baltic Sea Ice and Sea Surface Temperatures (BASIS) ice, is based on hand-drawn maps that were collected and then digitised in 1981 in a joint project of the Finnish Institute of Marine Research (today the Finnish Meteorological Institute (FMI)) and the Swedish Meteorological and Hydrological Institute (SMHI). BASIS ice was designed for storage on punch cards and all ice information is encoded by five digits. This makes the data hard to access. Here we present a post-processed product based on the original five-digit code. Specifically, we convert to standard ice quantities (including information on ice types), which we distribute in the current and free Network Common Data Format (NetCDF). Our post-processed data set will help to assess numerical ice models and provide easy-to-access unique historical reference material for sea ice in the Baltic Sea. In addition we provide statistics showcasing the data quality. The website http://www.baltic-ocean.org hosts the post-processed data and the conversion code. The data are also archived at the Data Publisher for Earth & Environmental Science, PANGAEA (doi:10.1594/PANGAEA.832353)

Description: Observations and simple theoretical arguments suggest that the Northern Hemisphere (NH) stratospheric polar vortex is stronger in winters following major volcanic eruptions. However, recent studies show that climate models forced by prescribed volcanic aerosol fields fail to reproduce this effect. We investigate the impact of volcanic aerosol forcing on stratospheric dynamics, including the strength of the NH polar vortex, in ensemble simulations with the Max Planck Institute Earth System Model. The model is forced by four different prescribed forcing sets representing the radiative properties of stratospheric aerosol following the 1991 eruption of Mt. Pinatubo: two forcing sets are based on observations, and are commonly used in climate model simulations, and two forcing sets are constructed based on coupled aerosol–climate model simulations. For all forcings, we find that simulated temperature and zonal wind anomalies in the NH high latitudes are not directly impacted by anomalous volcanic aerosol heating. Instead, high-latitude effects result from enhancements in stratospheric residual circulation, which in turn result, at least in part, from enhanced stratospheric wave activity. High-latitude effects are therefore much less robust than would be expected if they were the direct result of aerosol heating. Both observation-based forcing sets result in insignificant changes in vortex strength. For the model-based forcing sets, the vortex response is found to be sensitive to the structure of the forcing, with one forcing set leading to significant strengthening of the polar vortex in rough agreement with observation-based expectations. Differences in the dynamical response to the forcing sets imply that reproducing the polar vortex responses to past eruptions, or predicting the response to future eruptions, depends on accurate representation of the space–time structure of the volcanic aerosol forcing.

Description: In this study we explore the correlation of I/Ca ratios in three calcitic and one aragonitic foraminiferal species. I/Ca ratios are evaluated as possible proxies for changes in ambient redox conditions across the Peruvian oxygen minimum zone to the ambient oxygen concentrations in the habitat of the foraminiferal species studied. We test cleaning and measurement methods to determine I/Ca ratios in benthic foraminifera from the Peruvian oxygen minimum zone. All species show a positive trend in their I/Ca ratios as a function of higher oxygen concentrations and these trends are all statistically significant except for the aragonitic species Hoeglundina elegans. The most promising species appears to be Uvigerina striata which shows a highly statistically significant correlation between I/Ca ratios and bottom water (BW) oxygenation (I/Ca = 0.032(± 0.004)[O2]BW + 0.29(± 0.03), R2 = 0.61, F = 75, P 〈 0.0001). Although I/Ca ratios in benthic foraminifera might prove to be a valuable proxy for changing redox-conditions the iodine volatility in acidic solutions, the species dependency ofI/Ca–[O2]BW correlations, and the individual variability of single tests severely interfere with the observed I/Ca–[O2]BW relationship.

Description: Large tectonic plates are known to be susceptible to internal deformation, leading to a range of phenomena including intraplate volcanism. However, the space and time dependence of intraplate deformation and its relationship with changing plate boundary configurations, subducting slab geometries, and absolute plate motion is poorly understood. We utilise a buoyancy driven Stokes flow solver, BEM-Earth, to investigate the contribution of subducting slabs through time on Pacific Plate motion and plate-scale deformation, and how this is linked to intraplate volcanism. We produce a series of geodynamic models from 62 to 42 Ma in which the plates are driven by the attached subducting slabs and mantle drag/suction forces. We compare our modelled intraplate deformation history with those types of intraplate volcanism that lack a clear age progression. Our models suggest that changes in Cenozoic subduction zone topology caused intraplate deformation to trigger volcanism along several linear seafloor structures, mostly by reactivation of existing seamount chains, but occasionally creating new volcanic chains on crust weakened by fracture zones and extinct ridges. Around 55 Ma subduction of the Pacific-Izanagi ridge reconfigured the major tectonic forces acting on the plate by replacing ridge push with slab pull along its north-western perimeter, causing lithospheric extension along pre-existing weaknesses. Large scale deformation observed in the models coincides with the seamount chains of Hawaii, Louisville, Tokelau, and Gilbert during our modelled time period of 62 to 42 Ma. We suggest that extensional stresses between 72 and 52 Ma are the likely cause of large parts of the formation of the Gilbert chain and that localised extension between 62 and 42 Ma could cause late-stage volcanism along the Musicians Volcanic Ridges. Our models demonstrate that early Cenozoic changes in Pacific plate driving forces only cause relatively minor changes in Pacific absolute plate motions, and cannot be responsible for the Hawaii-Emperor Bend (HEB), confirming previous interpretations that the 47 Ma HEB does not reflect an absolute plate motion event.

Description: Aspects of hydrography and large-scale circulation observed in the Mediterranean Sea during the M84/3 and P414 cruises (April and June 2011, respectively) are presented. In contrast to most of the recent expeditions, which were limited to special areas of the basin, these two cruises, especially the M84/3, offered the opportunity of delineating a quasi-synoptic picture of the distribution of the relevant physical parameters along a section extending through the whole Mediterranean, from the Lebanese coast up to the Strait of Gibraltar. The foci of our analysis are the observed water mass properties and velocity fields. The first are investigated through T–S diagrams and an optimum multiparameter (OMP) analysis and the results are discussed also in the context of recently identified modes of variability; the second are studied by comparing the velocity fields observed using a vessel-mounted Acoustic Doppler Current Profiler and those calculated from the observed density fields. Overall, a distribution of temperature, salinity and geostrophic velocities emerges, which is far from that observed before the beginning of the so-called "Eastern Mediterranean Transient", a major climatic shift in the hydrography and circulation of the Mediterranean Sea which began at the end of the 1980s. The picture which emerges helps to further address the complexity of long-term evolution of hydrography and large-scale circulation of the Mediterranean Sea.

Description: Free-ocean CO2 enrichment (FOCE) systems are designed to assess the impact of ocean acidification on biological communities in situ for extended periods of time (weeks to months). They overcome some of the drawbacks of laboratory experiments and field observations by enabling (1) precise control of CO2 enrichment by monitoring pH as an offset of ambient pH, (2) consideration of indirect effects such as those mediated through interspecific relationships and food webs, and (3) relatively long experiments with intact communities. Bringing perturbation experiments from the laboratory to the field is, however, extremely challenging. The main goal of this paper is to provide guidelines on the general design, engineering, and sensor options required to conduct FOCE experiments. Another goal is to introduce xFOCE, a community-led initiative to promote awareness, provide resources for in situ perturbation experiments, and build a user community. Present and existing FOCE systems are briefly described and examples of data collected presented. Future developments are also addressed as it is anticipated that the next generation of FOCE systems will include, in addition to pH, options for oxygen and/or temperature control. FOCE systems should become an important experimental approach for projecting the future response of marine ecosystems to environmental change.

Description: Vertebrate innate immunity is the first line of defense against an invading pathogen and has long been assumed to be largely unspecific with respect to parasite/pathogen species. However, recent phenotypic evidence suggests that immunogenetic variation, i.e. allelic variability in genes associated with the immune system, results in host-parasite genotype-by-genotype interactions and thus specific innate immune responses. Immunogenetic variation is common in all vertebrate taxa and this reflects an effective immunological function in complex environments. However, the underlying variability in host gene expression patterns as response of innate immunity to within-species genetic diversity of macroparasites in vertebrates is unknown. We hypothesized that intra-specific variation among parasite genotypes must be reflected in host gene expression patterns. Here we used high-throughput RNA-sequencing to examine the effect of parasite genotypes on gene expression patterns of a vertebrate host, the three-spined stickleback (Gasterosteus aculeatus). By infecting naïve fish with distinct trematode genotypes of the species Diplostomum pseudospathaceum we show that gene activity of innate immunity in three-spined sticklebacks depended on the identity of an infecting macroparasite genotype. In addition to a suite of genes indicative for a general response against the trematode we also find parasite-strain specific gene expression, in particular in the complement system genes, despite similar infection rates of single clone treatments. The observed discrepancy between infection rates and gene expression indicates the presence of alternative pathways which execute similar functions. This suggests that the innate immune system can induce redundant responses specific to parasite genotypes.

Description: The Boknis Eck (BE) time series station, initiated in 1957, is one of the longest-operated time series stations worldwide. We present the first statistical evaluation of a data set of nine physical, chemical and biological parameters in the period of 1957–2013. In the past three to five decades, all of the measured parameters underwent significant long-term changes. Most striking is an ongoing decline in bottom water oxygen concentration, despite a significant decrease of nutrient and chlorophyll a concentrations. Temperature-enhanced oxygen consumption in the bottom water and a prolongation of the stratification period are discussed as possible reasons for the ongoing oxygen decline despite declining eutrophication. Observations at the BE station were compared with model output of the Kiel Baltic Sea Ice Ocean Model (BSIOM). Reproduced trends were in good agreement with observed trends for temperature and oxygen, but generally the oxygen concentration at the bottom has been overestimated.

Description: Sulfurimonas denitrificans was originally isolated from coastal marine sediments. It can grow with thiosulfate and nitrate or sulfide and oxygen. Recently sequencing of its genome revealed that it encodes periplasmic and cytoplasmic [NiFe]-hydrogenases but the role of hydrogen for its metabolism has remained unknown. We show the first experimental evidence that S. denitrificans can indeed express a functional hydrogen uptake active hydrogenase and can grow on hydrogen. In fact, under the provided conditions it grew faster and denser on hydrogen than on thiosulfate alone and even grew with hydrogen in the absence of reduced sulfur compounds. In our experiments, at the time points tested, the hydrogen uptake activity appeared to be related to the periplasmic hydrogenase and not to the cytoplasmic hydrogenase. Our data suggest that under the provided conditions S. denitrificans can grow more efficiently with hydrogen than with thiosulfate.

Description: Mineral dust supplied to remote ocean regions stimulates phytoplankton growth through delivery of micronutrients, notably iron (Fe). Although attention is usually paid to Fe (hydr)oxides as major sources of available Fe, Fe-bearing clay minerals are typically the dominant phase in mineral dust. The mineralogy and chemistry of clay minerals in dust particles, however, are largely unknown. We conducted microscopic identification and chemical analysis of the clay minerals in Asian and Saharan dust particles. Cross-sectional slices of dust particles were prepared by focused ion beam (FIB) techniques and analyzed by transmission electron microscopy (TEM) combined with energy dispersive X-ray spectroscopy (EDXS). TEM images of FIB slices revealed that clay minerals occurred as either nano-thin platelets or relatively thick plates. Chemical compositions and lattice fringes of the nano-thin platelets suggested that they included illite, smectite, illite–smectite mixed layers, and their nanoscale mixtures (illite–smectite series clay minerals, ISCMs) which could not be resolved with an electron microbeam. EDXS chemical analysis of the clay mineral grains revealed that the average Fe content was 5.8% in nano-thin ISCM platelets assuming 14% H2O, while the Fe content of illite and chlorite was 2.8 and 14.8%, respectively. In addition, TEM and EDXS analyses were performed on clay mineral grains dispersed and loaded on micro-grids. The average Fe content of clay mineral grains was 6.7 and 5.4% in Asian and Saharan dusts, respectively. A comparative X-ray diffraction analysis of bulk dusts showed that Saharan dust was more enriched in clay minerals than Asian dust, while Asian dust was more enriched in chlorite. Clay minerals, in particular nanocrystalline ISCMs and Fe-rich chlorite, are probably important sources of Fe to remote marine ecosystems. Further detailed analyses of the mineralogy and chemistry of clay minerals in global mineral dusts are required to evaluate the inputs of Fe to surface ocean microbial communities.

Description: Fast Repetition Rate fluorometry (FRRf) measurements of phytoplankton photophysiology from an acrossbasin South Atlantic cruise (as part of the GEOTRACES programme) characterised two dominant ecophysiological regimes which were interpreted on the basis of nutrient limitation. South of the South Subtropical Convergence (SSTC) in the northern sub-Antarctic sector of the Antarctic Circumpolar Current (ACC) in the Eastern Atlantic Basin, waters are characterised by elevated chlorophyll concentrations, a dominance by larger phytoplankton cells, and low apparent photochemical efficiency (Fv / Fm). Shipboard 24 h iron (Fe) addition incubation experiments confirmed that Fe stress was primarily responsible for the low Fv / Fm, with Fe addition to these waters, either within the artificial bottle additions or naturally occurring downstream enrichment from Gough Island, significantly increasing Fv / Fm values. To the north of the SSTC at the southern boundary of the South Atlantic Gyre, phytoplankton are characterised by high values of Fv / Fm which, coupled with the low macronutrient concentrations and increased presence of picocyanobacteria, are interpreted as conditions of Fe replete, balanced macronutrient-limited growth. Spatial correlation was found between Fv / Fm and Fe:nitrate ratios, supporting the suggestion that the relative supply ratios of these two nutrients can control patterns of limitation and consequently the ecophysiology of phytoplankton in subtropical gyre and ACC regimes.

Description: This paper is the outcome of a workshop held in Rome in November 2011 on the occasion of the 25th anniversary of the POEM (Physical Oceanography of the Eastern Mediterranean) program. In the workshop discussions, a number of unresolved issues were identified for the physical and biogeochemical properties of the Mediterranean Sea as a whole, i.e., comprising the Western and Eastern sub-basins. Over the successive two years, the related ideas were discussed among the group of scientists who participated in the workshop and who have contributed to the writing of this paper. Three major topics were identified, each of them being the object of a section divided into a number of different sub-sections, each addressing a specific physical, chemical or biological issue: 1. Assessment of basin-wide physical/biochemical properties, of their variability and interactions. 2. Relative importance of external forcing functions (wind stress, heat/moisture fluxes, forcing through straits) vs. internal variability. 3. Shelf/deep sea interactions and exchanges of physical/biogeochemical properties and how they affect the sub-basin circulation and property distribution. Furthermore, a number of unresolved scientific/methodological issues were also identified and are reported in each sub-section after a short discussion of the present knowledge. They represent the collegial consensus of the scientists contributing to the paper. Naturally, the unresolved issues presented here constitute the choice of the authors and therefore they may not be exhaustive and/or complete. The overall goal is to stimulate a broader interdisciplinary discussion among the scientists of the Mediterranean oceanographic community, leading to enhanced collaborative efforts and exciting future discoveries.

Description: We suggest the application of a flux parameterization commonly used over terrestrial areas for calculation of CO2 fluxes over sea ice surfaces. The parameterization is based on resistance analogy. We present a concept for parameterization of the CO2 fluxes over sea ice suggesting to use properties of the atmosphere and sea ice surface that can be measured or calculated on a routine basis. Parameters, which can be used in the conceptual model, are analysed based on data sampled from a seasonal fast-ice area, and the different variables influencing the exchange of CO2 between the atmosphere and ice are discussed. We found the flux to be small during the late winter with fluxes in both directions. Not surprisingly we find that the resistance across the surface controls the fluxes and detailed knowledge of the brine volume and carbon chemistry within the brines as well as knowledge of snow cover and carbon chemistry in the ice are essential to estimate the partial pressure of pCO(2) and CO2 flux. Further investigations of surface structure and snow cover and driving parameters such as heat flux, radiation, ice temperature and brine processes are required to adequately parameterize the surface resistance.

Description: The termination of the African Humid Period in northeastern Africa during the early Holocene was marked by the southward migration of the rain belt and the disappearance of the Green Sahara. This interval of drastic environmental changes was also marked by the initiation of food production by North African huntergatherer populations and thus provides critical information on human-environment relationships. However, existing records of regional climatic and environmental changes exhibit large differences in timing and modes of the wet/dry transition at the end of the African Humid Period. Here we present independent records of changes in river runoff, vegetation and erosion in the Nile River watershed during the Holocene obtained from a unique sedimentary sequence on the Nile River fan using organic and inorganic proxy data. This high-resolution reconstruction allows to examine the phase relationship between the changes of these three parameters and provides a detailed picture of the environmental conditions during the Paleolithic/Neolithic transition. The data show that river runoff decreased gradually during the wet/arid transition at the end of the AHP whereas rapid shifts of vegetation and erosion occurred earlier between 8.7 and ,6 ka BP. These asynchronous changes are compared to other regional records and provide new insights into the threshold responses of the environment to climatic changes. Our record demonstrates that the degradation of the environment in northeastern Africa was more abrupt and occurred earlier than previously thought and may have accelerated the process of domestication in order to secure sustainable food resources for the Neolithic African populations.

Description: A growing body of work on the neuroethology of cubozoans is based largely on the capabilities of the photoreceptive tissues, and it is important to determine the molecular basis of their light sensitivity. The cubozoans rely on 24 special purpose eyes to extract specific information from a complex visual scene to guide their behavior in the habitat. The lens eyes are the most studied photoreceptive structures, and the phototransduction in the photoreceptor cells is based on light sensitive opsin molecules. Opsins are photosensitive transmembrane proteins associated with photoreceptors in eyes, and the amino acid sequence of the opsins determines the spectral properties of the photoreceptors. Here we show that two distinct opsins (Tripedalia cystophora-lens eye expressed opsin and Tripedalia cystophora-neuropil expressed opsin, or Tc-leo and Tc-neo) are expressed in the Tripedalia cystophora rhopalium. Quantitative PCR determined the level of expression of the two opsins, and we found Tc-leo to have a higher amount of expression than Tc-neo. In situ hybridization located Tc-leo expression in the retinal photoreceptors of the lens eyes where the opsin is involved in image formation. Tc-neo is expressed in a confined part of the neuropil and is probably involved in extraocular light sensation, presumably in relation to diurnal activity.

Description: Significant changes in the overturning circulation of the Mediterranean Sea has been observed during the last few decades, the most prominent phenomena being the Eastern Mediterranean Transient (EMT) in the early 1990s and the Western Mediterranean Transition (WMT) during the mid-2000s. During both of these events unusually large amounts of deep water were formed, and in the case of the EMT, the deep water formation area shifted from the Adriatic to the Aegean Sea. Here we synthesize a unique collection of transient tracer (CFC-12, SF6 and tritium) data from nine cruises conducted between 1987 and 2011 and use these data to determine temporal variability of Mediterranean ventilation. We also discuss biases and technical problems with transient tracer-based ages arising from their different input histories over time; particularly in the case of time-dependent ventilation. We observe a period of low ventilation in the deep eastern (Levantine) basin after it was ventilated by the EMT so that the age of the deep water is increasing with time. In the Ionian Sea, on the other hand, we see evidence of increased ventilation after year 2001, indicating the restarted deep water formation in the Adriatic Sea. This is also reflected in the increasing age of the Cretan Sea deep water and decreasing age of Adriatic Sea deep water since the end of the 1980s. In the western Mediterranean deep basin we see the massive input of recently ventilated waters during the WMT. This signal is not yet apparent in the Tyrrhenian Sea, where the ventilation seems to be fairly constant since the EMT. Also the western Alboran Sea does not show any temporal trends in ventilation.

Description: This study presents the first quantitative estimate of the mineral dust emission associated with atmospheric depressions and mobile cyclones in North Africa. Atmospheric depressions are automatically tracked at 925 hPa based on ERA-Interim data from the European Centre for Medium-Range Weather Forecasts for 1989–2008. A set of filter criteria is applied to identify mobile cyclones, i.e. migrating and long-lived cyclones. The shorter term cyclone is used as a synonym for mobile cyclones. Dust emission is calculated with a dust emission model driven by 10 m winds and soil moisture from ERA-Interim. Emission peaks during winter and spring with spatial averages of 250–380 g m−2 per month. Comparison of the dust source activation frequency from the model against SEVIRI satellite observation shows a good agreement in the Bodélé Depression but differences in the north and west of North Africa. Depressions are abundant, particularly in summer when the Saharan heat low is situated over West Africa and during spring in the lee of the Atlas Mountains. Up to 90% (55% annually and spatially averaged) of dust emission occurs within 10 degrees of these depressions, with embedded mechanisms such as nocturnal low-level jets playing a role. Cyclones are rarer and occur primarily north of 20° N in spring in agreement with previous studies and over summertime West Africa consistent with near-surface signatures of African Easterly Waves. Dust emission within 10 degrees of cyclones peaks over Libya with up to 25% in spring. Despite the overall small contribution of 4% annually and spatially averaged, cyclones coincide with particularly intense dust emission events exceeding the climatological mean by a factor of four to eight. Soil moisture weakens dust emission during cyclone passage by about 10%.

Description: We present a new coupled ocean-circulation–ice model configuration of the Baltic Sea. The model features, contrary to most existing configurations, a high horizontal resolution of ≈ 1 nautical mile (≈ 1.85 km), which is eddy-resolving over much of the domain. The vertical discretisation comprises a total of 47 vertical levels. Results from a 1987 to 1999 hindcast simulation show that the model's fidelity is competitive. As suggested by a comparison with sea surface temperatures observed from space, this applies especially to near-surface processes. Hence, the configuration is well suited to serve as a nucleus of a fully fledged coupled ocean-circulation–biogeochemical model (which is yet to be developed). A caveat is that the model fails to reproduce major inflow events. We trace this back to spurious vertical circulation patterns at the sills which may well be endemic to high-resolution models based on geopotential coordinates. Further, we present indications that – so far neglected – eddy/wind effects exert significant control on wind-induced up- and downwelling.

Description: The increasing pCO2 in seawater is a serious threat for marine calcifiers and alters the biogeochemistry of the ocean. Therefore, the reconstruction of past-seawater properties and their impact on marine ecosystems is an important way to investigate the underlying mechanisms and to better constrain the effects of possible changes in the future ocean. Cold-water coral (CWC) ecosystems are biodiversity hotspots. Living close to aragonite-undersaturation, these corals serve as living laboratories as well as archives to reconstruct the boundary conditions of their calcification under the carbonate system of the ocean. We investigated the reef-building CWC Lophelia pertusa as a recorder of intermediate ocean seawater pH. This species-specific field calibration is based on a unique sample set of live in-situ collected L. pertusa and corresponding seawater samples. These data demonstrate that uranium speciation and skeletal incorporation for azooxanthellate scleractinian CWCs is pH dependent. However, this also indicates that internal pH up-regulation of the coral does not play a role in uranium incorporation into the majority of the skeleton of L. pertusa. This study suggests L. pertusa provides a new archive for the reconstruction of intermediate water mass pH and hence may help to constrain tipping points for ecosystem dynamics and evolutionary characteristics in a changing ocean.

Description: Cold-water coral reefs are known to locally enhance the diversity of deep-sea fauna as well as of microbes. Sponges are among the most diverse faunal groups in these ecosystems, and many of them host large abundances of microbes in their tissues. In this study, twelve sponge species from three cold-water coral reefs off Norway were investigated for the relationship between sponge phylogenetic classification (species and family level), as well as sponge type (high versus low microbial abundance), and the diversity of sponge-associated bacterial communities, taking also geographic location and water depth into account. Community analysis by Automated Ribosomal Intergenic Spacer Analysis (ARISA) showed that as many as 345 (79%) of the 437 different bacterial operational taxonomic units (OTUs) detected in the dataset were shared between sponges and sediments, while only 70 (16%) appeared purely sponge-associated. Furthermore, changes in bacterial community structure were significantly related to sponge species (63% of explained community variation), sponge family (52%) or sponge type (30%), whereas mesoscale geographic distances and water depth showed comparatively small effects (〈5% each). In addition, a highly significant, positive relationship between bacterial community dissimilarity and sponge phylogenetic distance was observed within the ancient family of the Geodiidae. Overall, the high diversity of sponges in cold-water coral reefs, combined with the observed sponge-related variation in bacterial community structure, support the idea that sponges represent heterogeneous, yet structured microbial habitats that contribute significantly to enhancing bacterial diversity in deep-sea ecosystems.

Description: In order to test the influences of ocean acidification on the ocean pelagic ecosystem, so far the largest CO2 manipulation mesocosm study (European Project on Ocean Acidification, EPOCA) was performed in Kings Bay (Kongsfjorden), Spitsbergen. During a 30 day incubation, bacterial diversity was investigated using DNA fingerprinting and clone library analysis of bacterioplankton samples. Terminal restriction fragment length polymorphism (T-RFLP) analysis of the PCR amplicons of the 16S rRNA genes revealed that general bacterial diversity, taxonomic richness and community structure were influenced by the variation of productivity during the time of incubation, but not the degree of ocean acidification. A BIOENV analysis suggested a complex control of bacterial community structure by various biological and chemical environmental parameters. The maximum apparent diversity of bacterioplankton (i.e., the number of T-RFs) in high and low pCO2 treatments differed significantly. A negative relationship between the relative abundance of Bacteroidetes and pCO2 levels was observed for samples at the end of the experiment by the combination of T-RFLP and clone library analysis. Our study suggests that ocean acidification affects the development of bacterial assemblages and potentially impacts the ecological function of the bacterioplankton in the marine ecosystem.

Description: Cold-water coral (CWC) reefs are heterogeneous ecosystems comprising numerous microhabitats. A typical European CWC reef provides various biogenic microhabitats (within, on and surrounding colonies of coral species such as Lophelia pertusa, Paragorgia arborea and Primnoa resedaeformis, or formed by their remains after death). These microhabitats may be surrounded and intermixed with non-biogenic microhabitats (soft sediment, hard ground, gravel/pebbles, steep walls). To date, studies of distribution of sessile fauna across CWC reefs have been more numerous than those investigating mobile fauna distribution. In this study we quantified shrimp densities associated with key CWC microhabitat categories at the Røst Reef, Norway, by analysing image data collected by towed video sled in June 2007. We also investigated shrimp distribution patterns on the local scale (〈40 cm) and how these may vary with microhabitat. Shrimp abundances at the Røst Reef were on average an order of magnitude greater in biogenic reef microhabitats than in non-biogenic microhabitats. Greatest shrimp densities were observed in association with live Paragorgia arborea microhabitat (43 shrimp m−2, SD = 35.5), live Primnoa resedaeformis microhabitat (41.6 shrimp m−2, SD = 26.1) and live Lophelia pertusa microhabitat (24.4 shrimp m−2, SD = 18.6). In non-biogenic microhabitat, shrimp densities were 〈2 shrimp m−2. CWC reef microhabitats appear to support greater shrimp densities than the surrounding non-biogenic microhabitats at the Røst Reef, at least at the time of survey.

Description: Submarine canyons are known as one of the seafloor morphological features where living cold-water coral (CWC) communities develop in the Mediterranean Sea. We investigated the CWC community of the two westernmost submarine canyons of the Gulf of Lions canyon system: the Cap de Creus Canyon (CCC) and Lacaze-Duthiers Canyon (LDC). Coral associations have been studied through video material recorded by means of a manned submersible and a remotely operated vehicle. Video transects have been conducted and analyzed in order to obtain information on (1) coral bathymetric distribution and density patterns, (2) size structure of coral populations, and (3) coral colony position with respect to the substrate. Madrepora oculata was the most abundant CWC in both canyons, while Lophelia pertusa and Dendrophyllia cornigera mostly occurred as isolated colonies or in small patches. An important exception was detected in a vertical cliff in LDC where a large L. pertusa framework was documented. This is the first record of such an extended L. pertusa framework in the Mediterranean Sea. In both canyons coral populations were dominated by medium and large colonies, but the frequent presence of small-sized colonies also indicate active recruitment. The predominant coral orientation (90° and 135°) is probably driven by the current regime as well as by the sediment load transported by the current flows. In general, no clear differences were observed in the abundance and in the size structure of the CWC populations between CCC and LDC, despite large differences in particulate matter between canyons.

Description: All known photoreceptor cells adapt to constant light stimuli, fading the retinal image when exposed to an immobile visual scene. Counter strategies are therefore necessary to prevent blindness, and in mammals this is accomplished by fixational eye movements. Cubomedusae occupy a key position for understanding the evolution of complex visual systems and their eyes are assumedly subject to the same adaptive problems as the vertebrate eye, but lack motor control of their visual system. The morphology of the visual system of cubomedusae ensures a constant orientation of the eyes and a clear division of the visual field, but thereby also a constant retinal image when exposed to stationary visual scenes. Here we show that bell contractions used for swimming in the medusae refresh the retinal image in the upper lens eye of Tripedalia cystophora. This strongly suggests that strategies comparable to fixational eye movements have evolved at the earliest metazoan stage to compensate for the intrinsic property of the photoreceptors. Since the timing and amplitude of the rhopalial movements concur with the spatial and temporal resolution of the eye it circumvents the need for post processing in the central nervous system to remove image blur.

Description: The Mediterranean Sea is a semi-enclosed sea characterized by high salinities, temperatures and densities. The net evaporation exceeds the precipitation, driving an anti-estuarine circulation through the Strait of Gibraltar, contributing to very low nutrient concentrations. The Mediterranean Sea has an active overturning circulation, one shallow cell that communicates directly with the Atlantic Ocean, and two deep overturning cells, one in each of the two main basins. It is surrounded by populated areas and is thus sensitive to anthropogenic forcing. Several dramatic changes in the oceanographic and biogeochemical conditions have been observed during the past several decades, emphasizing the need to better monitor and understand the changing conditions and their drivers. During 2011 three oceanographic cruises were conducted in a coordinated fashion in order to produce baseline data of important physical and biogeochemical parameters that can be compared to historic data and be used as reference for future observational campaigns. In this article we provide information on the Mediterranean Sea oceanographic situation, and present a short review that will serve as background information for the special issue in Ocean Science on "Physical, chemical and biological oceanography of the Mediterranean Sea". An important contribution of this article is the set of figures showing the large-scale distributions of physical and chemical properties along the full length

Description: Recent studies on the impacts of ocean acidification on pelagic communities have identified changes in carbon to nutrient dynamics with related shifts in elemental stoichiometry. In principle, mesocosm experiments provide the opportunity of determining temporal dynamics of all relevant carbon and nutrient pools and, thus, calculating elemental budgets. In practice, attempts to budget mesocosm enclosures are often hampered by uncertainties in some of the measured pools and fluxes, in particular due to uncertainties in constraining air–sea gas exchange, particle sinking, and wall growth. In an Arctic mesocosm study on ocean acidification applying KOSMOS (Kiel Off-Shore Mesocosms for future Ocean Simulation), all relevant element pools and fluxes of carbon, nitrogen and phosphorus were measured, using an improved experimental design intended to narrow down the mentioned uncertainties. Water-column concentrations of particulate and dissolved organic and inorganic matter were determined daily. New approaches for quantitative estimates of material sinking to the bottom of the mesocosms and gas exchange in 48 h temporal resolution as well as estimates of wall growth were developed to close the gaps in element budgets. However, losses elements from the budgets into a sum of insufficiently determined pools were detected, and are principally unavoidable in mesocosm investigation. The comparison of variability patterns of all single measured datasets revealed analytic precision to be the main issue in determination of budgets. Uncertainties in dissolved organic carbon (DOC), nitrogen (DON) and particulate organic phosphorus (POP) were much higher than the summed error in determination of the same elements in all other pools. With estimates provided for all other major elemental pools, mass balance calculations could be used to infer the temporal development of DOC, DON and POP pools. Future elevated pCO2 was found to enhance net autotrophic community carbon uptake in two of the three experimental phases but did not significantly affect particle elemental composition. Enhanced carbon consumption appears to result in accumulation of dissolved organic carbon under nutrient-recycling summer conditions. This carbon over-consumption effect becomes evident from mass balance calculations, but was too small to be resolved by direct measurements of dissolved organic matter. Faster nutrient uptake by comparatively small algae at high CO2 after nutrient addition resulted in reduced production rates under future ocean CO2 conditions at the end of the experiment. This CO2 mediated shift towards smaller phytoplankton and enhanced cycling of dissolved matter restricted the development of larger phytoplankton, thus pushing the system towards a retention type food chain with overall negative effects on export potential.

Description: Here we report on data from an oceanographic cruise on the German research vessel Meteor covering large parts of the Mediterranean Sea during spring of 2011. The main objectives of this cruise was to conduct measurements of physical, chemical and biological variables on a section across the Mediterranean Sea with the goal of producing a synoptic picture of the distribution of relevant physical and biogeochemical properties, in order to compare those to historic data sets. During the cruise, a comprehensive data set of relevant variables following the guide lines for repeat hydrography outlined by the GO-SHIP group (http://www.go-ship.org/) was collected. The measurements include; salinity and temperature (CTD), an over-determined carbonate system, inorganic nutrients, oxygen, transient tracers (CFC-12, SF6), Helium isotopes and tritium, and carbon isotopes. The cruise sampled all major basins of the Mediterranean Sea following roughly an east-to-west section from the coast of Lebanon to through the Strait of Gibraltar, and to the coast of Portugal. Also a south-to-north section from the Ionian Sea to the Adriatic Sea was carried out. Additionally, sampling in the Aegean, Adriatic and Tyrrhenian Seas were carried out. The sections roughly followed lines and positions that have been sampled previously during other programs, thus providing the opportunity for comparative investigations of the temporal development of various parameters.

Description: The oceanic biological carbon pump is an important factor in the global carbon cycle. Organic carbon is exported from the surface ocean mainly in the form of settling particles derived from plankton production in the upper layers of the ocean. The large variability in current estimates of the global strength of the biological carbon pump emphasises that our knowledge of a major planetary carbon flux remains poorly constrained. We present a database of 723 estimates of organic carbon export from the surface ocean derived from the 234Th technique. The dataset is archived on the data repository PANGEA® (www.pangea.de) under doi:10.1594/PANGAEA.809717. Data were collected from tables in papers published between 1985 and early 2013. We also present sampling dates, publication dates and sampling areas. Most of the open ocean provinces are represented by multiple measurements. However, the western Pacific, the Atlantic Arctic, South Pacific and the southern Indian Ocean are not well represented. There is a variety of integration depths ranging from surface to 300 m. Globally the fluxes ranged from 0 to 1500 mg C m−2 d−1.

Description: Variability and trends in seasonal and interannual ice area export out of the Laptev Sea between 1992 and 2011 are investigated using satellite-based sea ice drift and concentration data. We found an average total winter (October to May) ice area transport across the northern and eastern Laptev Sea boundaries (NB and EB) of 3.48 × 10hoch5 km2. The average transport across the NB (2.87 × 10hoch5 km2)is thereby higher than across the EB (0.61 × 10hoch5 km2), with a less pronounced seasonal cycle. The total Laptev Sea ice area flux significantly increased over the last decades (0.85 × 10hoch5 km2 decade−1, p 〉 0.95), dominated by increasing export through the EB (0.55 × 10hoch5 km2 decade−1, p 〉 0.90), while the increase in export across the NB is smaller (0.3 × 10hoch5 km2 decade−1) and statistically not significant. The strong coupling between across-boundary SLP gradient and ice drift velocity indicates that monthly variations in ice area flux are primarily controlled by changes in geostrophic wind velocities, although the Laptev Sea ice circulation shows no clear relationship with large-scale atmospheric indices. Also there is no evidence of increasing wind velocities that could explain the overall positive trends in ice export. The increased transport rates are rather the consequence of a changing ice cover such as thinning and/or a decrease in concentration. The use of a back-propagation method revealed that most of the ice that is incorporated into the Transpolar Drift is formed during freeze-up and originates from the central and western part of the Laptev Sea, while the exchange with the East Siberian Sea is dominated by ice coming from the central and southeastern Laptev Sea. Furthermore, our results imply that years of high ice export in late winter (February to May) have a thinning effect on the ice cover, which in turn preconditions the occurence of negative sea ice extent anomalies in summer.

Description: To examine the potentially competing influences of microzooplankton and calcite mineral ballast on organic matter remineralization, we incubated diatoms in darkness in rolling tanks with and without added calcite minerals (coccoliths) and microzooplankton (rotifers). Concentrations of particulate organic matter (POM in suspension or in aggregates), of dissolved organic matter (DOM), and of dissolved inorganic nutrients were monitored over 8 days. The presence of rotifers enhanced the remineralization of ammonium and phosphate, but not dissolved silicon, from the biogenic particulate matter, up to 40% of which became incorporated into aggregates early in the experiment. Added calcite resulted in rates of excretion of ammonium and phosphate by rotifers that were depressed by 67% and 36%, respectively, demonstrating the potential for minerals to inhibit the destruction of POM by zooplankton in the water column. Lastly, the presence of the rotifers and added calcite minerals resulted in a more rapid initial rate of aggregation, although not a greater overall amount of aggregation during the experiment.

Description: Meridional and vertical distributions of several biogeochemical parameters were studied along a section in the southeastern Atlantic and the Southern Ocean south of South Africa during the austral summer 2008 of the International Polar Year to characterize the biogeochemical provinces and to assess the seasonal net diatom production. Based on analyses of macro-nutrients, ammonium (NH4), chlorophyll a, (Chl a), phaeopigments, biogenic silica (BSi), particulate inorganic carbon (PIC), and particulate organic carbon and nitrogen (POC and PON, respectively), four biogeochemical domains were distinguished along the section: the subtropical Atlantic, the confluence zone of the subtropical and subantarctic domains, the Polar Frontal Zone (PFZ) in the Antarctic Circumpolar Current (ACC), and the north-eastern branch of the Weddell Gyre. The subtropical region displayed extremely low nutrient concentrations featuring oligotrophic conditions, and sub-surface maxima of Chl a and phaeopigments never exceeded 0.5 µg L−1 and 0.25 µg L−1, respectively. The anticyclonic and cyclonic eddies crossed in the Cape Basin were characterized by a deepening and a rise, respectively, of the nutrients isoclines. The confluence zone of the subtropical domain and the northern side of the ACC within the subantarctic domain displayed remnant nitrate and phosphate levels, whereas silicate concentrations kept to extremely low levels. In this area, Chl a level of 0.4–0.5 µg L−1 distributed homogenously within the mixed layer, and POC and PON accumulated to values up to 10 µM and 1.5 µM, respectively, indicative of biomass accumulation along the confluence zone during the late productive period. In the ACC domain, the Polar Frontal Zone was marked by a post-bloom of diatoms that extended beyond the Polar Front (PF) during this late summer condition, as primarily evidenced by the massive depletion of silicic acid in the surface waters. The accumulation of NH4 to values up to 1.25 µM at 100 m depth centred on the PF and the accumulation of BSi up to 0.5 µM in the surface waters of the central part of the PFZ also featured a late stage of the seasonal diatom bloom. The silica daily net production rate based on the seasonal depletion of silicic acid was estimated to be 11.9 ± 6.5 mmol m−2 d−1 in the domain of the vast diatom post-bloom, agreeing well with the previously recorded values in this province. The Weddell Gyre occasionally displayed relative surface depletion of silicic acid, suggesting a late stage of a relatively minor diatom bloom possibly driven by iceberg drifting releases of iron. In this domain the estimated range of silica daily net production rate (e.g. 21.1 ± 8.8 mmol m−2 d−1) is consistent with previous studies, but was not significantly higher than that in the Polar Front region.

Description: Samoylov Island is centrally located within the Lena River Delta at 72° N, 126° E and lies within the Siberian zone of continuous permafrost. The landscape on Samoylov Island consists mainly of late Holocene river terraces with polygonal tundra, ponds and lakes, and an active floodplain. The island has been the focus of numerous multidisciplinary studies since 1993, which have focused on climate, land cover, ecology, hydrology, permafrost and limnology. This paper aims to provide a framework for future studies by describing the characteristics of the island's meteorological parameters (temperature, radiation and snow cover), soil temperature, and soil moisture. The land surface characteristics have been described using high resolution aerial images in combination with data from ground-based observations. Of note is that deeper permafrost temperatures have increased between 0.3 to 1.3 °C over the last five years. However, no clear warming of air and active layer temperatures is detected since 1998, though winter air temperatures during recent years have not been as cold as in earlier years.