Stareso MedSeA mesocosm experiment: field and perturbation studies 2012

A joint mesocosm experiment took place in June/July 2012 in Corsica (bay of Calvi, Stareso station;http://www.stareso.com/) as part of the european MedSeA project. Nine mesocosms (52 m**3) were deployed over a 20 days period and 6 different levels of pCO2 and 3 control mesocosms (about 450 µatm), were used, in order to cover the range of pCO2 anticipated for the end of the present century. During this experiment, the potential effects of these perturbations on chemistry, planktonic community composition and dynamics including: eucaryotic and prokaryotic species composition, primary production, nutrient and carbon utilization, calcification, diazotrophic nitrogen fixation, organic matter exudation and composition, micro-layer composition and biogas production were studied by a group of about 25 scientists from 8 institutes and 6 countries. This is one of the first mesocosm experiments conducted in oligotrophic waters. A blog dedicated to this experiment can be viewed at: http://medseastareso2012.wordpress.com/.

Calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea

The distribution of calcareous dinoflagellate cysts in surface sediments from the Mediterranean Sea was quantitatively analysed. The samples contain 11 cyst species and the vegetative coccoid Thoracosphaera heimii. Cyst abundance increases towards the deeper parts of the basins and is generally higher in the eastern Mediterranean Sea. Three major distribution characteristics exist: (1) different assemblages in oceanic and neritic regions, (2) little agreement with the associations of areas studied so far like the Atlantic Ocean, and (3) a unique oceanic assemblage in the eastern Mediterranean Sea. A gradual change in cyst assemblages from the western to the eastern Mediterranean Sea was observed and statistically compared with the main environmental gradients in the upper water column. Temperature, nitrate concentration and possibly salinity appear to be the most important factors controlling cyst production. Three groups containing cysts with similar environmental preferences can be distinguished: (1) an eastern Mediterranean group related to relatively high temperature and salinity but low nitrate concentration, (2) a group of more or less consistently abundant cosmopolitan species tolerating or even preferring relatively low temperature and salinity but high nitrate concentration, and (3) a group containing species that are possibly adapted to neritic environments and have probably been transported from coastal areas into the studied regions. In contrast to other calcareous plankton, calcareous dinoflagellate cysts correlate strongly with the main environmental gradients in the Mediterranean Sea, bearing a high potential for palaeoenvironmental reconstructions.

Distribution of organic-walled dinoflagellate cysts in surface sediments of the Benguela upwelling system

To obtain insight in the relationship between the spatial distribution of organic-walled dinoflagellate cysts (dinocysts) and local environmental conditions, fifty-eight surface sediment samples from the coastal shelf off SW Africa were investigated on their dinocyst content with special focus on the two main river systems and the active upwelling that characterise this region. To avoid possible overprint by species-selective preservation, samples have been selected mainly from shelf sites where high sedimentation rates and/or low bottom water oxygen concentrations prevail. Multivariate ordination analyses have been carried out to investigate the relationship between the distribution patterns of individual species to environmental parameters of the upper water column and sediment transport processes. The main oceanographical variables at the surface (temperature, salinity, nutrients chlorophyll-a) in the region show onshore-offshore gradients. This pattern is reflected in the dinocyst associations with high relative abundances of heterotrophic dinocyst species in neritic regions characterised by high chlorophyll-aand low salinity conditions in surface waters. Phototrophic dinocyst species, notably Operculodinium centrocarpum, dominate in the more oceanic area. Differences in the distribution of phototrophic dinocyst species can be related to sea surface salinity and sea surface temperature gradients and to a lesser extent to chlorophyll-a concentrations. Apart from longitudinal gradients the dinocyst distribution clearly reflects regional environmental features. Six groups of species can be distinguished, characteristic for (1) coastal regions (cysts of Polykrikos kofoidii and Selenopemphix quanta), (2) the vicinity of active upwelling (Brigantedinium spp., Echinidinium aculeatum, Echinidinium spp. and Echinidinium transparantum), (3) river mouths (Lejeunecysta oliva, cysts of Protoperidinium americanum, Selenopemphix nephroides and Votadinium calvum), (4) slope and open ocean sediments (Dalella chathamense, Impagidinium patulum and Operculodinium centrocarpum, (5) the southern Benguela region (south of 24°S) (Spiniferites ramosus) and (6) the northern Benguela region (north of 24°S) (Nematosphaeropsis labyrinthus and Pyxidinopsis reticulata). No indication of overprint of the palaeo-ecological signal by lateral transport of allochthonous species could be observed.

Individual dinoflagellate cysts species of surface sediment samples from the Mediterranean Sea

To determine the relationship between the spatial dinoflagellate cyst distribution and oceanic environmental conditions, 34 surface sediments from the Eastern and Western Mediterranean Sea have been investigated for their dinoflagellate cyst content. Multivariate ordination analyses identified sea-surface temperature, chlorophyll-a , nitrate concentration, salinity, and bottom oxygen concentration as the main factors affecting dinoflagellate cyst distribution in the region. Based on the relative abundance data, two associations can be distinguished that can be linked with major oceanographic settings. (1) An offshore eastern Mediterranean regime where surface sediments are characterized by oligotrophic, warm, saline surface water, and high oxygen bottom water concentrations (Impagidinium species, Nematosphaeropsis labyrinthus, Pyxidinopsis reticulata and Operculodinium israelianum). Based on the absolute abundance, temperature is positively related to the cyst accumulation of Operculodinium israelianum. Temperature does not form a causal factor influencing the accumulation rate of the other species in this association. Impagidinium species and Nematosphaeropsis labyrinthus show a positive relationship between cyst accumulation and nitrate availability in the upper waters. (2) Species of association 2 have highest relative abundances in the Western Mediterranean Sea, Strait of Sicily/NW Ionian Sea, and/or the distal ends of the Po/Nile/Rhône River plumes. At these stations, surface waters are characterized by (relative to the other regime) higher productivity associated with lower sea-surface temperature, salinity, and lower bottom water oxygen concentrations (Selenopemphix nephroides, Echinidinium spp., Selenopemphix quanta, Quinquecuspis concreta, Brigantedinium spp. and Lingulodinium machaerophorum). Based on both the absolute and relative abundances, Selenopemphix nephroides is suggested to be a suitable indicator to trace changes in the trophic state of the upper waters. The distribution of Lingulodinium machaerophorum is related to the presence of river-influenced surface waters, notably the Nile River. We suggest that this species might form a suitable marker to trace past variations in river discharge, notably from the Nile.

Environmental controls on the Emiliania huxleyi calcite mass

Although ocean acidification is expected to impact (bio)calcification by decreasing the seawater carbonate ion concentration, [CO3]2-, there exists evidence of non-uniform response of marine calcifying plankton to low seawater [CO3]2-. This raises questions on the role of environmental factors other than acidification and on the complex physiological responses behind calcification. Here we investigate the synergistic effect of multiple environmental parameters, including temperature, nutrient (nitrate and phosphate) availability, and seawater carbonate chemistry on the coccolith calcite mass of the cosmopolitan coccolithophore Emiliania huxleyi, the most abundant species in the world ocean. We use a suite of surface (late Holocene) sediment samples from the South Atlantic and southwestern Indian Ocean taken from depths lying well above the modern lysocline. The coccolith calcite mass in our results presents a latitudinal distribution pattern that mimics the main oceanographic features, thereby pointing to the potential importance of phosphorus and temperature in determining coccolith mass by affecting primary calcification and possibly driving the E. huxleyi morphotype distribution. This evidence does not necessarily argue against the potentially important role of the rapidly changing seawater carbonate chemistry in the future, when unabated fossil fuel burning will likely perturb ocean chemistry beyond a critical point. Rather our study highlights the importance of evaluating the combined effect of several environmental stressors on calcifying organisms to project their physiological response(s) in a high CO2 world and improve interpretation of paleorecords.

Geographic distribution of dinoflagellate cysts in surface sediments

Dinoflagellate cysts are useful for reconstructing upper water conditions. For adequate reconstructions detailed information is required about the relationship between modern day environmental conditions and the geographic distribution of cysts in sediments. This Atlas summarises the modern global distribution of 71 organicwalled dinoflagellate cyst species. The synthesis is based on the integration of literature sources together with data of 2405 globally distributed surface sediment samples that have been preparedwith a comparable methodology and taxonomy. The distribution patterns of individual cyst species are being comparedwith environmental factors that are knownto influence dinoflagellate growth, gamete production, encystment, excystment and preservation of their organic-walled cysts: surface water temperature, salinity, nitrate, phosphate, chlorophyll-a concentrations and bottom water oxygen concentrations. Graphs are provided for every species depicting the relationship between seasonal and annual variations of these parameters and the relative abundance of the species. Results have been compared with previously published records; an overview of the ecological significance as well as information about the seasonal production of each individual species is presented. The relationship between the cyst distribution and variation in the aforementioned environmental parameters was analysed by performing a canonical correspondence analysis. All tested variables showed a positive relationship on the 99% confidence level. Sea-surface temperature represents the parameter corresponding to the largest amount of variance within the dataset (40%) followed by nitrate, salinity, phosphate and bottom-water oxygen concentration, which correspond to 34%, 33%, 25% and 24% of the variance, respectively. Characterisations of selected environments as well as a discussion about how these factors could have influenced the final cyst yield in sediments are included.

Compilation of downward flux observations from sediment trap deployments in the Atlantic Ocean - Contribution to EURO-BASIN's Data integration

We provide a compilation of downward fluxes (total mass, POC, PON, BSiO2, CaCO3, PIC and lithogenic/terrigenous fluxes) from over 6000 sediment trap measurements distributed in the Atlantic Ocean, from 30 degree North to 49 degree South, and covering the period 1982-2011. Data from the Mediterranean Sea are also included. Data were compiled from different sources: data repositories (BCO-DMO, PANGAEA), time series sites (BATS, CARIACO), published scientific papers and/or personal communications from PI's. All sources are specifed in the data set. Data from the World Ocean Atlas 2009 were extracted to provide each flux observation with contextual environmental data, such as temperature, salinity, oxygen (concentration, AOU and percentage saturation), nitrate, phosphate and silicate.

Total phospholipid fatty acid concentrations structrures of ODP Hole 191-1179 samples (Table 1)

Sediment samples ranging from 0.05 to 278 m below sea floor (mbsf) at a Northwest Pacific deep-water (5564 mbsl) site (ODP Leg 191, Site 1179) were analyzed for phospholipid fatty acids (PLFAs). Total PLFA concentrations decreased by a factor of three over the first meter of sediment and then decreased at a slower rate to approximately 30 mbsf. The sharp decrease over the first meter corresponds to the depth of nitrate and Mn(IV) reduction as indicated by pore water chemistry. PLFA-based cell numbers at site 1179 had a similar depth profile as that for Acridine orange direct cell counts previously made on ODP site 1149 sediments which have a similar water depth and lithology. The mole percentage of straight chain saturated PLFAs increases with depth, with a large shift between the 0.95 and 3.95 mbsf samples. PLFA stable carbon isotope ratios were determined for sediments from 0.05 to 4.53 mbsf and showed a general trend toward more depleted d13C values with depth. Both of these observations may indicate a shift in the bacterial community with depth across the different redox zones inferred from pore water chemistry data. The PLFA 10me16:0, which has been attributed to the bacterial genera Desulfobacter in many marine sediments, showed the greatest isotopic depletion, decreasing from -20 to -35 per mil over the first meter of sediment. Pore water chemistry suggested that sulfate reduction was absent or minimal over this same sediment interval. However, 10me16:0 has been shown to be produced by recently discovered anaerobic ammonium oxidizing (anammox) bacteria which are known chemoautotrophs. The increasing depletion in d13C of 10me16:0 with the unusually lower concentration of ammonium and linear decrease of nitrate concentration is consistent with a scenario of anammox bacteria mediating the oxidation of ammonium via nitrite, an intermediate of nitrate reduction.

Modelled niche centres and niche breadths of open ocean phytoplankton taxa

This data contains realized ecological niche estimates of phytoplankton taxa within the mixed layer of the open ocean. The estimates are based on data from the MARine Ecosystem DATa (MAREDAT) initiative, and cover five phytoplankton functional types: coccolithophores (40 species), diatoms (87 species), diazotrophs (two genera), Phaeocystis (two species) and picophytoplankton (two genera). Considered as major niche dimensions were temperature (°C), mixed layer depth (MLD; m), nitrate concentration (µmoles/L), mean photosynthetically active radiation in the mixed layer (MLPAR; µmoles/m**2/s), salinity, and the excess of phosphate versus nitrate relative to the Redfield ratio (P*; µmoles/L). For each niche dimension at a time, conditions at presence locations of the taxa were contrasted with conditions in 12 000 randomly sampled points from the open ocean using MaxEnt models. We used the quartiles of the response curves of these models to parameterize realized niche centers and niche breadths: the median (q50) of the response curves was considered to be the niche center and the distance between the lower quartile (q25) and the upper quartile (q75) was used as a rough estimate of niche breadth. We only reported meaningful niche estimates, i.e., estimates based on MaxEnt models that perform significantly better than random, as indicated by an area under the curve (AUC) score significantly larger than 0.5.

Nitrite consumption and associated isotope changes during a river flood event in the Elbe river

In oceans, estuaries, and rivers, nitrification is an important nitrate source, and stable isotopes of nitrate are often used to investigate recycling processes (e.g. remineralisation, nitrification) in the water column. Nitrification is a two-step process, where ammonia is oxidised via nitrite to nitrate. Nitrite usually does not accumulate in natural environments, which makes it difficult to study the single isotope effect of ammonia oxidation or nitrite oxidation in natural systems. However, during an exceptional flood in the Elbe River in June 2013, we found a unique co-occurrence of ammonium, nitrite, and nitrate in the water column, returning towards normal summer conditions within 1 week. Over the course of the flood, we analysed the evolution of d15N-[NH4]+ and d15N-[NO2]- in the Elbe River. In concert with changes in suspended particulate matter (SPM) and d15N SPM, as well as nitrate concentration, d15N-NO3 - and d18O-[NO3] -, we calculated apparent isotope effects during net nitrite and nitrate consumption. During the flood event, > 97 % of total reactive nitrogen was nitrate, which was leached from the catchment area and appeared to be subject to assimilation. Ammonium and nitrite concentrations increased to 3.4 and 4.4 µmol/l, respectively, likely due to remineralisation, nitrification, and denitrification in the water column. d15N-[NH4]+ values increased up to 12 per mil, and d15N-[NO2]- ranged from -8.0 to -14.2 per mil. Based on this, we calculated an apparent isotope effect 15-epsilon of -10.0 ± 0.1 per mil during net nitrite consumption, as well as an isotope effect 15-epsilon of -4.0 ± 0.1 per mil and 18-epsilon of -5.3 ± 0.1 per mil during net nitrate consumption. On the basis of the observed nitrite isotope changes, we evaluated different nitrite uptake processes in a simple box model. We found that a regime of combined riparian denitrification and 22 to 36 % nitrification fits best with measured data for the nitrite concentration decrease and isotope increase.

Average nutrient concentration, and growth characteristics of bacteria during USCGC Healy cruises in 2002 and 2004

Standing stocks and production rates for phytoplankton and heterotrophic bacteria were examined during four expeditions in the western Arctic Ocean (Chukchi Sea and Canada Basin) in the spring and summer of 2002 and 2004. Rates of primary production (PP) and bacterial production (BP) were higher in the summer than in spring and in shelf waters than in the basin. Most surprisingly, PP was 3-fold higher in 2004 than in 2002; ice-corrected rates were 1581 and 458 mg C/m**2/d respectively, for the entire region. The difference between years was mainly due to low ice coverage in the summer of 2004. The spatial and temporal variation in PP led to comparable variation in BP. Although temperature explained as much variability in BP as did PP or phytoplankton biomass, there was no relationship between temperature and bacterial growth rates above about 0°C. The average ratio of BP to PP was 0.06 and 0.79 when ice-corrected PP rates were greater than and less than 100 mg C/m**2/d, respectively; the overall average was 0.34. Bacteria accounted for a highly variable fraction of total respiration, from 3% to over 60% with a mean of 25%. Likewise, the fraction of PP consumed by bacterial respiration, when calculated from growth efficiency (average of 6.9%) and BP estimates, varied greatly over time and space (7% to >500%). The apparent uncoupling between respiration and PP has several implications for carbon export and storage in the western Arctic Ocean.

Effects of increased CO2 concentration on nutrient limited coastal summer plankton depend on temperature

Increasing seawater temperature and CO2 concentrations both are expected to increase coastal phytoplankton biomass and carbon to nutrient ratios in nutrient limited seasonally stratified summer conditions. This is because temperature enhances phytoplankton growth while grazing is suggested to be reduced during such bottom-up controlled situations. In addition, enhanced CO2 concentrations potentially favor phytoplankton species, that otherwise depend on costly carbon concentrating mechanisms (CCM). The trophic consequences for consumers under such conditions, however, remain little understood. We set out to experimentally explore the combined effects of increasing temperature and CO2 concentration for phytoplankton biomass and stoichiometry and the consequences for trophic transfer (here for copepods) on a natural nutrient limited Baltic Sea summer plankton community. The results show, that warming effects were translated to the next trophic level by switching the system from a bottom-up controlled to a mainly top-down controlled one. This was reflected in significantly down-grazed phytoplankton and increased zooplankton abundance in the warm temperature treatment (22.5°C). Additionally, at low temperature (16.5°C) rising CO2 concentrations significantly increased phytoplankton biomass. The latter effect however, was due to direct negative impact of CO2 on copepod nauplii which released phytoplankton from grazing in the cold but not in the warm treatments. Our results suggest that future seawater warming has the potential to switch trophic relations between phytoplankton and their grazers under nutrient limited conditions with the consequence of potentially disguising CO2 effects on coastal phytoplankton biomass.