Carbon and nitrogen stable isotope analysis of three types of oyster tissue in an impacted estuary

The stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) of the muscle, ctenidia and viscera of the Sydney rock oyster, Saccostrea glomerata, showed the dilution and assimilation of tertiary treated sewage along an estuarine gradient. The enriched ¹⁵N values of oyster ctenidia and viscera from within 50 m of the sewage outfall indicated the use of ¹⁵N-enriched tertiary treated sewage effluent (16 ± 2.3‰) as a nutrient source. The effect of sewage nitrogen on oyster δ¹⁵N was localised, with oysters 5 km upstream and downstream of the outfall not significantly enriched. Viscera δ¹⁵N was most sensitive to sewage nutrients and δ¹³C significantly defined an ocean-to-estuarine gradient. High variance in isotope ratios of viscera compromised its use as an indicator of anthropogenic nutrients, and this also reduced the utility of whole-body stable isotope ratios. Ctenidia was the most useful indicator tissue of sewage discharge at the scale of this study, being consistently and significantly enriched in δ¹⁵N close to the sewage outfall and δ¹³C clearly defined an estuarine gradient with less internal variability than viscera. Muscle δ¹⁵N was least sensitive to sewage effluent and showed the least variability, making it more suited to investigations of anthropogenic nutrient enrichment over larger spatio-temporal scales.     

Isotopic and elemental indicators of nutrient sources and status of coastal habitats in the Caribbean Sea,Yucatan Peninsula,Mexico

Nutrient inputs associated with coastal population growth threaten the integrity of coastal ecosystems around the globe. In order to assess the threat posed by rapid growth in tourism, we analyzed the nutrient concentrations as well as the δ¹⁵N of NO₃⁻ and macrophytes to detect wastewater nitrogen (N) at 6 locations along a groundwater-dominated coastal seagrass bed on the Caribbean coast of Mexico. We predicted that locations with greater coastal development would have higher concentrations of dissolved inorganic nitrogen (DIN) and phosphorus (P), as well as δ¹⁵N of NO₃⁻, reflecting wastewater sources of N. However, concentrations of NO₃⁻ were not significantly different between developed (3.3 ± 5.3 μM NO₃⁻) and undeveloped (1.1 ± 0.7 μM) marine embayments. The most important control on DIN concentration appeared to be mixing of fresh and salt water, with DIN concentrations negatively correlated with salinity. The δ¹⁵N of NO₃⁻ was elevated at an inland pond (7.0 ± 0.42‰) and a hydrologically-connected tide pool (7.6 ± 0.57‰) approximately 1 km downstream of the pond. The elevated δ¹⁵N of NO₃⁻ at the pond was paralleled by high δ¹⁵N values of Cladophora sp., a ubiquitous green alga (10 ± 1‰). We hypothesize that inputs of nitrogen rich (NO₃⁻ > 30 μM) groundwater, characterized by ¹⁵N enriched signatures, flow through localized submarine groundwater discharges (SGD) and contribute to the elevated δ¹⁵N signatures observed in many benthic macrophytes. However, changes in nitrogen concentrations and isotope values over the salinity gradient suggest that other processes (e.g. denitrification) could also be contributing to the ¹⁵N enrichments observed in primary producers. More measurements are needed to determine the relative importance of nitrogen transformation processes as a source of ¹⁵N to groundwaters; however, it is clear that continued inputs of anthropogenic N via SGD have the potential to severely impact ecologically and economically valuable seagrass meadows and coral reefs along the Caribbean coast of Mexico.     

Methane and methane carbon isotope ratios in the Northeast Atlantic including the Mid-Atlantic Ridge (50°N)

Previous work has shown that methane anomalies frequently occur within the rift valley of the Mid-Atlantic Ridge (MAR). The plumes appear confined within the high, steep walls of the valley, and it is not known whether methane may escape to the open ocean outside. In order to investigate this question, the concentration and ¹³C/¹²C ratio of methane together with CCl₃F concentration were measured in the northeastern Atlantic including the rift valley near 50°N. This segment contained methane plumes centered several 100 m above the valley floor with δ¹³C values mostly between –15‰ and –10‰. A limited number of helium isotope measurements showed that δ³He increased to 17% at the bottom of the valley, which suggests the helium and methane sources may be spatially separated. In the eastern Atlantic away from the ridge (48°N, 20°W), the methane concentration decreased monotonically from the surface to the bottom, but the methane δ¹³C exhibited a mid-water maximum of about –25‰. The bottom water methane contained a significantly lower δ¹³C of about –36‰. Thus, it appears that isotopically heavy methane escapes from the MAR into North Atlantic Deep Water (NADW) that contacts the ridge crest while circulating to the east. The formation of NADW supplies isotopically light methane that dilutes the input of heavy carbon from the ridge. We employed a time-dependent box model to calculate the extent of isotope dilution and thereby the flux of MAR methane into the NADW circulation. The degree of methane oxidation, which affects the ¹³C/¹²C of methane through kinetic isotope fractionation, was estimated by comparing methane and CFC-11 model results with observations. The model calculations indicate a MAR methane source of about 0.06×10⁻⁹ mol L⁻¹ yr⁻¹ to waters at the depth of the ridge crest. Assuming this extends to a 500 m thick layer over half of the entire Atlantic, the amount of methane escaping from the MAR to the open ocean is estimated to be about 1×10⁹ mol yr⁻¹. The total production of methane within the rift valley is likely much greater than the flux from the valley to the outside because of local oxidation. This implies that serpentinization of ultramafic rocks supports much of methane production in the rift valley because the amount expected from basalt degassing in association with mantle helium (<0.6×10⁹ mol CH₄ yr⁻¹) is less than even the net amount escaping from the valley. The model results also indicate the methane specific oxidation rate is about 0.05 yr⁻¹ in open waters of the northern Atlantic.     

Tracing seafloor methane emissions with benthic foraminifera: Results from the Ana submarine landslide (Eivissa Channel,Western Mediterranean Sea)

The hypothesis that benthic foraminifera are useful proxies of local methane emissions from the seafloor has been verified on sediment core KS16 from the headwall of the Ana submarine landslide in the Eivissa Channel, Western Mediterranean Sea. The core MS312 from a nearby location with no known methane emissions is utilised as control. The core was analysed for biostratigraphy, benthic foraminiferal assemblages, Hyalinea balthica and Uvigerina peregrina carbon and oxygen stable isotope composition, and sedimentary structures. The upper part of the core records post-landslide deglacial and Holocene normal marine hemipelagic sediments with highly abundant benthic foraminifera species that are typical of outer neritic to upper bathyal environment. In this interval, the δ¹³C composition of benthic foraminifera indicates normal marine environment analogous to those found in the control core. Below the sedimentary hiatus caused by the emplacement of the slide, the foraminiferal assemblages are characterised by lower density and higher Shannon Index. Markedly negative δ¹³C shifts in benthic foraminifera are attributed to the release of methane through the seabed. The mean values of the ¹³C anomaly in U. peregrina are ? 0.951 ± 0.208 in the pre-landslide sediments, and ? 0.269 ± 0.152 in post-slide reworked sediments deposited immediately above the hiatus. The δ¹³C anomaly in Hyalinea balthica is ? 2.497 ± 0.080 and ? 2.153 ± 0.087, respectively. To discard the diagenetic effects on the δ¹³C anomaly, which could have been induced by Ca–Mg replacement and authigenic carbonate overgrowth on foraminifera tests, a benthic foraminifera subsample has been treated following an oxidative and reductive cleaning protocol. The cleaning has resulted, only in some cases, in a slight reduction of the anomaly by 0.95% for δ¹³C and < 0.80% for δ¹⁸O. Therefore, the first conclusion is that the diagenetic alteration is minor and it does not alter significantly the overall carbon isotopic anomaly in the core. Consequently, the pre-landslide sediments have been subject to pervasive methane emissions during a time interval of several thousand years. Methane emissions continued during and immediately after the occurrence of Ana Slide at about 61.5 ka. Subsequently, methane emissions decreased and definitely ceased during the last deglaciation and the Holocene.     

Authigenic carbonates from methane seeps of the northern Congo fan: Microbial formation mechanism

Authigenic carbonates were collected from methane seeps at Hydrate Hole at 3113 m water depth and Diapir Field at 2417 m water depth on the northern Congo deep-sea fan during RV Meteor cruise M56. The carbonate samples analyzed here are nodules, mainly composed of aragonite and high-Mg calcite. Abundant putative microbial carbonate rods and associated pyrite framboids were recognized within the carbonate matrix. The δ¹³C values of the Hydrate Hole carbonates range from ?62.5‰ to ?46.3‰ PDB, while the δ¹³C values of the Diapir Field carbonate are somewhat higher, ranging from ?40.7‰ to ?30.7‰ PDB, indicating that methane is the predominant carbon source at both locations. Relative enrichment of ¹⁸O (δ¹⁸O values as high as 5.2‰ PDB) are probably related to localized destabilization of gas hydrate. The total content of rare earth elements (REE) of 5% HNO₃-treated solutions derived from carbonate samples varies from 1.6 ppm to 42.5 ppm. The shale-normalized REE patterns all display positive Ce anomalies (Ce/Ce* > 1.3), revealing that the carbonates precipitated under anoxic conditions. A sample from Hydrate Hole shows a concentric lamination, corresponding to fluctuations in δ¹³C values as well as trace elements contents. These fluctuations are presumed to reflect changes of seepage flux.     

The trophic significance of the invasive seaweed Sargassum muticum in sandy beaches

Native and exotic seaweeds frequently lie on the beach and sustain part of the benthic food web. However, the role of exotic seaweeds as food sources for beach consumers has been poorly studied. We studied the temporal and spatial variability in the trophic significance of the invasive brown seaweed Sargassum muticum on sandy beaches. We measured the stable isotopes (δ¹³C and δ¹⁵N) in the tissues of S. muticum and of invertebrate consumers and estimated the dietary biomass proportion of S. muticum during four sampling dates at two beaches and heights on the shore. Samples were collected from eight pitfall traps placed at a distance of 2 m from each other. Detrital macroalgae and seagrasses were also collected by hand within an area of 30 cm around each pitfall trap. We measured the spatial and temporal variability in the isotope composition of the beach consumers and of S. muticum using different models of analyses of variance. We then calculated the biomass proportion of S. muticum to the animal diet with a two-isotopic mixing model. The invasive alga S. muticum seemed to be one of the main food sources for the amphipod Talitrus saltator and, to a less extent, for the isopod Tylos europaeus. The importance of S. muticum was however temporally variable and decreased during spring (in March and May), probably due to the availability of native macrophytes. The supply of invasive wrack to beach food webs thus deserves more attention if we want to understand their role in influencing food web dynamics.     

Trophic relationships along a bathymetric gradient at the deep-sea observatory HAUSGARTEN

Deep-seafloor communities, especially those from the ice-covered Arctic, are subject to severe food limitation as the amount of particulate organic matter (POM) from the surface is attenuated with increasing depth. Here, we use naturally occurring stable isotope tracers (δ¹⁵N) to broaden our rudimentary knowledge of food web structure and the response of benthic organisms to decreasing food supplies along the bathymetric transect (～1300–5600 m water depth) of the deep-sea observatory HAUSGARTEN. Encompassing five trophic levels, the HAUSGARTEN food web is among the longest indicating continuous recycling of organic material typical of food-limited deep-sea ecosystems. The δ¹⁵N signatures ranged from 3.0‰ for Foraminifera to 21.4‰ (±0.4) for starfish (Poraniomorpha tumida). The majority of organisms occupied the second and third trophic level. Demersal fish fed at the third trophic level, consistent with results from stomach contents analysis. There were significant differences in the δ¹⁵N signatures of different functional groups with highest δ¹⁵N values in predators/scavengers (13.2±0.2‰) followed by suspension feeders (11.2±0.2‰) and deposit feeders (10.2±0.3‰). Depth (=increasing food limitation) affected functional groups in different ways. While the isotopic signatures of predators/scavengers did not change, those of suspension feeders increased with depth, and the reverse was found for deposit feeders. In contrast to the results of other studies, the δ¹⁵N signatures in POM samples obtained below 800 m did not vary significantly with depth indicating that changes in δ¹⁵N values are unlikely to be responsible for the depth-related δ¹⁵N signature changes observed for benthic consumers. However, the δ¹⁵N signatures of sediments decreased with increasing depth, which also explains the decrease found for deposit feeders. Suspension feeders may rely increasingly on particles trickling down the HAUSGARTEN slope and carrying higher δ¹⁵N signatures than the decreasing POM supplies, which elevates the δ¹⁵N value of their tissues. Our results imply that a depth-stratified approach should be taken to avoid a misinterpretation of data obtained at different depths.     

Fluid flow reconstruction in karstified Panormide platform limestones (north-central Sicily): Implications for hydrocarbon prospectivity in the Sicilian fold and thrust belt

Diagenetic analysis based on field and petrographic observations, isotope and microthermometric data was used to reconstruct the fluid flow history of the Cretaceous shallow water limestones from the Panormide platform exposed in north-central Sicily. Analysis focused on diagenetic products in cavities and dissolution enlarged fractures of the karstified limestones that occur just below a regional unconformity. The fluid flow history could be broken down into five stages that were linked to the kinematic and burial history of the region. (1) Petrography (zoned cathodoluminescence and speleothem textures) and stable isotopes (6.5 < δ¹⁸O_V-PDB < ?3.5‰ and 0 < δ¹³C_V-PDB < ?14‰) indicate that the earliest calcite phase was associated with karstification during emergence of the platform. Limestone dissolution at this stage is important with regard to possible reservoir creation in the Panormide palaeogeographic domain. (2) Fine-grained micrite sedimentation, dated as latest Cretaceous by nannopalaeontology and its ⁸⁷Sr/⁸⁶Sr isotope ratio (0.7078), marks replacement by marine fluids during subsequent submergence of the karstified platform. (3) The following calcite cement was still precipitated by marine-derived fluids (?7.0 < δ¹⁸O_V-PDB < ?5.0‰ and ?3.0 < δ¹³C_V-PDB < 0.5‰/T_m = ?2 to ?5 °C), but at increasingly higher temperatures (T_h = 60–120 °C). This has been interpreted as precipitation during Oligocene foredeep burial. (4) Hot (T_h = 130–180 °C), low saline (T_m < ?2.5 °C) fluids with increasingly higher calculated δ¹⁸O_SMOW signatures (+6 to +14‰) subsequently invaded the karst system. These fluids most likely migrated during fold and thrust belt development. The low salinity and relatively high δ¹⁸O_SMOW signatures of the fluids are interpreted to be the result of clay dewatering reactions. The presence of bitumen and associated fluorite with hydrocarbon inclusions at this stage in the paragenesis constrains the timing of oil migration in the region. (5) Finally, high saline fluids with elevated ⁸⁷Sr/⁸⁶Sr (0.7095–0.7105) signatures invaded the karst system. This last fluid flow event was possibly coeval with localized dolomitization and calcite cementation along high-angle faults of Pliocene age, as suggested by identical radiogenic signatures of these diagenetic products.     

Nitrogen isotopic composition of sinking particles from the southern Bay of Bengal: Evidence for variable nitrogen sources

An 8-year record of N fluxes and δ¹⁵N of sinking particles from the deep southern Bay of Bengal, northern Indian Ocean, is presented. Fluxes and δ¹⁵N vary between ∼0.1 and 3 mg m⁻² day⁻¹ and ∼2‰ and 8‰, respectively. The seasonal variation is determined mainly by oceanographic processes coupled to the Indian monsoon system. The annual pattern of δ¹⁵N is characterized by minima during spring intermonsoon (∼March–May), when nutrient inputs to the euphotic zone should be low because of stratification, and lighter nitrate/particulate matter is expected to be advected from the central Bay. Highest δ¹⁵N are associated with peak fluxes during southwest monsoon (∼June–September), when the southern Bay comes under the influence of the SW monsoon current, which appears to advect particulate matter with distinctly higher δ¹⁵N. The impact of this process, however, varies interannually under the influence of factors such as ENSO and the Indian Dipole Mode. Weakened advection leads to relatively low N fluxes and reduced δ¹⁵N. The data highlight the necessity of multi-annual studies to comprehend the natural variability of a system.     

Depth habitats and seasonal distributions of recent planktic foraminifers in the Canary Islands region (29°N) based on oxygen isotopes

Seasonal depth stratified plankton tows, sediment traps and core tops taken from the same stations along a transect at 29°N off NW Africa are used to describe the seasonal succession, the depth habitats and the oxygen isotope ratios (δ¹⁸O_shell) of five planktic foraminiferal species. Both the δ¹⁸O_shell and shell concentration profiles show variations in seasonal depth habitats of individual species. None of the species maintain a specific habitat depth exclusively within the surface mixed layer (SML), within the thermocline, or beneath the thermocline. Globigerinoides ruber (white) and (pink) occur with moderate abundance throughout the year along the transect, with highest abundances in the winter and summer/fall season, respectively. The average δ¹⁸O_shell of G. ruber (w) from surface sediments is similar to the δ¹⁸O_shell values measured from the sediment-trap samples during winter. However, the δ¹⁸O_shell of G. ruber (w) underestimates sea surface temperature (SST) by 2 °C in winter and by 4 °C during summer/fall indicating an extension of the calcification/depth habitat into colder thermocline waters. Globigerinoides ruber (p) continues to calcify below the SML as well, particularly in summer/fall when the chlorophyll maximum is found within the thermocline. Its vertical distribution results in δ¹⁸O_shell values that underestimate SST by 2 °C. Shell fluxes of Globigerina bulloides are highest in summer/fall, where it lives and calcifies in association with the deep chlorophyll maximum found within the thermocline. Pulleniatina obliquiloculata and Globorotalia truncatulinoides, dwelling and calcifying a part of their lives in the winter SML, record winter thermocline (～180 m) and deep surface water (～350 m) temperatures, respectively. Our observations define the seasonal and vertical distribution of multiple species of foraminifera and the acquisition of their δ¹⁸O_shell.     

Nitrogen isotope patterns in the oxygen-deficient waters of the Eastern Tropical North Pacific Ocean

We analyzed the stable nitrogen isotope composition of an extensive set of samples of particulate matter (PM) and seawater nitrate collected during October/November 1997 along the Mexican coastline from 24° to 11.5°N. At the northern and southern end of our study area, the δ¹⁵N of PM ranged between 5 and 7‰ in the upper 200 m of the water column with higher values at intermediate depths. These data are very similar to those reported from other parts of the open ocean. In the oxygen minimum zone (OMZ), we found significantly higher δ¹⁵N values for suspended particles. Furthermore, the δ¹⁵N of nitrate (NO₃) was elevated within the OMZ and we found a strong relationship between the oxygen concentration, nitrate deficit and the ¹⁵N content of the nitrate. The core of the OMZ between 22°N, 105°W and 15°N, 110°W coincided with higher nitrate deficits and δ¹⁵N values relative to the stations near the boundaries. The δ¹⁵N of nitrate was highest, with values up to 18.7‰, where oxygen concentrations were below 1–2 μmol/l. This pattern is consistent with an overall nitrogen isotopic enrichment factor of 22.5‰ for denitrification in the core of the OMZ using the Rayleigh equation (closed-system approach). Results from a diffusion model (open-system approach), however, gave a fractionation factor of 30±7.5‰, implying that the Rayleigh formula only gives a lower estimate of the fractionation factor ε. The vertical flux of particles collected in short-term deployments (ca. 35 h) of a drifting sediment trap was not significantly correlated with the water column nitrate deficit. The isotopic signature of the nitrate within the gradient is very similar to the δ¹⁵N value of sedimenting particles, suggesting that there might be a strong link between the production and sedimentation of particles. Upward flux of nitrate across the thermocline can account for less than half of the particle flux leaving the mixed layer. Mixing and transport of nitrate across the lower boundary of the OMZ can lead to significant enrichment in the ¹⁵N content of deep waters, and our isotopic data imply that at least 14% of the nitrate in the waters below the OMZ originates from this source.     

Carbon and nitrogen dynamics of the intertidal seagrass, <Emphasis Type="Italic">Zostera japonica</Emphasis>, on the southern coast of the Korean peninsula

Seagrasses require a large amount of nutrient assimilation to support high levels of production, and thus nutrient limitation for growth often occurs in seagrass habitats. Seagrasses can take up nutrients from both the water column and sediments. However, since seagrasses inhabiting in the intertidal zones are exposed to the air during low tide, the intertidal species may exhibit significantly different carbon (C) and nitrogen (N) dynamics compared to the subtidal species. To examine C and N dynamics of the intertidal seagrass, Zostera japonica, C and N content and stable isotope ratios of above- and below-ground tissues were measured monthly at the three intertidal zones in Koje Bay on the southern coast of Korea. The C and N content and stable isotope (δ¹³C and δ¹⁵N) ratios of seagrass tissues exhibited significant seasonal variations. Both leaf and rhizome C content were not significantly correlated with productivity. Leaf δ¹³C values usually exhibited negative correlations with leaf productivity. These results of tissue C content and δ¹³C values suggest that photosynthesis of Z. japonica in the study site was not limited by inorganic C supply, and sufficient inorganic C was provided from the atmosphere. The tissue N content usually exhibited negative correlations with leaf productivity except at the upper intertidal zone, suggesting that Z. japonica growth was probably limited by N availability during high growing season. In the upper intertidal zone, no correlations between leaf productivity and tissue elemental content and stable isotope ratios were observed due to the severely suppressed growth caused by strong desiccation stress.     

Seasonal oxygen and carbon isotope variability in euthecosomatous pteropods from the Sargasso Sea

We examine seasonal variations in the stable carbon and oxygen isotopic composition of individual shells of the pteropods Limacina inflata and Styliola subula, collected from Oceanic Flux Program sediment traps (at 500 m depth) near Bermuda in the western Sargasso Sea. Calcification depths estimated from L. inflata δ¹⁸O vary between 200 and 650 m in late winter and spring, and between 50 and 250 m in late summer and fall. S. subula shows similar seasonal variability with calcification depths between 250 and 600 m in late winter and spring and 50–400 m in late summer and fall. These results suggest that both species calcify across a greater range of depths than indicated by previous geochemical studies. Furthermore, the data indicate that these species change their calcification depth in conjunction with changes in thermal stratification of the water column. Pteropod shell δ¹³C values vary inversely with δ¹³C_DIC but show a positive correlation with seawater [CO₃²⁻] and temperature after depth differences in δ¹³C_DIC are accounted for. We hypothesize that either the influence of temperature on metabolic CO₂ incorporation during shell growth and/or the influence of ambient [CO₃²⁻] on shell geochemistry can explain these relationships. Taken together, the individual shell δ¹⁸O and δ¹³C data suggest that shell calcification, and by inference the life cycle, of these pteropods is several months or less. Individual pteropod shell analyses have potential for contributing to our understanding of the environmental parameters that play a role in seasonal calcification depth shifts, as well as to our knowledge of past upper ocean thermal structure.     

Fish farming enhances biomass and nutrient loss in Posidonia oceanica (L.) Delile

Fish farming impact on the seasonal biomass, carbon and nutrient (nitrogen and phosphorus) balance of the endemic Mediterranean seagrass Posidonia oceanica was assessed in the Aegean Sea (Greece) in order to detect changes in magnitude and fate of seagrass production and nutrient incorporation with organic loading of the meadows. Phosphorus concentration in the leaves, rhizomes and roots was enhanced under the cages throughout the study. Standing biomass was diminished by 64% and carbon, nitrogen and phosphorus standing stock by 64%, 61% and 48%, respectively, under the cages in relation to those at the control. Seagrass production decreased by 68% and element (C, N, P) incorporation by 67%, 58% and 58%, respectively, under the cages. Leaf shedding was reduced by 81% and loss of elements (C, N, and P) through shedding by 82%, 74% and 72%, respectively, under the cages. Leaf and element (C, N, P) residual loss rate, accounting for grazing and mechanical breakage of leaves, was decreased by 79%, 85%, 100% and 96%, respectively, at the control station. At the control station, 13.98 g C m^?2 yr^?1, 1.91 g N m^?2 yr^?1 and 0.05 g P m^?2 yr^?1 were produced in excess of export and loss. In contrast, under the cages 12.69 g C m^?2 yr^?1, 0.31 g N m^?2 yr^?1 and 0.04 g P m^?2 yr^?1 were released from the meadow. Organic loading due to fish farm discharges transformed the seagrass meadow under the cages from a typical sink to a source of organic carbon and nutrients.     

Nutrient retention in the sediments and the submerged aquatic vegetation of the coastal lagoon of the Ria de Aveiro,Portugal

A decrease in the areas covered by seagrasses within the Ria de Aveiro, Portugal, has been observed over the past five decades, resulting in a corresponding increase of the areas of uncovered sediment supporting the growth of sparse macroalgae populations only. Presently, several macroalgae (Ulva spp., Gracilaria sp.) and one seagrass species (Nanozostera noltii (Hornem.) Toml. & Posl.) comprise the submerged aquatic vegetation (SAV) adapted to this shallow, high-energy environment, characterised by fast tidal currents and turbid waters and in which large areas of the bed are exposed during low tide. This study shows that there is a strong inter-relation between the SAV and the surface sediment in intertidal areas. The sediment covered by N. noltii was finer (median grain size 95 µm) and had a high percentage of organic matter (mean value 7.6%), compared with the sediment colonised by macroalgae (median grain size 239 µm; mean organic content 3.2%). The concentrations of both total nitrogen and phosphorous were significantly greater (P < 0.001) in surface sediments covered by N. noltii. Thus, sediments within N. noltii appear to act as a large reservoir of N and P by accumulating greater concentrations of fine sediment particles (silt and clay) and organic matter when compared with the coarser sediment covered with macroalgae only. Hence, the reduction in the area covered by seagrasses will likely result in a gradual loss of nutrients and fine sediment from the Ria de Aveiro channels.     

Distribution and burial of organic carbon in sediments from the Indian Ocean upwelling region off Java and Sumatra,Indonesia

Sediments were sampled and oxygen profiles of the water column were determined in the Indian Ocean off west and south Indonesia in order to obtain information on the production, transformation, and accumulation of organic matter (OM). The stable carbon isotope composition (δ¹³C_org) in combination with C/N ratios depicts the almost exclusively marine origin of sedimentary organic matter in the entire study area. Maximum concentrations of organic carbon (C_org) and nitrogen (N) of 3.0% and 0.31%, respectively, were observed in the northern Mentawai Basin and in the Savu and Lombok basins. Minimum δ¹⁵N values of 3.7‰ were measured in the northern Mentawai Basin, whereas they varied around 5.4‰ at stations outside this region. Minimum bottom water oxygen concentrations of 1.1 mL L^?1, corresponding to an oxygen saturation of 16.1%, indicate reduced ventilation of bottom water in the northern Mentawai Basin. This low bottom water oxygen reduces organic matter decomposition, which is demonstrated by the almost unaltered isotopic composition of nitrogen during early diagenesis. Maximum C_org accumulation rates (CARs) were measured in the Lombok (10.4 g C m^?2 yr^?1) and northern Mentawai basins (5.2 g C m^?2 yr^?1). Upwelling-induced high productivity is responsible for the high CAR off East Java, Lombok, and Savu Basins, while a better OM preservation caused by reduced ventilation contributes to the high CAR observed in the northern Mentawai Basin. The interplay between primary production, remineralisation, and organic carbon burial determines the regional heterogeneity. CAR in the Indian Ocean upwelling region off Indonesia is lower than in the Peru and Chile upwellings, but in the same order of magnitude as in the Arabian Sea, the Benguela, and Gulf of California upwellings, and corresponds to 0.1–7.1% of the global ocean carbon burial. This demonstrates the relevance of the Indian Ocean margin off Indonesia for the global OM burial.     

Stable isotope constraints on the nitrogen cycle of the Mediterranean Sea water column

We used the nitrogen isotope ratio of algae, suspended particles and nitrate in the water column to track spatial variations in the marine nitrogen cycle in the Mediterranean Sea. Surface PON (5–74 m) was more depleted in ¹⁵N in the eastern basin (−0.3±0.5‰) than in the western basin (+2.4±1.4‰), suggesting that nitrogen supplied by biological N₂ fixation may be an important source of new nitrogen in the eastern basin, where preformed nitrate from the Atlantic Ocean could have been depleted during its transit eastward. The δ¹⁵N of nitrate in the deep Mediterranean (∼3‰ in the western-most Mediterranean and decreasing toward the east) is significantly lower than nitrate at similar depths from the North Atlantic (4.8–5‰), also suggesting an important role for N₂ fixation. The eastward decrease in the δ¹⁵N of surface PON is greater than the eastward decrease in the δ¹⁵N of the subsurface nitrate, implying that the amount of N₂ fixation in the eastern Mediterranean is great enough to cause a major divergence in the δ¹⁵N of phytoplankton biomass from the δ¹⁵N of the nitrate upwelled from below. Variations in productivity associated with frontal processes, including shoaling of the nitracline, did not lead to detectable variations in the δ¹⁵N of PON. This indicates that no differential fertilization or productivity gradient occurred in the Almerian/Oran area. Our results are consistent with a lack of gradient in chlorophyll-a (chl-a) and nitrate concentration in the Alboran Sea. ¹⁵N enrichment in particles below 500 m depth was detected in the Alboran Sea with respect to surface PON, reaching an average value of +7.4±0.7‰. The δ¹⁵N in sinking particles caught at 100 m depth (4.9–5.6‰) was intermediate between suspended surface and suspended deep particles. We found a consistent difference in the isotopic composition of nitrogen in PON compared with that of chlorophyll (Δδ¹⁵N[PON-chlorin]=+6.4±1.4‰) in the surface, similar to the offset reported earlier in cultures for cellular N and chl-a. This indicates that δ¹⁵N of phytoplankton biomass was retained in surface PON, and that alteration of the isotopic signal of PON at depth was due to heterotrophic activity.     

Trophic relationships in deep-water decapods of Le Danois bank (Cantabrian Sea,NE Atlantic): Trends related with depth and seasonal changes in food quality and availability

The trophic relationships of decapod crustaceans on Le Danois bank (NE of Iberian Peninsula, NE Atlantic Ocean) were studied within the framework of the multidisciplinary project ECOMARG during two surveys, one in October 2003 and the other in April 2004. The diets of eleven species of decapods were analyzed and, within a rather continuous gradient of food source exploitation, 3 trophic groups were identified: (1) plankton feeders, comprising the shrimps Acanthephyra pelagica, Sergia robusta, and Pasiphaea tarda, which preyed on meso-bathypelagic taxa such as euphausiids and calanoids; (2) benthos feeders, comprising the crangonids Pontophilus norvegicus and Pontophilus spinosus, the crab Geryon trispinosus and the shrimp Aristeus antennatus; and (3) an intermediate group, including the rest of species, with mixed diets that included detritus. Among the third group, anomurans (Munida tenuimana, Pagurus alatus, and Parapagurus pilosimanus) consumed phytoplanktonic detritus in April, suggesting a link with peaks of surface Chl a occurring between March and April in the study area. Gut pigment and isotopic (δ¹³C/δ¹⁵N correlations) analyses revealed that assemblages inhabiting the top of the bank (455–612 m) and the inner basin (642–1048 m, close to the Lastres canyon head) had different food sources, with species inhabiting the deepest region exhibiting a stronger dependence on marine snow derivatives. These results are consistent with the higher proportion of mud and sediment organic matter (OM) content in the inner basin (82.2% pellites; 6.3% OM at 1028 m) compared to the top of the Le Danois bank (only 13.9% pellites; 2.8% OM at 485 m), which is a hydrodynamically more active zone. Exploitation of different food sources is also consistent with differences in the trophic level of species, inferred from stable δ¹⁵N isotope analyses, which yield values ranging from 6.88‰ for the hermit crab P. alatus to 13.52‰ for the crangonid shrimp P. norvegicus. Stomach fullness was higher in April 2004 than in October 2003, both between and within species of the dominant decapods, including detritus feeders (M. tenuimana) and benthos feeders (e.g. G. trispinosus, P. norvegicus). Most species exhibited a parallel increase in their density in April 2004, with a significant positive correlation between density and stomach fullness. This increase coincides with a peak of surface Chl a concentration occurring in March–April.     

Seasonal influences on size-fractionated chlorophyll a concentrations and primary production in the north-west Indian Ocean

Chlorophyll a (chl a) concentrations and primary production by the 0.2–2, 2–18 and >18 μm phytoplankton size-fractions were estimated along a transect in the NW Indian Ocean extending from the coast of Oman to 8°N 68°E during the late SW monsoon and autumn intermonsoonal seasons in 1994. Primary production was estimated using the ¹⁴C technique with either in situ or simulated in situ incubations. During the late monsoon season, maximal chl a and production values were recorded in the coastal upwelling zone with values of 69 mg m^-2 and 3800 mg C m^-2 d^-1, respectively. The maxima, which were distributed patchily in this region, were dominated by the >18 μm size-fraction. Over the remainder of the transect chl a concentrations and production averaged 30 mg m^-2 and 1500 mg C m^-2 d^-1, respectively, with approximately equal contributions by the three size-fractions in the case of chl a at the majority of stations, but in general, with a maximum in production in the 0.2–2 μm fraction. Immediately following cessation of the SW monsoon wind, chl a and production values over the northern part of the transect decreased to values similar to those over the southern part of the transect at the time of the SW monsoon, with the contributions by the three size-fractions being approximately equal. During the following intermonsoonal season, both chl a concentrations and production across the section were dominated by the 0.2–2 μm size-fraction, with average chl a and production values of the order of 20 mg m^-2 and 750 mg C m^-2 d^-1, respectively. Considerable variation in production values, however, was exhibited across the transect. A clearly defined subsurface chl a maximum was only recorded at the southernmost stations of the transect in oligotrophic waters: the feature did not develop universally across the transect during the intermonsoon.     

Compositional and fluorescence characteristics of the giant diatom Ethmodiscus along a 3000 km transect (28°N) in the central North Pacific gyre

The giant diatom Ethmodiscus was examined along an east–west transect at 28–30°N during 2002 and 2003 to determine if abundance, chemical composition or physiological status of this largest of diatoms varied on the scale of 100's–1000's of km in North Pacific gyre. Abundance ranged from <0.1–>2.0 cells m⁻³ and supported the notion of an abundance mosaic reported previously. However, there was only minimal support for the relationship between abundance and nutrient concentration at 125 m reported previously. Cellular chlorophyll varied little along the transect (7.3–10.9 ng chl cell⁻¹) except at the westernmost station. Cellular N and P quotas co-varied 3–4.5 fold (mean=50.8±3.7 and 3.7±0.8 nmol N and P cell⁻¹) and yielded N:P ratios that closely clustered around the Redfield ratio (average=14.6±1.1). Only low levels of chlorophyll-normalized alkaline phosphatase (APase) activity were observed (0.4–2.5 nmol P μg chl⁻¹ h⁻¹) with APase activity lower than that in either the bulk water, or co-occurring Trichodesmium spp. and Pyrocystis noctiluca. The active fluorescence parameter F_v:F_m, a property sensitive to Fe stress, was uniformly high at all stations (average=0.73±0.04 for 2003, and 0.69±0.05 for 2002), indicating sufficient Fe for optimum photosynthetic competence. These results contrasted sharply with results from Rhizosolenia mats reported along the same transect where there was a significant decline westward in F_v:F_m. Both ferredoxin (Fd) and flavodoxin accumulated in cells of Ethmodiscus, resulting in Fd Index values of<0.6. Iron cell quotas ranged from 0.7–5.1 pmol Fe cell⁻¹. When normalized to cytoplasmic volume, the Fe μm⁻³ was comparable to that of Escherichia coli. We note that the disproportionate contribution of the vacuole (with its high organic content) to total volume typical of large diatoms is a potentially significant source of error in Fe:C ratios and suggest that Fe should be normalized to cytoplasmic volume whenever possible to permit valid intercomparisons between studies. The composition, F_v:F_m data and Fe:C ratio suggest a relatively uniform population experiencing little N, P or Fe stress. The uncoupling of the Fd Index from these measures is consistent with previous findings showing that the expression of flavodoxin can be characterized as an early stress response and that its accumulation is not necessarily correlated with physiological deficit. Ethmodiscus appears to be well adapted to some of the most oligotrophic waters in the ocean. Because it is an important sedimentary marker, the biology of living Ethmodiscus provides insights into the source of extensive Ethmodiscus oozes. Mass sedimentation after frontal accumulation has been suggested as a source for these oozes. Our data contain no evidence that the flux is linked directly to Fe, N or P stress.