Depth-dependence in stable isotope ratio δ15N of benthic POM consumers: The role of particle dynamics and organism trophic guild

The stable nitrogen isotope ratio (δ¹⁵N) is an established indicator of trophic hierarchy in marine food-web studies. Most of these studies presume that spatial variation in the primary food source is negligible, although a water-depth-related increase in δ¹⁵N of particulate organic matter (POM) has been found in many systems. We used the high-Antarctic Weddell Sea shelf and slope ecosystem to test whether such a depth-related change in δ¹⁵N is reflected at higher trophic levels, i.e., benthic consumers of POM. In suspension feeders (SF) we found a significant increase in δ¹⁵N with water depth of up to 9.8‰, whereas in deposit feeders (DF) a depth effect was barely detectable. Particle-size preferences of the two feeding guilds combined with particle-size-dependent sinking velocities and biogeochemical reworking of POM are discussed as the major causes of these differences. It is essential to marine food-web studies to take into account the general depth effect on POM δ¹⁵N as well as potential feeding-guild-specific differences in the response of POM consumer tissue δ¹⁵N to avoid serious bias and misinterpretation of stable-isotope-based trophic information.     

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.     

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.     

Food web structure of two Mediterranean lagoons under varying degree of eutrophication

The food web structure and functioning of two north-western Mediterranean lagoons exhibiting contrasting degrees of eutrophication and marine influences were compared through δ¹³C and δ¹⁵N analysis of major potential food sources and consumers. The Lapalme Lagoon is well preserved and has kept a natural and temporary connection with the open sea. Conversely, the Canet Lagoon is heavily eutrophicated and its water exchange with the open sea has been artificially reduced. In Lapalme, all potential food sources and consumers exhibited δ¹⁵N values indicative of pristine coastal areas. Suspended particulate organic matter (POM) and sediment organic matter (SOM) pools seemed to constitute the main food sources of most primary consumers. Both primary producers and all consumers were much more ¹⁵N-enriched (by ∼ 10‰) and more ¹³C-depleted in Canet than in Lapalme. This reflected: (1) the assimilation of important amounts of anthropogenic nitrogen in the food web, and (2) a marked and uniform influence of ¹³C-depleted allochtonous sources of carbon. Based on the mean δ¹⁵N of primary consumers, we found rather similar food web lengths in both lagoons with top consumers at trophic levels 3.6 and 4.0 in Canet and Lapalme, respectively. However, the eutrophication of the Canet Lagoon resulted in a simplification of the food web structure (i.e., a single trophic pathway from a ¹⁵N-enriched fraction of the SOM pool to top predators) compared to what was observed in Lapalme Lagoon where additional ¹³C-enriched food sources played a significant trophic role. Moreover, some consumers of Canet tended to exploit primary producers to a larger extent (and thus to exhibit lower trophic levels) than in Lapalme.     

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.     

Food-web structure and trophodynamics of mesopelagic–suprabenthic bathyal macrofauna of the Algerian Basin based on stable isotopes of carbon and nitrogen

The trophodynamics of mesopelagic (macrozooplankton/micronekton) and benthic boundary layer (suprabenthos=hyperbenthos) faunas from the Algerian Basin were characterized on a seasonal scale through stable carbon and nitrogen isotopic analyses of a total of 34 species and two broad taxa (Copepoda and Cumacea). This is the first study simultaneously focused on trophodynamics of deep-sea zooplankton and suprabenthos. Samples were collected southeast of Mallorca (Algerian Basin, Western Mediterranean), on the continental slope close to Cabrera Archipelago, at 650–780 m depths, ca. bi-monthly between August 2003 and June 2004. Mean δ¹³C values of suprabenthos ranged from ?21.1‰ (Munnopsurus atlanticus) to ?16.7‰ (Cyclaspis longicaudata). Values of δ¹⁵N ranged from 2.8‰ (Lepechinella manco) to 9.9‰ (larvae of Gnathia sp.). The stable isotope ratios of suprabenthic fauna displayed a continuum of values, confirming a wide spectrum of feeding guilds (from filter feeders/surface deposit feeders to predators). According to the available information on diets for suprabenthic species, the highest annual mean δ¹⁵N values were found for the hematophagous isopod Gnathia sp. parasite on fish (represented by Praniza larvae) and carnivorous amphipods (e.g. Rhachotropis spp., Nicippe tumida) consuming copepods, and the lowest δ¹⁵N values were found for two cumaceans (Cyclaspis longicaudata and Platysympus typicus) feeding on detritus. Assuming a ¹⁵N-enrichment factor of 2.5‰ and deposit feeders as baseline, we found three trophic levels in suprabenthic food webs. δ¹³C ranges were particularly wide among deposit feeders (ranging from ?21.8% to ?17.3‰) and omnivores (from ?20.5% to ?18.8‰), suggesting exploitation of particulate organic matter (POM) of different characteristics. Our isotopic analyses revealed lower ranges of δ¹³C and δ¹⁵N for macrozooplankton/micronekton, compared with suprabenthos. δ¹³C values of zooplankton taxa ranged from ?21.1‰ (the hyperiid Phrosina semilunata) to ?19.9‰ (the decapod Pasiphaea multidentata), while δ¹⁵N values ranged from 3.9‰ (P. semilunata) to 7.5‰ (P. multidentata). Among zooplankton, more enriched δ¹⁵N values were found among carnivores (e.g. the fish Cyclothone spp. and Pasiphaea multidentata) preying on copepods, hyperiids, euphausiids and small fish. The lowest δ¹⁵N values were found for hyperiids that feed on the mucus nets of salps (e.g. Vibilia armata). After contrasting isotope analysis with dietary data, we conclude there were two trophic levels among zooplankton/micronekton. Strong correlation between the mean annual δ¹⁵N and δ¹³C values was found for zooplankton (R²=0.7), but not for suprabenthos, which suggests a single source of carbon for plankton. We found a general seasonal trend for δ¹³C enrichment from late autumn (November) to late winter–spring (February–April) for both suprabenthos and zooplankton. The δ¹³C enrichment in February–April was correlated in zooplankton with higher surface chlorophyll a concentration 1 month before sampling. As evidenced by δ¹³C–δ¹⁵N correlations, the response of zooplankton to the peak of surface primary production was almost immediate (an increase of δ¹³C–δ¹⁵N correlations in February), and stronger than for suprabenthos. The response among suprabenthos was weak, with slight increase in δ¹³C–δ¹⁵N relationships in April–June.     

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.     

Pelagic nitrogen dynamics in the Vietnamese upwelling area according to stable nitrogen and carbon isotope data

Upwelling and nitrogen (N) fixation provide new N for primary production off southern central Vietnam. Here we evaluate the roles of both N sources for zooplankton nutrition by comparing δ¹⁵N and δ¹³C values in nitrate, particulate organic matter (POM), and six net-plankton size fractions from monsoon and intermonsoon seasons. The δ¹³C values in POM and the net-plankton size fractions differed by 2–4‰ at any time. We assume that plankton from the POM filters was dominated by nano-and picoplankton as opposed to micro- and mesoplankton in the net-samples. The implications of this are discussed in terms of size differential pathways of C and N in the planktonic food web. We used δ¹⁵N to estimate the differences in N nutrition between the actual upwelling region and the oligotrophic area further offshore. The δ¹⁵N values of the net-plankton size fractions were depleted in δ¹⁵N by ca. 2‰ outside compared to inside the upwelling area during the monsoon season. We attribute these patterns to the additional utilization of N derived from N fixation. The concomitant findings of high N fixation rates reported earlier and low subthermocline nitrate (nitrate_sub) values of 2.9–3.6‰ support this conclusion. Net-plankton δ¹⁵N values increased with size, pointing to the dominance of higher trophic levels in the larger size fractions. According to a two source mixing model N fixation may have provided up to 13% of the N demand in higher trophic levels.     

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.     

Trophic ecology and vertical patterns of carbon and nitrogen stable isotopes in zooplankton from oxygen minimum zone regions

The unique physical and biogeochemical characteristics of oxygen minimum zones (OMZs) influence plankton ecology, including zooplankton trophic webs. Using carbon and nitrogen stable isotopes, this study examined zooplankton trophic webs in the Eastern Tropical North Pacific (ETNP) OMZ. δ¹³C values were used to indicate zooplankton food sources, and δ¹⁵N values were used to indicate zooplankton trophic position and nitrogen cycle pathways. Vertically stratified MOCNESS net tows collected zooplankton from 0 to 1000 m at two stations along a north-south transect in the ETNP during 2007 and 2008, the Tehuantepec Bowl and the Costa Rica Dome. Zooplankton samples were separated into four size fractions for stable isotope analyses. Particulate organic matter (POM), assumed to represent a primary food source for zooplankton, was collected with McLane large volume in situ pumps.The isotopic composition and trophic ecology of the ETNP zooplankton community had distinct spatial and vertical patterns influenced by OMZ structure. The most pronounced vertical isotope gradients occurred near the upper and lower OMZ oxyclines. Material with lower δ¹³C values was apparently produced in the upper oxycline, possibly by chemoautotrophic microbes, and was subsequently consumed by zooplankton. Between-station differences in δ¹⁵N values suggested that different nitrogen cycle processes were dominant at the two locations, which influenced the isotopic characteristics of the zooplankton community. A strong depth gradient in zooplankton δ¹⁵N values in the lower oxycline suggested an increase in trophic cycling just below the core of the OMZ. Shallow POM (0–110 m) was likely the most important food source for mixed layer, upper oxycline, and OMZ core zooplankton, while deep POM was an important food source for most lower oxycline zooplankton (except for samples dominated by the seasonally migrating copepod Eucalanus inermis). There was no consistent isotopic progression among the four zooplankton size classes for these bulk mixed assemblage samples, implying overlapping trophic webs within the total size range considered.     

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.     

Shell architecture,element composition,and stable isotope signature of the giant deep-sea oyster Neopycnodonte zibrowii sp. n. from the NE Atlantic

A conspicuous new deep-sea oyster, Neopycnodonte zibrowii sp. n., is described from the Azores Archipelago, where it thrives in 420 to >500 m water depth in high densities concealed underneath overhangs. The new species reaches a relatively large size, which may exceed 20 cm, and is characterised by a very unusual hinge line morphology, straight without a bulge of the resilium. It is compared to the extant Indo-Pacific Empressostrea kostini Huber and Lorenz, 2007 and to the cosmopolitan Neopycnodonte cochlear (Poli, 1791), which has a broadly sympatric distribution at shallower depths in the Azores and Bay of Biscay. Radiocarbon dating reveals that individuals reach an impressive lifespan of one to more than five centuries, placing them among the longest-lived molluscs known to date. They often grow on top of each other, forming stacks that resemble dish piles—an effective measure to optimise shell stability with minimal biomineralisation effort, but with the drawback of increased bioerosion ultimately leading to detachment.Three microstructure types are developed in N. zibrowii: (1) the cross-foliated, calcitic, dorsal to central endostracum and aragonitic ligostracum, (2) the porous vesicular structure of the calcitic ventral endostracum, and (3) the simple prismatic aragonitic myostracum. Foliated and vesicular shell portions show sub-millimetre-scale first-order increments delineated by conchiolin-rich growth breaks (interpreted as reproductive cyclicity), and less distinct second-order increments (interpreted as annual in nature). This pattern is clearly reflected by the elemental composition with the primary growth breaks lacking Ca and Sr but including Mg and S as organic matrix constituents. The second-order increments within the calcite are mirrored by moderately co-varying Mg/Ca and S/Ca fluctuations at stable Sr concentrations, reflecting varying proportions of organic matrix. Dorsal and central endostracum transects reveal a low inter-valve, but considerable inter-specimen variability with high Mg/Ca molar ratios and fluctuations (22.5±17.6 mmol/mol), low Sr/Ca values (0.2±0.1 mmol/mol), and a typical to high S/Ca content (6.9±2.2 mmol/mol), when compared to other calcitic bivalves.Unlike short-lived, shallow-water oysters, N. zibrowii thrives under very stable environmental conditions. Minimal temperature fluctuations and stable open marine salinity provide an optimal basis to recognise biological fractionation processes. Strong Mg/Ca fluctuations indicate a physiological control related to metabolism and biomineralisation, prohibiting the use of this ratio as a temperature or ocean chemistry proxy. Low Sr/Ca ratios indicate rather constant and low long-term accretion rates, while short-scale fluctuations may be attributed to short-term variations in growth rate and Mg incorporation. Oxygen isotopes yield a considerable spread of 1.8‰ with a mean of 2.0±0.3‰ δ¹⁸O V-PDB, and low correlation between different contemporaneous parts of the shell and between specimens. These values surprisingly exceed expected equilibrium conditions, calculated from in situ temperature data (annual mean 12.3±0.3 °C) and seawater isotopic composition (0.5±0.1‰ δ¹⁸O SMOW), by 0.5‰ on average. Such positively shifting vital effects, previously reported for limpets and barnacles, are often overlooked in high-temperature and high-amplitude settings and may be more common than is currently believed. Carbon isotopes range from 0.2 to 3.5‰ δ¹³C V-PDB (mean 1.8±0.7‰) and show an ontogenetic decrease, but may incorporate an environmental signal in adult portions, indicated by a strong correspondence of peaks between specimens. This signal is likely driven by a complex interplay of different factors, such as primary production, current-based food supply and metabolism.     

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.     

基于稳定性同位素的普里兹湾南极大磷虾不同发育阶段食性转变分析

As one of the most common and dominant species in the Southern Ocean, Antarctic krill(Euphausia superba)play a significant role in food web structure and the process of energy flow. The diet of Antarctic krill in the Prydz Bay during austral summer of 2012/2013 was investigated and the ontogenetic shift in krill diet was evaluated using the stable isotope method. The nitrogen stable isotope values(δ~(15) N) of adults((2.78±0.58)‰) were much higher than those of juveniles((1.69±0.70)‰), whereas the carbon stable isotope values(δ~(13) C) of adults(–(28.26±1.08)‰) were slightly lower than those of juveniles(–(27.48±1.35)‰). Particulate organic matter(POM)from 0, 25, and 50 m depth combined(0/25/50 m) represented phytoplankton food items. The results showed that phytoplankton food items in surface water and mesozooplankton were two essential food items for Antarctic krill in the Prydz Bay during summer. POM(0/25/50 m) contributes 56%–69% and 26%–34% to the diet of juvenile and adult krill, respectively, whereas mesozooplankton composes 13%–34% and 58%–71% of the diet of juvenile and adult krill, respectively. Thus, an ontogenetic diet shift from POM(0/25/50 m), which consists mainly of phytoplankton, to a higher trophic level diet containing mesozooplankton, was detected. The capacity for adults to consume more zooplankton food items may minimize their food competition with juveniles, which rely mostly on phytoplankton food items. This suggests "diet shift with ontogeny" which may somehow help krill keep their dietary energy budget balanced and well adapted to the Antarctic marine ecosystem as a dominant species.     

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.     

Elemental and stable isotope composition of Zostera noltii (Horneman) leaves during the early phases of decay in a temperate mesotidal lagoon

The aim of this study was to assess the changes in the elemental and stable isotope composition of Zostera noltii leaves during the early phases of decay. The litter biomass, the C, N and S elemental contents, the C:N ratios and the δ¹³C, δ¹⁵N and δ³⁴S values were monitored in a litterbag experiment for 60 days at Ria Formosa lagoon, southern Portugal in two different field conditions, permanently submerged in a salt marsh pond and at high intertidal. The elemental and stable isotope composition of both attached leaves and leaf wrack of Z. noltii was also determined. The overall decay rate of Z. noltii leaf material throughout the whole experimental period was k = 0.016 ± 0.001 d⁻¹. Even though the biomass loss of the Z. noltii leaves after 30 days was about 25% higher within the pond than at the high intertidal, at the end of the experiment there were no significant differences between sites. The decay of the C and N content was of the same magnitude and thus the C:N molar ratio did not vary significantly in the experiment. The elemental content of the leaf wrack was in the range of the element content of leaf litter after 30–60 days of incubation. Contrary to other vascular plants rich in lignin, which degrades slowly, and thus forces the decrease of the δ¹³C values during decay, the δ¹³C values of Z. noltii leaves did not vary significantly over time or between incubation sites. The δ¹⁵N values of leaves did not increase as has been reported for vascular plants with high C:N ratio. This suggests that during the early phase of the Z. noltii decomposition the community of decomposers relied mostly on the endogenous nitrogen of the plants, not needing to immobilize exogenous N. Stable isotope studies to assess the contribution of this species to the food web can rely on the natural δ¹³C and δ¹⁵N abundances of living tissues without any correction to account for decomposition. The analysis of the δ³⁴S values of Z. noltii detritus was not useful probably due to the contamination of the samples with pyrite, which has a depleted δ³⁴S signal.     

Sea-ice algae: Major contributors to primary production and algal biomass in the Chukchi and Beaufort Seas during May/June 2002

Sea-ice and water samples were collected at 14 stations on the shelves and slope regions of the Chukchi and Beaufort Seas during the spring 2002 expedition as part of the Shelf–Basin Interaction Studies. Algal pigment content, particulate organic carbon and nitrogen, and primary productivity were estimated for both habitats based on ice cores, brine collection and water samples from 5-m depth. The pigment content (0.2–304.3 mg pigments m⁻²) and primary productivity (0.1–23.0 mg C m⁻³ h⁻¹) of the sea-ice algae significantly exceeded water-column parameters (0.2 and 1.0 mg pigments m⁻³; <0.1–0.4 mg C m⁻³ h⁻¹), making sea ice the habitat with the highest food availability for herbivores in early spring in the Chukchi and Beaufort Seas. Stable isotope signatures for ice and water samples did not differ significantly for δ¹⁵N, but for δ¹³C (ice: −25.1‰ to −14.2‰; water: −26.1‰ to −22.4‰). The analysis of nutrient concentrations and the pulse-amplitude-modulated fluorescence signal of ice algae and phytoplankton indicate that nutrients were the prime limiting factor for sea-ice algal productivity. The estimated spring primary production of about 1–2 g C m⁻² of sea-ice algae on the shelves requires the use of substantial nutrient reservoirs from the water column.     

基于碳、氮稳定同位素的雅浦海沟底栖生物食物来源和营养级初探

为了分析雅浦海沟中底栖生物群落的食物来源和营养级,本研究分析了雅浦海沟真光层中浮游植物和浮游动物、海底沉积物和巨型底栖生物（海绵、海参、海蛇尾、海星、海葵和钩虾）中的碳、氮稳定同位素组成。研究发现雅浦海沟真光层中的浮游植物和浮游动物δ13C值[（-22.8±0.4）‰和（-21.8±0.8）‰]和δ15N值[（5.4±0.4）‰和（6.8±0.2）‰]与巨型底栖生物的δ13C值（-20.1‰~-16.8‰）和δ15N值（11.9‰~17.9‰）的差异超过了一个营养级,表明作为底栖生物的初始食物来源的浮游植物和浮游动物在向下输送的过程中经历了食物链传递和细菌的降解。巨型底栖生物的δ15N和δ13C值之间无显著的相关性,此外不同物种之间营养级也存在明显差异,表现为海绵的营养级相对较高（3.4~4.7）,海参（3.3~3.6）、海蛇尾（3.4~3.5）和海星（3.2~3.7）的营养级较为接近,钩虾（2.9~3.3）和海葵（3.1）的营养级则相对略低,反映了底栖生物不同物种之间食物来源的多样化。     

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.     

Vertical changes in abundance,biomass and community structure of copepods down to 3000 m in the southern Bering Sea

Vertical changes in abundance, biomass and community structure of copepods down to 3000 m depth were studied at a single station of the Aleutian Basin of the Bering Sea (53°28′N, 177°00′W, depth 3779 m) on the 14th June 2006. Both abundance and biomass of copepods were greatest near the surface layer and decreased with increase in depth. Abundance and biomass of copepods integrated over 0–3000 m were 1,390,000 inds. m^?2 and 5056 mg C m^?2, respectively. Copepod carcasses occurred throughout the layer, and the carcass:living specimen ratio was the greatest in the oxygen minimum layer (750–100 m, the ratio was 2.3). A total of 72 calanoid copepod species belonging to 34 genera and 15 families occurred in the 0–3000 m water column (Cyclopoida, Harpacticoida and Poecilostomatoida were not identified to species level). Cluster analysis separated calanoid copepod communities into 5 groups (A–E). Each group was separated by depth, and the depth range of each group was at 0–75 m (A), 75–500 m (B), 500–750 m (C), 750–1500 m (D) and 1500–3000 m (E). Copepods were divided into four types based on the feeding pattern: suspension feeders, suspension feeders in diapause, detritivores and carnivores. In terms of abundance the most dominant group was suspension feeders (mainly Cyclopoida) in the epipelagic zone, and detritivores (mainly Poecilostomatoida) were dominant in the meso- and bathypelagic zones. In terms of biomass, suspension feeders in diapause (calanoid copepods Neocalanus spp. and Eucalanus bungii) were the major component (ca. 10–45%), especially in the 250–3000 m depth. These results are compared with the previous studies in the same region and that down to greater depths in the worldwide oceans.