The trophic ecology of key megafaunal species at the Pakistan Margin: Evidence from stable isotopes and lipid biomarkers

The Arabian Sea is subject to intense seasonality resulting from biannual monsoons, which lead to associated large particulate fluxes and an abundance of organic carbon, a potential food source at the seafloor for benthic detritivores. We used the stable isotopes of carbon and nitrogen alongside lipid analyses to examine potential food sources (particulate and sedimentary organic matter, POM and SOM respectively) in order to determine trophic linkages for the twelve most abundant megafaunal species (Pontocaris sp., Solenocera sp., Munidopsis aff. scobina, Actinoscyphia sp., Actinauge sp., Echinoptilum sp., Pennatula aff. grandis, Astropecten sp. Amphiura sp. Ophiura euryplax, Phormosoma placenta and Hyalinoecia sp.) at the Pakistan Margin between 140 and 1400 m water depth. This transect spans a steep gradient in oxygen concentrations and POM flux. Ranges of δ¹³C and δ¹⁵N values were narrow in POM and SOM (4‰ and 2‰ for δ¹³C and δ¹⁵N, respectively) with little evidence of temporal variability. Labile lipid compounds in SOM originating from phytoplankton did exhibit seasonal change in their concentrations at the shallowest sites, 140 and 300 m. Benthic megafauna had broad ranges in δ¹³C and δ¹⁵N (>10‰ and >8‰ for δ¹³C and δ¹⁵N, respectively) suggesting they occupy several trophic levels and utilize a variety of food sources. There is evidence for feeding niche separation between and within trophic groups. Lipid biomarkers in animal tissues indicate a mixture of food sources originating from both phytoplankton (C_20:5(n-3) and C_22:6(n-3)) and invertebrate prey (C_20:1 and C_22:1). Biomarkers originating from phytodetritus are conserved through trophic transfer to the predator/scavengers. Six species (Pontocaris sp., Solenocera sp., Actinoscyphia sp., Echinoptilum sp., Amphiura sp. and Hyalinoecia sp.) showed a significant biochemical response to the seasonal supply of food and probably adapt their trophic strategy to low food availability. Biotransformation of assimilated lipids by megafauna is evident from polyunsaturated fatty acid distributions, for example, Echinoptilum sp. converts C_20:5(n-3) to C_24:6(n-3).     

Isotopic variation of fishes in freshwater and estuarine zones of a large subtropical coastal lagoon

We used stable C and N isotope ratios of tissues from 29 fish species from a large subtropical lagoon in southern Brazil to examine spatial variability in isotopic composition and vertical trophic structure across freshwater and estuarine habitats. Nitrogen isotope ratios indicated a smooth gradation in trophic positions among species, with most fishes occupying the secondary and tertiary consumer level. Fish assemblages showed a significant shift in their carbon isotopic signatures between freshwater and estuarine sites. Depleted carbon signatures (from −24.7‰ to −17.8‰) were found in freshwater, whereas more enriched signatures (from −19.1‰ to −12.3‰) were obtained within the estuarine zone downstream. Based on our survey of the C₃ and C₄ plants and isotopic values for phytoplankton and benthic microalgae reported for ecosystems elsewhere, we hypothesized that the observed δ¹³C differences in the fish assemblage between freshwater and estuarine sites is due to a shift from assimilating organic matter ultimately derived from C₃ freshwater marsh vegetation and phytoplankton at the freshwater site (δ¹³C ranging from −25‰ to −19‰), to C₄ salt-marsh (e.g. Spartina) and widgeon grass (Ruppia maritima), benthic microalgae and marine phytoplankton at the estuarine sites (from −18‰ to −12‰). Our results suggested that fish assemblages are generally supported by autochthonous primary production. Freshwater fishes that likely were displaced downstream into the estuary during periods of high freshwater discharge had depleted δ¹³C values that were characteristic of the upper lagoon. These results suggest that spatial foodweb subsidies can occur within the lagoon.     

Oceanographic controls on the carbon isotopic compositions of sinking particles from the Cariaco Basin

This study examined the relationship between carbon isotopic composition of sinking organic matter (OM) and the biological, physical and chemical properties of the surface ocean in the Cariaco Basin. The ¹³C/¹²C ratio of OM (δ¹³C_org) in sinking particles was determined on sediment trap samples from four depths collected from 1996 to 1999 as part of the CArbon Retention In A Colored Ocean time series. Water column properties, including temperature, productivity, chlorophyll and concentration of dissolved CO₂, were concurrently measured on monthly cruises. The δ¹³C_org varied from a high of –17.7‰ to a low of –22.6‰ during the study period. The variation of the δ¹³C_org throughout seasonal cycles was directly proportional to the strength of upwelling and was negatively correlated with temperature (r²=0.64). During the 1996–1997 upwelling event, the strongest during the study period, the δ¹³C_org increased by 4.4‰ whereas during the 1998–1999 upwelling event, the weakest during the study period, the δ¹³C_org only increased by 3.3‰. Contrary to most previous studies, we observed a negative relationship (r²=0.53) between [CO_2 aq] and the estimated isotopic fractionation factor (ε_p). However, there was no correlation between ε_p and the calculated growth rates indicating that there was non-diffusive uptake of carbon into phytoplankton cells. It thus appears that [CO_2 aq] does not control the δ¹³C_org in the water column of the study site. The best explanation for the isotopic enrichment observed is a carbon concentrating mechanism (CCM) in phytoplankton. The existence of a CCM in phytoplankton has major implications for the interpretation of the δ¹³C_org in the Cariaco Basin.     

Stable carbon isotope ratios of plankton carbon and sinking organic matter from the Atlantic sector of the Southern Ocean

The stable carbon isotope composition of particulate organic carbon (POC) from plankton, sediment trap material and surface sediments from the Atlantic sector of the Southern Ocean were determined. Despite low and constant water temperatures, large variations in the δ¹³C values of plankton were measured. ¹³C enrichments of up to 10‰ coincided with a change in the diatom assemblage and a two-fold increase in primary production. Increased CO₂ consumption as a result of rapid carbon fixation may result in diffusion limitation reducing the magnitude of the isotope fractionation. The δ¹³C values of plankton from sea-ice cores display a relationship with the chlorophyll a content. High ‘ice-algae’ biomass, in combination with a limited exchange with the surrounding seawater, results in values of about − 18 to − 20‰. It is assumed that these values are related to a reduced CO₂ availability in the sea-ice system. In comparison with plankton, sinking krill faeces sampled by traps can be enriched by 2–5‰ in ¹³C (e.g. central Bransfield Strait). In contrast, the transport of particles in other faeces, diatom aggregates or chains results in minor isotope changes (e.g. Drake Passage, Powell Basin, NW Weddell Sea). A comparison between the δ¹³C values of sinking matter and those of surface sediments reveals that ¹³C enrichments of up to 3–4‰ may occur at the sediment-water boundary layer. These isotopic changes are attributed to high benthic respiration rates.     

Sources and transport of dissolved and particulate organic carbon in the Mississippi River estuary and adjacent coastal waters of the northern Gulf of Mexico

Dissolved organic carbon (DOC), stable carbon isotopic (δ¹³C) compositions of DOC and particulate organic carbon (POC), and elemental C/N ratios of POC were measured for samples collected from the lower Mississippi and Atchafalaya rivers and adjacent coastal waters in the northern Gulf of Mexico during the low flow season in June 2000 and high flow season in April 2001. These isotopic and C/N results combined with DOC measurements were used to assess the sources and transport of terrestrial organic matter from the Mississippi and Atchafalaya rivers to the coastal region in the northern Gulf of Mexico. δ¹³C values of both POC (−23.8‰ to −26.8‰) and DOC (−25.0‰ to −29.0‰) carried by the two rivers were more depleted than the values measured for the samples collected in the offshore waters. Strong seasonal variations in δ¹³C distributions were observed for both POC and DOC in the surface waters of the region. Fresh water discharge and horizontal mixing played important roles in the distribution and transport of terrestrial POC and DOC offshore. Our results indicate that both POC and DOC exhibited non-conservative behavior during the mixing especially in the mid-salinity range. Based on a simple two end-member mixing model, the comparison of the measured DOC-δ¹³C with the calculated conservative isotopic mixing curve indicated that there was a significant in situ production of marine-derived DOC in the mid- to high-salinity waters consistent with our in situ chlorophyll-a measurements. Our DOC-δ¹³C data suggest that a removal of terrestrial DOC mainly occurred in the high-salinity (>25) waters during the mixing. Our study indicates that the mid- to high- (10–30) salinity range was the most dynamic zone for organic carbon transport and cycling in the Mississippi River estuary. Variability in isotopic and elemental compositions along with variability in DOC and POC concentrations suggest that autochthonous production, bacterial utilization, and photo-oxidation could all play important roles in regulating and removing terrestrial DOC in the northern Gulf of Mexico and further study of these individual processes is warranted.     

Carbon and nitrogen isotope composition of particulate organic matter in relation to mucilage formation in the northern Adriatic Sea

Carbon and nitrogen stable isotope ratios of particulate organic matter (POM) were studied approximately weekly during spring and summer 2003 and 2004 in the Gulf of Trieste (northern Adriatic Sea) in order to track the temporal variations and differences between two years. In parallel, particulate organic carbon (POC) and particulate nitrogen (PN), phytoplankton biomass (chlorophyll a), and N and P nutrients were monitored. All studied parameters, especially N and P nutrients and chlorophyll a, showed higher concentrations and larger variability in spring 2004. As a consequence the macroaggregates were produced in late spring 2004. The C and N isotope composition of POM was not directly linked to phytoplankton biomass dynamics. The δ¹³C_POC values covaried with temperature. In 2004, δ¹³C_POC variations followed the δ¹⁵N_PN values as well as the δ¹³C_DIC values which were probably more dependent on the photosynthetic use of ¹²C. Variations in δ¹⁵N_POM values were most probably the consequence of variations in N nutrient sources used in phytoplankton assimilation. The significant correlation between δ¹⁵N_PN values and nitrate concentrations in 2004 implies intense nitrate assimilation in the presence of higher nitrate concentration. This suggests nitrate as the key nutrient in the »new primary production«, later producing macroaggregates with a mean δ¹³C and δ¹⁵N values of − 19‰ and 5‰, respectively. A low fractionation factor ε, < 1‰, lower than that reported in other marine and lacustrine systems, was found probably to be a consequence of distinct phytoplankton species, i.e. several classes of autotrophic nanoflagellates, and specific growth conditions present in the Gulf of Trieste. The tentative use of C isotope composition of POM revealed a higher contribution of allochthonous organic matter in 2004 compared to 2003 due to higher riverine inflow.     

Origin and decomposition of sinking particulate organic matter in the deep water column inferred from the vertical distributions of its δN, δ and δ

Sinking particles were analyzed for their nitrogen isotopic ratio δ¹⁵N) of total particulate nitrogen (PN), stable carbon isotopic ratio (δ¹³C) and radioactive isotopic ratio (δ¹⁴C) of total particulate organic carbon (POC), at three different latitudinal (temperate, subpolar and equatorial) and geomorphological (trench, proximal abyssal plain and distal abyssal plain) sites in the western North Pacific Ocean using year-long time series sediment trap systems, to clarify the common vertical trends of the isotopic signals in deep water columns. Although the δ¹⁵N and δ¹³C values of sinking particulate organic matter (POM) were partly affected by the resuspension of sedimentary POM from the sea floor, especially in the trench, the changes in δ¹⁵N and δ¹³C values owing to the resuspension could be corrected by calculation of the isotopic mass balance from δ¹⁴C of sinking POC. After this correction, common downward decreasing trends in δ¹⁵N and δ¹³C values were obtained in the deep water columns, irrespective of the latitudes and depths. These coincidental isotopic signals between δ¹⁵N and δ¹³C values provide new constraints for the decomposition process of sinking POM, such as the preferential degradation of ¹⁵N- and ¹³C-rich compounds and the successive re-formation of the sinking particles by higher trophic level organisms in the deep water column.     

Stable carbon isotope distribution of particulate organic matter in the ocean: a model study

The stable carbon isotopic composition of particulate organic matter in the ocean, δ¹³C_POC, shows characteristic spatial variations with high values in low latitudes and low values in high latitudes. The lowest δ¹³C_POC values (−32‰ to −35‰) have been reported in the Southern Ocean, whereas in arctic and subarctic regions δ¹³C_POC values do not drop below −27‰. This interhemispheric asymmetry is still unexplained. Global gradients in δ¹³C_POC are much greater than in δ¹³C_DIC, suggesting that variations in isotopic fractionation during organic matter production are primarily responsible for the observed range in δ¹³C_POC. Understanding the factors that control isotope variability is a prerequisite when applying δ¹³C_POC to the study of marine carbon biogeochemistry. The present model study attempts to reproduce the δ¹³C_POC distribution pattern in the ocean. The three-dimensional (3D) Hamburg Model of the Oceanic Carbon Cycle version 3.1 (HAMOCC3.1) was combined with two different parametrizations of the biological fractionation of stable carbon isotopes. In the first parametrization, it is assumed that the isotopic fractionation between CO₂ in seawater and the organic material produced by algae, _P, is a function of the ambient CO₂ concentration. The two parameters of this function are derived from observations and are not based on an assumption of any specific mechanism. Thus, this parametrization is purely empirical. The second parametrization is based on fractionation models for microalgae. It is supported by several laboratory experiments. Here the fractionation, _P, depends on the CO₂ concentration in seawater and on the (instantaneous) growth rates, μ_i, of the phytoplankton. In the Atlantic Ocean, where most field data are available, both parametrizations reproduce the latitudinal variability of the mean δ¹³C_POC distribution. The interhemispheric asymmetry of δ¹³C_POC can mostly be attributed to the interhemispheric asymmetry of CO₂ concentration in the water. However, the strong seasonal variations of δ¹³C_POC as reported by several authors, can only be explained by a growth rate-dependent fractionation, which reflects variations in the cellular carbon demand.     

Distribution and sources of organic carbon, nitrogen and their isotopes in sediments of the subtropical Pearl River estuary and adjacent shelf, Southern China

The isotopic composition (δ¹³C and δ¹⁵N) and organic carbon (OC) and total nitrogen (TN, organic plus inorganic) content of 37 carbonate-free surficial sediments of the subtropical Pearl River estuary and the adjacent shelf of South China Sea (SCS) was determined. The δ¹³C values indicate that the sediment organic material is a mixture from two sources, terrestrial and marine. Several of the sediments have extremely low (< 4) OC / TN ratios, which could be due to low OC contents and/or to a significant fraction of the TN present as inorganic nitrogen adsorbed on clays. In general, the spatial patterns of OC, TN, δ¹³C and δ¹⁵N are similar. Values are low at the river mouth and on the western coast, suggesting proportionally greater accumulation of terrestrial particulate organic matter relative to marine phytodetritus, which is limited by low productivity in the turbid plume of the Pearl River. Algal-derived organic carbon (al-OC) content is estimated to be low (≤ 0.06%) at the river mouth and higher (up to 0.57%) on the adjacent inner shelf based on a mixing model of end members.     

Sub-recent nitrogen-isotope trends in sediments from Skagerrak (North Sea) and Kattegat: Changes in N-budgets and N-sources?

We determined ¹⁵N/¹⁴N ratios of total nitrogen in surface sediments and dated sediment cores to reconstruct the history of N-loading of the North Sea. The isotopic N composition in modern surface sediments is equivalent to and reflects the isotopic mixture of oceanic nitrate on the one hand (δ¹⁵N = 5‰) and the imprint of river-borne nitrogen input into the SE North Sea (δ¹⁵N up to 12‰ in estuaries of the SE North Sea) on the other hand. We compare the results with δ¹⁵N records from pre-industrial sediment intervals in cores from the Skagerrak and Kattegat areas, which both constitute significant depositional centres for N in the North Sea and the Baltic Sea/North Sea transition. As expected, isotopically enriched anthropogenic nitrogen was found in the two records from the Kattegat area, which is close to eutrophication sources on land. Enrichment of δ¹⁵N in cores from the Skagerrak – the largest sediment sink for nitrogen in the entire North Sea – was not significant and values were similar to those found in sediment layers representing pre-industrial conditions. We interpret this isotopic uniformity as an indication that most riverine reactive nitrogen with its characteristic isotopic signature is removed by denitrification in shallow shallow-water sediments before reaching the main sedimentary basin of the North Sea.     

Food web structure in a near-pristine mangrove area of the Australian Wet Tropics

Carbon and nitrogen isotopic composition was used to identify the main sources of carbon and describe the main trophic pathways in Deluge Inlet, a near-pristine mangrove estuary in tropical north Queensland, Australia. Producers' δ¹³C varied from −28.9‰ for mangroves to −18.6‰ for seagrass. Animals were also well separated in δ¹³C (−25.4‰ to −16.3‰ for invertebrates and −25.2‰ to −17.2‰ for fish), suggesting considerable differences in ultimate sources of carbon, from a substantial reliance on mangrove carbon to an almost exclusive reliance on seagrass. In general, invertebrates had lower δ¹⁵N than fish, indicating lower trophic levels. Among fish, δ¹⁵N values reflected well the assumed trophic levels, as species from lower trophic levels had lower δ¹⁵N than species from higher trophic levels. Trophic levels and trophic length were estimated based on δ¹⁵N of invertebrate primary consumers (6.1‰), with results suggesting a food web with four trophic levels. There was also evidence of a high level of diet overlap between fish species, as indicated by similarities in δ¹³C for fish species of higher trophic levels. Stable isotope data was also useful to construct a general model for this food web, where five main trophic pathways were identified: one based on both mangrove and microphytobenthos, one on plankton, two on both microphytobenthos and seagrass, and one based mainly on seagrass. This model again suggested the presence of four trophic levels, in agreement with the value calculated based on the difference in δ¹⁵N between invertebrate primary consumers and top piscivores.     

Sulfur and iron speciation in surface sediments along the northwestern margin of the Black Sea

The speciation of sedimentary sulfur (pyrite, acid volatile sulfides (AVS), S⁰, H₂S, and sulfate) was analyzed in surface sediments recovered at different water depths from the northwestern margin of the Black Sea. Additionally, dissolved and dithionite-extractable iron were quantified, and the sulfur isotope ratios in pyrite were measured. Sulfur and iron cycling in surface sediments of the northwestern part of the Black Sea is largely influenced by (1) organic matter supply to the sediment, (2) availability of reactive iron compounds and (3) oxygen concentrations in the bottom waters. Biologically active, accumulating sediments just in front of the river deltas were characterized by high AVS contents and a fast depletion of sulfate concentration with depth, most likely due to high sulfate reduction rates (SRR). The δ³⁴S values of pyrite in these sediments were relatively heavy (−8‰ to −21‰ vs. V-CDT). On the central shelf, where benthic mineralization rates are lower, re-oxidation processes may become more important and result in pyrite extremely depleted in δ³⁴S (−39‰ to −46‰ vs. V-CDT). A high variability in δ³⁴S values of pyrite in sediments from the shelf-edge (−6‰ to −46‰ vs. V-CDT) reflects characteristic fluctuations in the oxygen concentrations of bottom waters or varying sediment accumulation rates. During periods of oxic conditions or low sediment accumulation rates, re-oxidation processes became important resulting in low AVS concentrations and light δ³⁴S values. Anoxic conditions in the bottom waters overlying shelf-edge sediments or periods of high accumulation rates are reflected in enhanced AVS contents and heavier sulfur isotope values. The sulfur and iron contents and the light and uniform pyrite isotopic composition (−37‰ to −39‰ vs. V-CDT) of sediments in the permanently anoxic deep sea (1494 m water depth) reflect the formation of pyrite in the upper part of the sulfidic water column and the anoxic surface sediment. The present study demonstrates that pyrite, which is extremely depleted in ³⁴S, can be found in the Black Sea surface sediments that are positioned both above and below the chemocline, despite differences in biogeochemical and microbial controlling factors.     

Stable isotope analysis of some representative fish and invertebrates of the Newfoundland and Labrador continental shelf food web

We examined stable carbon and nitrogen isotopic signatures of 17 fish and 16 invertebrate taxa common to the Newfoundland and Labrador (NL) continental shelf food web. Particular sampling emphasis was placed on Atlantic cod (Gadus morhua) and related prey species (e.g. shrimp, Pandalus borealis, and capelin, Mallotus villosus). We found highly significant (p < 0.0001) differences between near-shore (bays) and offshore (shelf edge) δ¹⁵N signatures for cod, ‘other fish’ (pooled) and invertebrates (pooled). In contrast, there were only minor differences in δ¹³C signatures of ‘other fish’ (p < 0.05) and no difference for cod and invertebrates among the two habitats. We sampled at two times of the year (January and June) and found no systematic effect of season on both δ¹³C and δ¹⁵N in cod, ‘other fish’ and invertebrates. We calculated isotopic fractionation factors for cod from the entire shelf (mixed diet) and for cod with diets composed mainly of capelin or shrimp. These values ranged between 2.2‰ and 3.9‰ for δ¹⁵N and −0.4‰ and 0.8‰ for δ¹³C and, for δ¹⁵N, may reflect diet-related differences in bioenergetic status. We discuss potential mechanisms for near-shore versus offshore enrichment of δ¹⁵N signatures, and demonstrate the implications of this spatial variation on δ¹⁵N-derived trophic position estimates.     

Carbon and nitrogen stable isotopes in suspended matter and sediments from the Schelde Estuary

The C/N and stable C and N isotope ratios (δ¹³C, δ¹⁵N) of sedimentary and suspended particulate matter were determined in the Schelde Estuary. Suspended matter was divided into 2 to 5 size fractions by centrifugation. Four major pools of organic matter were recognized: riverine, estuarine, marine and terrestrial materials. Terrestrial organic matter (δ¹³C≈−26‰, δ¹⁵N≈3.5‰, C/N≈21) is important for the sedimentary pool, but suspended matter is dominated by the marine (δ¹³C≈−18‰, δ¹⁵N≈9‰, C/N≈8), riverine (δ¹³C≈−30‰, δ¹⁵N≈9‰, C/N≈7.5) and estuarine (δ¹³C≈−29‰, δ¹⁵N≈15‰, C/N≈8) end-members. In the upper estuary, the suspended matter size fractions vary systematically in their carbon and nitrogen biogeochemistry, with the small particles having low C/N ratios, depleted δ¹³C and enriched δ¹⁵N values relative to large particles. Moreover, sedimentary and suspended matter differ significantly in terms of C/N ratios (17 vs. 8.9), δ¹³C (−26.3 vs. −28.9‰) and δ¹⁵N (+6.9 vs. 12.0‰). In the lower estuary, suspended matter fractions are similar and sedimentary and suspended organic matter differ only in terms of δ¹³C (−23.5 vs. −20.1‰). Our data indicate that autochthonous organic matter contributes significantly to the total suspended matter and that the suspended organic matter composition cannot be explained in terms of conservative mixing of riverine and terrestrial sources on the one hand and marine sources on the other hand.     

The nearshore western Beaufort Sea ecosystem: Circulation and importance of terrestrial carbon in arctic coastal food webs

The nearshore shelf of the Beaufort Sea is defined by extreme physical and biological gradients that have a distinctive influence on its productivity and trophic structure. Massive freshwater discharge from the Mackenzie River, along with numerous smaller rivers and streams elsewhere along the coast, produce an environment that is decidedly estuarine in character, especially in late spring and summer. Consequently, the Beaufort coast provides a critical habitat for several species of amphidromous fishes, some of which are essential to the subsistence lifestyle of arctic native populations. Because of its low in situ productivity, allochthonous inputs of organic carbon, identifiable on the basis of isotopic composition, are important to the functioning of this arctic estuarine system. Coastal erosion and river discharge are largely responsible for introducing high concentrations of suspended sediment from upland regions into the nearshore zone. The depletion in the ¹³C content of invertebrate and vertebrate consumers, which drops about 4–5‰ eastward along the eastern Alaskan Beaufort Sea coast, may reflect the assimilation of this terrestrial organic matter into local food webs. In addition, the large range in ¹³C values of fauna collected in the eastern Beaufort (nearly 8‰) compared to the same species in the northeastern Chukchi (3‰), indicate a lower efficiency of carbon transfer between trophic levels in the eastern Beaufort. The wider spread in stable isotope values in the eastern Beaufort may also reflect a decoupling between benthic and pelagic components. Isotopic tracer studies of amphidromous fishes in the Simpson Island barrier island lagoon revealed that terrestrial (peat) carbon may contribute as much as 30–50% of their total dietary requirements. On the eastern Alaska Beaufort Sea coast, the δ¹³C values of arctic cod collected in semi-enclosed lagoons were more depleted, by 3–4‰, compared to fish collected in the coastal Beaufort Sea. Calculations from isotopic mixing equations indicate cod from lagoons may derive 70% of their carbon from terrestrial sources. The δ¹⁵N values of lagoon fish were also 4‰ lower than coastal specimens, reflective of the lower δ¹⁵N values of terrestrially derived nitrogen (0–1.5‰ compared to 5–7‰ for phytoplankton). The role of terrestrial carbon in arctic estuarine food webs is especially important in view of the current warming trend in the arctic environment and the role of advective processes that transport carbon along the nearshore shelf. Biogeochemical studies of the arctic coastal estuarine environment may provide more insights into the function of these biologically complex ecosystems.     

Nitrogen sources in the South China Sea, as discerned from stable nitrogen isotopic ratios in rivers, sinking particles, and sediments

Stable nitrogen isotopic ratios were measured in sinking particles and surface sediments from the South China Sea (SCS) in order to study recent nitrogen sources and degradation. Average δ¹⁵N values of 16 sediment traps deployed at seven locations in the northern, central and southern SCS were uniformly low, ranging between 2.7 and 4.5‰ with a winter minimum in the northern and central SCS. Enhanced nitrogen contents and δ¹⁵N values were noted in samples affected by swimmers, comprising between 5 and 20% of total nitrogen fluxes. Nitrate sources were subsurface waters from the western Pacific, which were isotopically depleted due to the remineralization of nitrogen from nitrogen fixation in surface waters. Nitrogen fixation in the SCS contributed up to 20% to the settling particles. In the southern SCS, resuspended matter close to the shelf added to the sinking particulates. The long-term trap record from the central SCS revealed decreasing δ¹⁵N values during the 1990s, which correspond with findings from the North Pacific Subtropical Gyre and may be attributable to increased nitrogen fixation due to global warming-related stratification. This trend may be restricted to the 1990s but could also persist due to the projection of more frequent occurrence of El Niño conditions.The δ¹⁵N increase from swimmer-free trap averages of 2.7–3.6‰ to values of 5–6‰ in underlying deep-sea sediments was in the same range as in other deep ocean areas. Similar to results from the northern Indian Ocean, this increase could be related to isotopic enrichment during amino acid degradation. The lowest sedimentary δ¹⁵N values characterize the Pinatubo ash layer deposited off Luzon in an event of mass sedimentation in 1991. The fast deposition of organic matter drawn from the surface waters with the ash in the form of vertical density currents evidently preserved the planktonic δ¹⁵N signal.     

Organic matter characterization in a tropical estuarine-mangrove ecosystem of India: Preliminary assessment by using stable isotopes and lignin phenols

In order to characterize the sources and fate of organic matter (OM) in the Pichavaram estuarine-mangrove ecosystem (east coast of India), stable isotope (δ¹³C and δ¹⁵N) ratios and molecular lignin analyses were conducted in plant litter, benthic algae, sediment, particulate matter and in a variety of benthic invertebrate species. The δ¹³C signature of plant litter ranges from −29.75‰ to −27.64‰ suggesting that mangrove trees follow the C₃ photosynthetic pathway. Sedimentary δ¹³C signature (−28.92‰ to −25.34‰) demonstrates the greater influence of plant litter organic matter on sedimentary organic matter. Suspended particulate organic pool was influenced by terrestrial source and also seems to be influenced by the marine phytoplankton. Enriched signature of δ¹⁵N in surface sediments (4.66–8.01‰; avg. 6.69‰) suggesting the influence of anthropogenic nitrogen from agricultural fields and human settlements. Spatial chemical variability in availability of nitrogen and plant associated microbial interactions demonstrate variability in δ¹⁵N signature in mangrove plant litter. Two (lower and higher) trophic levels of invertebrates were identified with and observed >4‰ gradient in δ¹³C signal between these two trophic groups. The observed δ¹³C values suggest that the lower level invertebrates feed on phytoplankton and higher level organisms have a mixed source of diet, phytoplankton, sediment and particulate organic matter. Lignin phenol analyses explain that the benthic surface layer was almost free of lignin. The ratio between syringyl phenols to vanillyl phenols (S/V) is 1.14–1.32 (avg. 1.23) and cinnamyl phenols to vanillyl phenols (C/V) is 0.17–0.31 (avg. 0.24), demonstrate non-woody angiosperm tissues was the major sources of lignin to this ecosystem, while aldehyde to acid ratios (Ad/Al) describe diagenetic nature of sediment and is moderately to less degraded. A two-end-member mixing model indicate that the terrigenous OM was dominant in the estuarine zones, while in the mangrove zone terrigenous supply accounts for 60% and marine input accounts for 40%.     

On the sulphur chemistry of a super-anoxic fjord, Framvaren, South Norway

The concentrations of total carbonate (C_t), sulphate, sulphide, thiols and oxygen, the ratio between the stable sulphur isotopes ³⁴S and ³²S in sulphate and sulphide, and the density (used to calculate salinity) were determined on samples from the water column of Framvaren, a superanoxic fjord in southern Norway. From a depth of 18m (the oxic-anoxic boundary) the initial sulphate concentration, ([SO₄]_init), as calculated from salinity, is significantly higher than the sum of the measured sulphur species. This is attributed to a loss of sulphur from the water column. The amount of total carbonate produced, corrected for the initial concentration (C_t - 2.4 Sal/35) is found to be proportional to the amount of sulphate consumed, ([SO₄]_init - [SO₄]), according to the following relation C_t- 2.4 Sal/35 = 1.84 ([SO₄]_init - [SO₄]). Isotopic fractionation caused by bacterial sulphate reduction in the anoxic part of the water column produces sulphide with a δ³⁴S 40‰ lower than the δ³⁴S for sulphate at corresponding depths. The isotopic fractionation also results in δ³⁴S value for the remaining sulphate at depths below 80 m being considerably higher than the mean value for ocean water, which is close to + 20‰. The δ³⁴S values for sulphate at depths between 10 and 50 m were lower than + 20‰ which indicates oxidation of sulphide, which follows upon diffusion of sulphide from deeper parts of the water column and inflow of oxygenated seawater over the sill into the anoxic water of the fjord. A conclusive scenario of the Framvaren sulphur chemistry is presented.     

Benthic remineralization at the land–ocean interface: A case study of the Rhône River (NW Mediterranean Sea)

Biogeochemical processes in sediments under the influence of the Rhône River plume were studied using both in situ microelectrodes and ex situ sediment core incubations. Organic carbon (OC) and total nitrogen (TN) content as well as stable carbon isotopic composition of OC (δ¹³C_OC) were analysed in 19 surface sediments to determine the distribution and sources of organic matter in the Rhône delta system. Large spatial variations were observed in both the total O₂ uptake (5.2 to 29.3 mmol m⁻² d⁻¹) and NH₄⁺ release (−0.1 to −3.5 mmol m⁻² d⁻¹) rates at the sediment–water interface. The highest fluxes were measured near the Rhône River mouth where sedimentary OC and TN contents reached 1.81% and 0.23% respectively. Values of δ¹³C_OC ranged from −26.83‰ to −23.88‰ with a significant seawards enrichment tracing the dispersal of terrestrial organic matter on the continental shelf. The amount of terrestrial-derived OC reaches 85% in sediments close to the Rhône mouth decreasing down to 25% in continental shelf sediments. On the prodelta, high terrestrial OC accumulation rates support high oxygen uptake rates and thus indicating that a significant fraction of terrestrial OC is remineralized. A particulate organic carbon (POC) mass balance indicates that only 3% of the deposited POC is remineralized in prodelta sediments while 96% is recycled on the continental shelf. It was calculated that a large proportion of the Rhône POC input is either buried (52%) or remineralized (8%), mostly on the prodelta area. The remaining fraction (40%) is either mineralized in the water or exported outside the Rhône delta system in dissolved or particulate forms.     

Redefining the trophic importance of seagrasses for fauna in tropical Indo-Pacific meadows

Fauna species living in seagrass meadows depend on different food sources, with seagrasses often being marginally important for higher trophic levels. To determine the food web of a mixed-species tropical seagrass meadow in Sulawesi, Indonesia, we analyzed the stable isotope (δ¹³C and δ¹⁵N) signatures of primary producers, particulate organic matter (POM) and fauna species. In addition invertebrates, both infauna and macrobenthic, and fish densities were examined to identify the important species in the meadow. The aims of this study were to identify the main food sources of fauna species by comparing isotopic signatures of different primary producers and fauna, and to estimate qualitatively the importance of seagrass material in the food web. Phytoplankton and water column POM were the most depleted primary food sources for δ¹³C (range −23.1 to −19.6‰), but no fauna species depended only on these sources for carbon. Epiphytes and Sargassum sp. had intermediate δ¹³C values (−14.2 to −11.9‰). Sea urchins, gastropods and certain fish species were the main species assimilating this material. Seagrasses and sedimentary POM had the least depleted values (−11.5 to −5.7‰). Between the five seagrass species significant differences in δ¹³C were measured. The small species Halophila ovalis and Halodule uninervis were most depleted, the largest species Enhalus acoroides was least depleted, while Thalassia hemprichii and Cymodocea rotundata had intermediate values. Fourteen fauna species, accounting for 10% of the total fauna density, were shown to assimilate predominantly (>50%) seagrass material, either directly or indirectly by feeding on seagrass consumers. These species ranged from amphipods up to the benthic top predator Taeniura lymma. Besides these species, about half of the 55 fauna species analyzed had δ¹³C values higher than the least depleted non-seagrass source, indicating they depended at least partly for their food on seagrass material. This study shows that seagrass material is consumed by a large number of fauna species and is important for a large portion of the food web in tropical seagrass meadows.