Trophic ecology of European sardine Sardina pilchardus and European anchovy Engraulis encrasicolus in the Bay of Biscay (north-east Atlantic) inferred from δ13C and δ15N values of fish and identified mesozooplanktonic organisms

European sardine (Sardina pilchardus) and European anchovy (Engraulis encrasicolus) are two species of economical and ecological significance in the Bay of Biscay (north-east Atlantic). However, the trophic ecology of both species is still poorly known in the area, and more generally, few studies have considered the potential trophic overlap between sardines and anchovies worldwide. This study aims to highlight the trophic links between the mesozooplankton and adults of these two pelagic fish in the Bay of Biscay, through carbon and nitrogen stable isotope analysis (SIA). Mesozooplankton and individuals of sardines and anchovies were collected during one season (spring 2010), over spatially contrasted stations within the study area. First, the potential effect of preservation (ethanol vs. freezing) and of delipidation (by cyclohexane) on mesozooplankton δ¹³C and δ¹⁵N values was assessed. Results demonstrated the necessity to correct for the preservation effect and for lipid contents in mesozooplankton for further analyses of sardines' and anchovies' diet through SIA. Next, this study highlighted the interest of working on identified mesozooplanktonic organisms instead of undetermined assemblages when unravelling food sources of planktivorous fish using stable isotopes. The inter-specific variability of isotope values within a planktonic assemblage was effectively high, probably depending on the various feeding behaviours that can occur among mesozooplankton species. Intra-specific variability was also significant and related to the spatial variations of baseline signatures in the area. To investigate the foraging areas and potential diet overlap of S. pilchardus and E. encrasicolus, mixing models (SIAR) were applied. Both fish species appeared to feed mainly in the neritic waters of the Bay of Biscay in spring and to select mainly small- to medium-sized copepods (e.g. Acartia sp., Temora sp.). However, E. encrasicolus showed a greater trophic plasticity by foraging more offshore and on a wider range of prey sizes, while S. pilchardus seemed more limited to coastal areas and the mesozooplanktonic species of these waters for feeding.     

δC and δN biogeographic trends in rocky intertidal communities along the coast of South Africa: Evidence of strong environmental signatures

Ecosystem dynamics driven by top-down controls have been well documented in rocky intertidal communities, while the effects of bottom-up influences are comparatively poorly understood. We hypothesized that large-scale signatures of the physical environment may be identifiable along the South African coastline as it is subject to two very different current systems (Benguela and Agulhas Currents) that profoundly influence primary production and thus both food type and availability. Through stable isotope analysis, we examined biogeographic patterns in multiple trophic levels at four sites along a 1400-km stretch of South African coastline and investigated the dietary role of macroalgal-derived organic carbon in rocky intertidal communities. The general positioning of trophic groups was comparable across all sites, with animals from the same trophic levels grouping together and with a δ¹⁵N fractionation of 1–2‰ between levels. The species found at all sites demonstrated east–west δ¹⁵N enrichment, presumably reflecting a biogeographic shift in nitrogen sources linked to upwelling on the west coast. Filter-feeders gave particularly clear results. Using discriminant analysis, mussels could be categorized into four geographic groups based on carbon and nitrogen signatures: east coast, southeast coast, south-west coast and west coast. Barnacles and polychaetes showed similar geographic groupings to mussels, but with shifts in actual values (1‰ depletion in δ¹³C and 3‰ enrichment in δ¹⁵N relative to mussels). This suggests that fractionation varies between species within a trophic level.     

Functional changes due to invasive species: Food web shifts at shallow Posidonia oceanica seagrass beds colonized by the alien macroalga Caulerpa racemosa

Multiple stable isotope analyses were used to examine the trophic shifts at faunal assemblages within the invading macroalga Caulerpa racemosa in comparison to established communities of Posidonia oceanica seagrass meadows. Sampling of macrobenthic invertebrates and their potential food sources of algal mats and seagrass meadows in Mallorca (NW Mediterranean) showed differences in species composition of faunal and primary producers among seagrass and C. racemosa. Accordingly, changes in food web structure and trophic guilds were observed, not only at species level but also at community level. The carbon and nitrogen isotope signatures of herbivores, detritivores and deposit feeders confirmed that the seagrass provided a small contribution to the macrofaunal organisms. δ¹³C at the P. oceanica seagrass and at the C. racemosa assemblages differed, ranging from −6.19 to −21.20‰ and −2.67 to −31.41‰, respectively. δ¹⁵N at the Caulerpa mats was lower (ranging from 2.64 to 10.45‰) than that at the seagrass meadows (3.51–12. 94‰). Significant differences in isotopic signatures and trophic level among trophic guilds at P. oceanica and C. racemosa were found. N fractionation at trophic guild level considerable differed between seagrass and macroalgae mats, especially for detritivores, deposit feeders, and herbivores. Filter feeders slightly differed with a relatively lower N signal at the seagrass and CR values at community level and at trophic guild level were higher in the C. racemosa invaded habitats indicating an increase in diversity of basal resource pools. C. racemosa did seem to broaden the niche diversity of the P. oceanica meadows it colonised at the base of the food web, may be due to the establishment of a new basal resource. The extent of the effects of invasive species on ecosystem functioning is a fundamental issue in conservation ecology. The observed changes in invertebrate and macrophytic composition, stable isotope signatures of concomitant species and consequent trophic guild and niche breadth shifts at invaded Caulerpa beds increase our understanding of the seagrass systems.     

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.     

Isotopic variation (δN, δC,and δS) with body size in post-larval estuarine consumers

Stable isotope analyses were conducted on fish and crabs in Apalachicola Bay, Florida (USA) to determine whether δ¹⁵N values are correlated with length within these species. Our objective was to define the smallest trophic unit of the system as a prelude to further isotopic evaluations of consumer trophic level in Apalachicola Bay. In some cases, the smallest trophic unit is the individual species; however, previous gut content analysis on organisms in this system indicated that subdivision by size class within species might be required. However, over the size ranges observed, there was no systematic change in the δ¹⁵N values of individual consumers. Analysis of the δ¹³C and δ³⁴S values of these species reveal no obvious shifts in the utilization of organic matter sources with length that could be obscuring a trend in the δ¹⁵N signal. We find no isotopic evidence for a systematic trophic level increase within the species examined over the size ranges observed in this estuary. These results indicate that species rather than size class is the best predictor of trophic habit among these Apalachicola Bay consumers.     

Variation in the mobilization of mercury into Black-winged Stilt Himantopus himantopus chicks in coastal saltpans,as revealed by stable isotopes

Causes of variation in mobilization of mercury into Black-winged Stilt Himantopus himantopus chicks were studied through analysis of stable isotope ratios of carbon and nitrogen. Blood and breast feathers were collected from chicks in coastal saltpans during successive breeding seasons. Detritus samples and potential prey (macroinvertebrates) were also collected. Total mercury concentrations and stable isotope signatures were measured using atomic absorption spectroscopy and isotope ratio mass spectrometry respectively. Mercury levels in Chironomidae, Corixidae and Hydrophilidae correlated with mercury levels in chick feathers. Differences of δ¹⁵N signatures between macroinvertebrate groups indicated that they belong to different trophic levels. δ¹⁵N signatures of invertebrates correlated with mercury levels in invertebrates and chicks, but not with δ¹⁵N signatures in chicks. Between-group and between-site differences of δ¹⁵N signatures and mercury levels in invertebrates suggested that they contribute differently to mercury mobilization into chicks, and their relative contribution depends on prey availability in each site. Inter-site differences in the biomagnification factor reinforced that idea. δ¹³C signatures in invertebrates marked a larger range of carbon sources than just detritus. Variation of water inflow regime and prey availability may cause between-group and between-site differences of δ¹³C signatures in prey. Discrepancies between feather and blood for δ¹³C signatures in Praias-Sado and Vaia suggested that temporal variation of prey availability may be the main factor affecting mercury mobilization into chicks in both those cases, since their water inflow regimes are the same. The lowest levels of δ¹³C signatures in Vau suggested that water inflow regime may be the main factor in this case, since no discrepancy existed in δ¹³C signatures between blood and feather.     

Stable isotope composition of modern bryozoan skeletal carbonate from the Otago Shelf,New Zealand

The oxygen (δ¹⁸O) and carbon (δ¹³C) isotope ratios of 10 species of living Bryozoa collected from the Otago Shelf, New Zealand were analysed to assess the extent to which isotopic equilibrium (relative to inorganic equilibrium isotope fractionation) is attained during the precipitation of skeletal calcium carbonate. The data reveal that whereas eight species of Bryozoa synthesise skeletal carbonate in apparent oxygen isotope equilibrium with respect to environmental conditions, two species (Celleporina grandis and Hippomonavella flexuosa) yield δ¹⁸O_calcite values which indicate significant disequilibrium oxygen isotope fractionation during calcification. Sufficient data are available from one species (C. grandis) to demonstrate that disequilibrium is probably related to kinetic factors associated with diffusion‐controlled transport of HCO₃‐ to the site of calcite precipitation. Carbon isotope signatures indicate significant departures from inorganic isotope equilibrium in all but one bryozoan species (Hippomenella vellicata). Although greater uncertainties are associated with estimates of the isotopic composition of total dissolved inorganic carbon (δ¹³C_SDIC), the data suggest that two factors—kinetic fractionation and incorporation of respiratory CO₂—are important in controlling carbon isotope disequilibrium. Where bryozoan species exhibit evidence for disequilibrium in both oxygen and carbon isotope systems (C. grandis, H. flexuosa), it is likely that kinetic factors are primarily responsible for observed departures from carbon isotope equilibrium. In contrast, the probable explanation for those species which display evidence for carbon isotope disequilibrium only, is that skeletal carbonate is precipitated from a DIC pool modified by the incorporation of respiratory CO₂. Differences between the carbon isotope composition of skeletal elements from the same species and co‐existing species living in the same community suggests that significant variations may occur in the extent to which marine DIC and respiratory CO₂ are utilised during calcification. Additional studies of carbon pathways associated with calcification are required to assess the relative effects of kinetic, metabolic, and environmental factors on the carbon isotopic composition of bryozoan skeletal carbonate.     

Stable isotope ratios in Cape gannets around the southern coasts of Africa reveal penetration of biogeographic patterns in oceanic signatures

The southern coasts of Africa are influenced by two major oceanic currents, leading to biogeographic patterns in inshore and offshore species assemblages, and in the stable isotope signatures of suspended particulate matter and filter-feeding mussels. We used the stable isotope ratios of carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) from the blood and feathers of adult and chick Cape gannets (Morus capensis) to investigate whether the geographic differences observed at the lower levels in the marine communities are deep penetrating effects that reach top predators. Additionally, we evaluated whether trophic segregation occurs between adult and reared chick gannets, and whether a shift to wintering habitat occurs in adults. The study was conducted during the 2006 breeding season on Bird Island in the Agulhas system, and on Malgas and Ichaboe Islands, in the south and north Benguela respectively. Our results showed significant differences in the isotope ratios of members of different colonies, but no intra-colony differences between tissues or age groups. These results indicate that there is neither age-related nor temporal segregation in the diet of members of the same colony. Feather isotopic values suggest that adults remain all year round in the same habitats, and do not undertake long migration after reproduction. Since all gannets tend to target similar prey, we attributed among-colony differences in isotope signatures mostly to the oceanic conditions experienced by the main prey of birds rather than substantial differences in diet composition. Overall, isotopic signatures segregate the two current systems, with depleted carbon values in the Agulhas and enriched nitrogen values in the upwelled waters of the Benguela. Within the Benguela birds from Ichaboe in the north had higher δ¹⁵N values than those from Malgas in the south, which we attributed to differences in the functioning of the upwelling cells in the vicinity of the two colonies. Finally, slight variation in the proportion of main prey and discards from fisheries may contribute to the variation in the stable-isotope signatures between colonies in the Benguela.     

Distinctive stable isotope ratios in important zooplankton species in relation to estuarine salinity gradients: Potential tracer of fish migration

To assess the potential of stable isotope ratios as an indicator of fish migration within estuaries, stable isotope ratios in important zooplankton species were analyzed in relation to estuarine salinity gradients. Gut contents from migratory juveniles of the euryhaline marine fish Lateolabrax japonicus were examined along the Chikugo River estuary of the Ariake Sea, which has the most developed estuarine turbidity maximum (ETM) in Japan. Early juveniles in March and April preyed primarily on two copepod species; Sinocalanus sinensis at lower salinities and Acartia omorii at higher salinities. Late juveniles (standard length > 40 mm) at lower salinities preyed exclusively on the mysid Acanthomysis longirostris until July and complementarily on the decapod Acetes japonicus in August. These prey species were collected along the estuary during the spring–summer seasons of 2003 and 2004, and their carbon and nitrogen stable isotope ratios (δ¹³C and δ¹⁵N) were evaluated. The δ¹³C values of prey species were distinct from each other and were primarily depleted within and in close proximity to the ETM (salinity < 10); S. sinensis (−26.6‰) < Acanthomysis longirostris (−23.3‰) < Acartia omorii (−21.1‰) < Acetes japonicus (−18.5‰). The overall gradient of δ¹³C with salinity occurred for all prey species and showed minor temporal fluctuations, while it was not directly influenced by the δ¹³C values in particulate organic matter along the estuary. In contrast to δ¹³C, the δ¹⁵N values of prey species did not exhibit any clear relationship with salinity. The present study demonstrated that δ¹³C has the potential for application as a tracer of fish migration into lower salinity areas including the ETM.     

Trophic functioning of the St. Lucia estuarine lake during a drought phase assessed using stable isotopes

The St. Lucia Estuary is Africa’s largest estuarine system and is currently experiencing the stress of prolonged freshwater deprivation, manifested by extremely low water levels and hypersalinity. These unprecedented conditions have raised questions regarding the trophic functioning of the ecosystem. Despite the substantial amount of research previously undertaken within this system, no studies of food web structure and function have yet been documented. This study therefore aimed to examine the food web structure of the St. Lucia estuary system through the use of carbon and nitrogen stable isotope analysis. Analysis of carbon isotope ratios indicates that benthic carbon sources are most utilised at sites with low water levels and generally higher salinity (Catalina Bay, Charter’s Creek). Conversely, the estuarine region of the mouth and Narrows, with its elevated water levels and lower salinity, still sustains a viable pelagic food web. Analysis of δ¹⁵N ratios indicates that the number of trophic transfers (food chain length) might be related to water levels. Overall, the study provides a greater understanding of the ecological processes of this complex estuarine lake, which may allow for future comparisons of trophic functioning under drought and normal/wet conditions to be made.     

Shifts in an epibenthic trophic web across a marine frontal area in the Southwestern Atlantic (Argentina)

Marine benthic trophic relationships and food web structures may be influenced by benthic–pelagic coupling processes, which could also be intensified by the physical dynamics of marine fronts. In this work, we employed stable isotope (δ¹³C and δ¹⁵N) analysis to investigate the influence of the Southwest (SW) Atlantic shelf-break front (SBF; 38–39°S, 55–56°W; Argentina) on an epibenthic trophic web. Epibenthic organisms were sampled, at depths of ~ 100 m, with a non-selective dredge from a sandy bottom community located in frontal (F) and marginal (M) areas. The SBF position and the chlorophyll-a (chl-a) concentrations were inferred using satellite data of the sea surface temperature (SST) and satellite chl-a concentration, respectively. The most noticeable shifts in stable isotopes between the sampled areas were those of the Patagonian scallop, Zygochlamys patagonica (δ¹³C), and those of the sea urchin, Sterechinus agassizi (δ¹⁵N). Diet analyses inferred from stable isotopes and mixing models demonstrated that the dominant component of this community, Z. patagonica, had variable contributions to higher trophic levels between areas. More importantly, the epibenthic assemblage in F areas showed δ¹³C-enriched and δ¹⁵N-depleted isotopic signatures with respect to the M areas. Collectively, this evidence suggests that frontal dynamics promotes the accumulation of δ¹³C-enriched phytoplankton in the seabed in F areas, while in M areas the more degraded organic matter becomes more important in the trophic web, decreasing the δ¹⁵N isotopic signature of the assemblage. Therefore, the trophic web was sustained by fresher food in F areas than in M areas, demonstrating the role of frontal dynamics in the shaping of these communities.     

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.     

South Atlantic and benthic foraminifer δ13C deviations: implications for reconstructing the Late Quaternary deep-water circulation

Benthic foraminifer species Cibicidoides wuellerstorfi and related genera are assumed to secrete calcite very close to the carbon isotope values of the ambient bottom-water ΣCO₂. Recently, attention has been focused on substantial productivity-linked δ¹³C depletions. To examine further the productivity effect on benthic δ¹³C deviations, we present data from the South Atlantic between 15 and 35°S, including water samples from 10 hydrographic stations and related surface-sediment measurements on C. wuellerstorfi. We compare open-ocean data with observations in the Namibia Upwelling area. As a result, δ¹³C_ΣCO2 values as well as phosphate concentrations in water samples of the upwelling realm differ significantly from those of the open-ocean realm at least in the upper and mid-depth water masses (SACW, AAIW, UCDW). However, deviations from the Redfield fractionation, caused by air–sea fractionation, remain constant within each water mass, which means that the carbon isotope changes toward upwelling areas are exclusively determined by biological cycling. In addition to lower δ¹³C_ΣCO2 values in upwelling areas, a depletion in the δ¹³C of epibenthic foraminifer calcite is observed, which is most likely explained by the decay of organic matter, reducing the ¹³C/¹²C ratio in the pore water and influencing the carbon isotopic composition of the C. wuellerstorfi shells of highly productive areas. The paleoceanographic implication of this effect for reconstructing the Late Quaternary deep-water circulation is discussed using carbon isotope records of several sediment cores within and outside the Namibia upwelling area.     

Origin and fate of particulate organic matter in the southern Beaufort Sea – Amundsen Gulf region,Canadian Arctic

To establish the relative importance of terrigenous and marine organic matter in the southern Beaufort Sea, we measured the concentrations and the stable isotopic compositions of organic carbon and total nitrogen in sediments and in settling particles intercepted by sediment traps. The organic carbon content of surface sediment in the Chukchi and southern Beaufort Seas ranged from 0.6 to 1.6% dry wt., without a clear geographical pattern. The C_ORG:N_TOT ratio ranged from 7.0 to 10.4 and did not vary significantly downcore at any one station. Values of δ¹³C_ORG and δ¹⁵N_TOT in the sediment samples were strongly correlated, with the highest values, indicative of a more marine contribution, in the Amundsen Gulf. In contrast, the organic matter content, elemental (C_ORG:N_TOT ratio) and isotopic (δ¹³C_ORG and δ¹⁵N_TOT) composition of the settling particles was different from and much more variable than in the bottom sediments. The isotopic signature of organic matter in the Beaufort Sea is well constrained by three distinct end-members: a labile marine component produced in situ by planktonic organisms, a refractory marine component, the end product of respiration and diagenesis, and a refractory terrigenous component. A three-component mixing model explains the scatter observed in the stable isotope signatures of the sediment trap samples and accommodates an apparent two-component mixing model of the organic matter in sediments. The suspended matter in the water column contains organic matter varying from essentially labile and marine to mostly refractory and terrigenous. As it settles through the water column, the labile marine organic matter is degraded, and its original stable isotope signature changes towards the signature of the marine refractory component. This process continues in the bottom sediment with the result that the sedimentary organic matter becomes dominated by the refractory terrigenous and marine components.     

Variability in the fractionation of stable isotopes during degradation of two intertidal red algae

Macroalgae contribute to intertidal food webs primarily as detritus, with unclear implications for food web studies using stable isotope analysis. We examined differences in the thallus parts of two South African rhodophytes (Gelidium pristoides and Hypnea spicifera) and changes in overall δ¹³C, δ¹⁵N signatures and C:N ratios during degradation in both the field and laboratory. We hypothesized that both degrading macroalgal tissue and macroalgal-derived suspended particulate material (SPM) would show negligible changes in δ¹³C, but enriched δ¹⁵N signatures and lower C:N ratios relative to healthy plants. Only C:N laboratory ratios conformed to predictions, with both species of macroalgae showing decomposition related changes in δ¹³C and significant depletions in δ¹⁵N in both the field and laboratory. In the laboratory, algal tissue and SPM from each species behaved similarly (though some effects were non-significant) but with differing strengths. Gelidium pristoides δ¹³C increased and C:N ratios decreased over time in tissue and SPM; δ¹⁵N became depleted only in SPM. Hypnea spicifera, δ¹³C, δ¹⁵N and C:N ratios all decreased during degradation in both SPM and algae.     

Seasonal food web structures and sympagic-pelagic coupling in the European Arctic revealed by stable isotopes and a two-source food web model

We simultaneously followed stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes in a two-source food web model to determine trophic levels and the relative importance of open water- and ice-associated food sources (phytoplankton vs. ice algae) in the lower marine food web in the European Arctic during four seasons. The model is based upon extensive seasonal data from 1995 to 2001.Phytoplankton, represented by samples of particulate organic matter from open water (Pelagic-POM) and ice algae, represented by samples from the underside of the ice (Ice-POM), were isotopically different. Ice-POM was generally dominated by the typical ice diatoms Nitzschia frigida and Melosira arctica and was more enriched than Pelagic-POM in ¹³C (δ¹³C = −20‰ vs. −24‰), but less enriched in ¹⁵N (δ¹⁵N = 1.8‰ vs. 4.0‰). However, when dominated by pelagic algae, Ice-POM was enriched in ¹³C and ¹⁵N similarly to Pelagic-POM.The derived trophic enrichment factors for δ¹⁵N (Δ_N = 3.4‰) and δ¹³C (Δ_C = 0.6‰) were similar in both pelagic and sympagic (ice-associated) systems, although the Δ_C for the sympagic system was variable.Trophic level (TL) range for zooplankton (TL = 1.8-3.8) was similar to that of ice fauna (TL = 1.9-3.7), but ice amphipods were generally less enriched in δ¹⁵N than zooplankton, reflecting lower δ¹⁵N in Ice-POM compared to Pelagic-POM. For bulk zooplankton, TLs and carbon sources changed little seasonally, but the proportion of herbivores was higher during May-September than in October and March. Overall, we found that the primary carbon source for zooplankton was Pelagic-POM (mean 74%), but depending on species, season and TL, substantial carbon (up to 50%) was supplied from the sympagic system. For bulk ice fauna, no major changes were found in TLs or carbon sources from summer to autumn. The primary carbon source for ice fauna was Ice-POM (mean 67%), although ice fauna with TL > 3 (adult Onisimus nanseni and juvenile polar cod) primarily utilized a pelagic food source.     

Nekton migration and feeding location in a coastal area – A stable isotope approach

Stable isotope analysis was used to investigate nekton movements and feeding location in a coastal area adjacent to a major European river, the Tagus, Portugal. Particulate organic matter isotopic signatures presented a gradient from the river towards the sea. Phytoplankton, zooplankton, polychaetes and the crab, Polybius henslowii, provided evidence of the incorporation of terrestrial organic matter into the lower levels of the food web, reflecting local isotopic signatures. Two fish species reflected the coastal isotopic gradient in δ¹³C, Diplodus vulgaris and Arnoglossus imperialis and the latter also presented isotopic differerences among the sites for δ¹⁵N. Alloteuthis subulata, Trisopterus luscus and Callionymus lyra were isotopicaly distinct among sites for δ¹⁵N. An increase of δ¹⁵N with length was detected for T. luscus and C. lyra, possibly showing ontogenic trophic level changes. Since A. subulata did not present differences in length and still showed isotopic distinction for δ¹⁵N, among areas, it was concluded that local biogeochemical factors may also have an influence. Diplodus bellottii and Dicologlossa cuneata did not reflect any isotopic signature reflecting their wide migration and feeding across the coastal area. Central isotopic ranges, defined as the site mean values for δ¹³C and δ¹⁵N ± 1‰ were determined for each species and site and those deviating from these were considered transient individuals. Central isotopic ranges accounted for 87% of A. imperialis, 80% of A. subulata, 77% of T. luscus, 67% of C. lyra and 50% of D. vulgaris. The number of individuals within each central isotopic range was surprisingly high for an open coastal area and comparable to those of more structured environments.     

Seasonal variability of δ15N in sinking particles in the Benguela upwelling region

Temporal changes in δ¹⁵N values of sinking particles collected with sediment traps in the Benguela upwelling regime off southwest Africa mirrored variations in the input of inorganic nitrogen to the surface water. Reductions in δ¹⁵N (to as low as 2.5‰) corresponded to low sea surface temperatures during austral spring and late austral autumn/early winter, indicating increased nitrate availability due to the presence of recently upwelled water. High particulate fluxes accompanied the low δ¹⁵N values and sea surface temperatures, reflecting increased productivity, fueled by the upwelled nutrients. High δ¹⁵N values (up to 13.1‰) coincided with high sea surface temperatures and low particle fluxes. In this area, the seaward extension of upwelling filaments, which usually occurs twice yearly, brings nutrient-rich water to the euphotic zone and leads to elevated productivity and relatively lower δ¹⁵N values of the particulate nitrogen. Satellite images of ocean chlorophyll show that productivity variations coincide with δ¹⁵N changes. The observed isotopic pattern does not appear to have been caused by variations in the species composition of the phytoplankton assemblage. Calculations based on δ¹⁵N of the sinking particulate nitrogen show that the surface nitrate pool was more depleted during late austral summer/early fall and mid-winter and that supply exceeded demand during the intense spring bloom and in late austral fall. The main uncertainty associated with these estimates is the effect of diagenesis on δ¹⁵N and possible variability in preservation of the isotope signal between periods of high and low particle flux.     

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.