Spatial organization of food webs along habitat gradients at deep-sea hydrothermal vents on Axial Volcano,Northeast Pacific

Deep-sea hydrothermal vents are characterized by steep spatial gradients and high temporal variability in habitat conditions. This leads to the organization of species distribution along spatial habitat gradients, which may constrain food resource utilization and food web structure. We conducted a stable-isotope-based study to test the hypothesis that food resource utilization is constrained by spatial habitat variability at diffuse hydrothermal vents on Axial Volcano, Northeast Pacific. Our study included the ten most biomass-prominent species and considered the temporal change in food web structure at recently created vent sites during three consecutive years. We related species average stable isotopic composition to their position between the center and the periphery of vent sites, using previously published data. Species spread widely along the δ¹³C axis, and showed a small variability in δ¹⁵N. This indicates that most species partition food resources between isotopically different carbon sources, and that they are not organized along predator–prey trophic chains. Particulate organic matter (POM) stable isotopic composition from a concomitant study corresponds to the signature of the expected diet for most organisms. Species average δ¹³C was significantly correlated to their relative position between the center and the periphery of vent sites. We relate this spatial variability in species isotopic composition to variability in the isotopic signature of both dissolved inorganic carbon (DIC) and POM. This spatial isotopic signal of consumers reveals the spatial structuring of food (POM) production and its consumption by the fauna. Accrual of species during the development of diffuse sites increased the inter-specific spread in δ¹³C, but did not increase the range in δ¹⁵N. Our results show that the spatial organization of species distribution results in a fragmented food web where species partition POM food resources according to their position in space. Shaping of species distribution by habitat gradients therefore constrains food web structure and the occurrence of predator–prey and competitive interactions.     

Trophic ecology of the rocky shore community associated with the Ascophyllum nodosum zone (Roscoff,France): A δC vs δN investigation

This study aimed to characterize the structure and functioning of the benthic food web associated with the Ascophyllum nodosum zone of the rocky shore of Roscoff by using δ¹³C and δ¹⁵N. Several characteristics of the trophic ecology of the invertebrates associated with this mid-littoral habitat and which belong to different functional groups (e.g., grazers, filter-feeders, predators and omnivores) were highlighted. In particular, the filter feeder species (including mostly sponges) used macroalgae-derived organic matter as a substantial food requirement. The results also pointed out an important stable isotopes variability for strict coexisting primary consumers which: (1) is directly related to the high δ¹⁵N range of the food sources; (2) makes it impossible to establish a unique trophic level scale based on δ¹⁵N values, as previously done in coastal environments; and (3) points out the existence of major co-occurring trophic pathways which characterise the Ascophyllum nodosum habitat.     

Utilization of carbon sources in a northern Brazilian mangrove ecosystem

Carbon and nitrogen stable isotope ratios (¹³C and ¹⁵N) and trophic level (TL) estimates based on stomach content analysis and published data were used to assess the contribution of autotrophic sources to 55 consumers in an intertidal mangrove creek of the Curuçá estuary, northern Brazil. Primary producers showed δ¹³C signatures ranging between −29.2 and −19.5‰ and δ¹⁵N from 3.0 to 6.3‰. The wide range of the isotopic composition of carbon of consumers (−28.6 to −17.1‰) indicated that different autotrophic sources are important in the intertidal mangrove food webs. Food web segregation structures the ecosystem into three relatively distinct food webs: (i) mangrove food web, where vascular plants contribute directly or indirectly via POM to the most ¹³C-depleted consumers (e.g. Ucides cordatus and zooplanktivorous food chains); (ii) algal food web, where benthic algae are eaten directly by consumers (e.g. Uca maracoani, mullets, polychaetes, several fishes); (iii) mixed food web where the consumers use the carbon from different primary sources (mainly benthivorous fishes). An IsoError mixing model was used to determine the contributions of primary sources to consumers, based on δ¹³C values. Model outputs were very sensitive to the magnitude of trophic isotope fractionation and to the variability in ¹³C data. Nevertheless, the simplification of the system by a priori aggregation of primary producers allowed interpretable results for several taxa, revealing the segregation into different food webs.     

Pathways of organic matter through food webs of diverse habitats in the regulated Nakdong River estuary (Korea)

The benthic macroinvertebrates of the Nakdong River estuary were sampled at three different habitats: two salt marsh (Scirpus triqueter and Phragmites australis) beds and a bare intertidal flat. Fishes were sampled in the main channel. The trophic importance of marsh vascular plants, microphytobenthos, and riverine and channel particulate organic matter to macroinvertebrate and fish production was studied using stable carbon and nitrogen isotope tracers. There was a dramatic change in coverage of macrophytes (salt marshes and seagrass) after the construction of an estuarine barrage in 1987 in the Nakdong River estuary, with the S. triqueter bed increasing, the P. australis bed decreasing, and Zostera marina habitats being nearly lost. Although the invertebrate δ¹³C were within a narrower range than those of the primary producers, the values varied considerably among consumers in these habitats. However, the isotope signatures of consumers showed similarities among different habitats. Cluster analysis based on their isotopic similarity suggested that the isotope variability among species was related more to functional feeding groups than to habitats or taxonomic groups. While δ¹³C values of suspension feeders were close to that of the channel POM (mainly phytoplankton), other benthic feeders and predators had δ¹³C similar to that of microphytobenthos. Isotopic mixing model estimates suggest that algal sources, including microphytobenthos and phytoplankton, play an important role in supporting the benthic food web. Despite the huge productivity of emergent salt marshes, the contribution of the marsh-derived organic matter to the estuarine food webs appears to be limited to some nutrition for some invertebrates just within marsh habitats, with little on the bare intertidal flats or in the channel fish communities. Isotope signatures of the channel fishes also confirm that algal sources are important in supporting fish nutrition. Our findings suggest that benthic and pelagic microalgae made a large contribution to consumer diets, while marsh plants may not have a large role in supporting food webs in this estuarine system.     

Tracing carbon transfer and assimilation by invertebrates and fish across a tropical mangrove ecosystem using stable isotopes

Mangroves are highly productive ecosystems that exhibit a diverse range of habitats, including tidal creeks and flats, forest gaps and interior forest with varying understory light intensity, tidal dynamics, geomorphological settings, and overall biological production. Within mangrove ecosystems, invertebrates and fish feed on heterogeneous food sources, the occurrence of which is unevenly distributed across the system. This provides a basis for testing models of carbon transfer across mangrove ecosystems. We hypothesized that the carbon transfer and assimilation by fish and invertebrates will vary across the different mangrove habitats and that such variations can be predicted by their stable isotope compositions. We analysed δ¹³C and δ¹⁵N signatures of consumers and their potential organic carbon sources across a tropical mangrove ecosystem in Vietnam. The δ¹³C values of crabs and snails significantly decreased from the tidal flat to interior forest, indicating that variations in carbon transfer and assimilation occurred at small scales <30 m. Reduced variation in δ¹³C of suspension‐feeding bivalves suggested that tidal water was a vector for large‐scale transport of carbon across the mangrove ecosystem. An analysis of co‐variance using habitat as a fixed factor and feeding habit and movement capacity of consumers as co‐variates indicated that habitat and feeding types were major features that affected the δ¹³C values of invertebrates and fish. The findings demonstrate that carbon transfer and assimilation across mangrove ecosystems occur as a diverse combination of small (<30 m) and large (>30 m) scale processes.     

Field evidence of differential food utilization of phytal harpacticoids collected from Fucus serratus indicated by δC and δN stable isotopes

The present study is the first to compare trophic relationships of several co-occurring phytal harpacticoid species, in their natural habitat, using both δ¹³C and δ¹⁵N signatures. Three phytal harpacticoid species/taxa (Zaus spinatus, Tisbe spp., and Parathalestris cf. intermedia) all collected from the alga Fucus serratus, at different times of the year, were analyzed. The results indicated that the harpacticoids were utilizing food sources differently. Specific food sources of the three species/taxa could not be accurately pinpointed, but there were strong indications that F. serratus and fragments from it did contribute significantly to the diet of P. cf. intermedia and Tisbe spp. Both of these harpacticoid species overlapped in δ¹³C and δ¹⁵N values with some of the macrofaunal species, collected from the same site in Hvassahraun, Iceland, while no overlap was seen for Z. spinatus. The signatures for Z. spinatus indicated that its food sources changed seasonally.     

Intraspecific diet shift in Talitrus saltator inhabiting exposed sandy beaches

Talitrid amphipods are the most abundant herbivores on exposed sandy beaches. Despite their important role as trophic intermediates between macrophytes and higher levels (i.e. insect and bird) of beach food webs, very little information is available on their feeding patterns. The main aim of this study was to investigate intraspecific differences in the feeding behaviour of Talitrus saltator. We tested the hypotheses that: (1) adult females and males showed different isotope signatures and therefore relied on different sources of food; and (2) patterns of variation of isotope signatures of juveniles differed from those of adult specimens, evidencing a diet shift during the development. We used stable isotope signatures and tested for differences upon the level on the shore, times of the year and beaches experiencing similar morpho-dynamic and environmental conditions. Finally, we investigated the trophic significance of macrophyte detritus in the diet of males, females and juveniles. Results showed that adult males had a more variable diet than females and juveniles (inferred from δ¹³C and δ¹⁵N values). Dual-isotope graphs suggested that Sargassum muticum and Cystoseira baccata wrack could be among the main food sources for both juvenile and adult stage.     

Habitat heterogeneity influences cold‐seep macrofaunal communities within and among seeps along the Norwegian margin – Part 2: contribution of chemosynthesis and nutritional patterns

The relative contribution of chemosynthesis in heterotrophic fauna at seeps is known to be influenced by depth and by habitat. Using stable isotopes of carbon and nitrogen, we investigated macro‐ and megafaunal nutritional patterns in Norwegian margin cold seeps by comparing food webs both among habitats within a seep site and between different sites. The very active Håkon Mosby mud volcano (HMMV) is characterized by geochemical gradients, microbial activity and faunal zonation from the centre to the periphery. The Storegga Slide (600–900 m depth) has pockmarks with patchy less active seeps, and also shows concentric zonation of habitats but at much smaller spatial scale. The dominant carbon source for macrofaunal nutrition in both areas was chemosynthetically fixed and the bulk of organic carbon was derived from sulphur‐oxidizing bacteria. In HMMV, food chains were clearly separated according to habitats, with significantly lighter δ¹³C signatures on microbial mats and adjacent sediment (?33.06 to ?50.62‰) than in siboglinid fields (?19.83 to ?35.03‰). Mixing model outputs revealed that the contribution of methane‐derived carbon was small in siboglinid fields (0–17%) but significant (39–61%) in the microbial mats. Moreover, the variability of macrofauna signatures within this later habitat suggests the co‐occurrence of two food chains, one based on primary production via methanotrophy and the other via sulphide oxidation. The length of the food chains also varied among habitats, with at least one more trophic level in the siboglinid fields located at the periphery of the volcano. Conversely, in Storrega pockmarks, faunal δ¹³C signatures did not vary among habitats but among species, although separate food chains seem to co‐occur. The small size of the seepage areas and their lower fluxes compared to HMMV allow more background species to penetrate the seep area, increasing the range of δ¹⁵N and the trophic level number. Probably due to the higher flux of photosynthetic particulate organic carbon, the overall chemosynthesis‐based carbon contribution in invertebrate nutrition was lower than that in HMMV.     

Assessing waterbird habitat use in coastal evaporative systems using stable isotopes (δC, δN and δD) as environmental tracers

Isotopic patterns of biota across salinity gradients in man-made evaporative systems could assist in determining the use of these habitats by animals. Here we report δ¹³C, δ¹⁵N and δD measurements of a euryhaline fish, the Mediterranean toothcarp (Aphanius fasciatus), inhabiting a range of salinities in the Thyna saltworks near Sfax (Tunisia). The contribution of these salinity niches to egg formation of two typically piscivorous bird species breeding in the area and feeding within saltworks, Little Tern (Sternula albifrons) and Little Egret (Egretta garzetta), was inferred trough a triple-isotope (δ¹³C, δ¹⁵N and δD) Bayesian mixing model. Isotopic trends for fish δ¹⁵N and δD across the salinity gradient followed the equations: δ¹⁵N = e^{(1.1 + 47.68/Salinity)} and δD = −175.74 + Salinity + Salinity²; whereas fish δ¹³C increased as salinity rose (δ¹³C = −10.83 + 0.02·Salinity), after a sudden drop in fish isotopic values for salinities >60 (Practical Salinity Scale) (average fish δ¹³C for salinities <60 = −5.92‰). Both bird species fed largely on low hypersalinity ponds (salinity = 43; average contribution = 37% and 22% for Little Egrets and Little Terns, respectively), although the use of intermediate hypersalinities (salinities 63 and 70) by Little Terns also occurred (16% and 21%, respectively). Isotopic patterns across salinity gradients allow the use of isotopic measurements to inform studies of habitat occupancy within evaporative systems and provide further insights into how wildlife communities interact with them.     

基于稳定同位素技术的光背团水虱食性分析

为了探究光背团水虱的食性特征,本研究利用碳、氮稳定同位素技术于2015年冬季和2016年夏季对广西北海廉州湾红树林中光背团水虱及其食物来源的碳、氮稳定同位素比值（δ13C值和δ15N值）进行分析。结果显示,冬季和夏季光背团水虱的δ13C值大小范围为-22.85‰~-21.87‰,平均值为（-22.46±0.35）‰;δ15N值大小范围为11.02‰~12.85‰,平均值为（11.88±0.56）‰;光背团水虱的δ13C值、δ15N值变化范围较小,表明其食物来源较为简单。单因素方差分析结果显示,冬季与夏季光背团水虱的平均δ13C值差异不显著（P>0.05）,而夏季的δ15N值普遍高于冬季δ15N值,差异显著（P<0.05）;不同生长阶段的光背团水虱δ13C值、δ15N值会随着体长的增长而增大,差异显著（P<0.05）,表明光背团水虱在生长的过程中可能发生了食性转变。光背团水虱的δ13C值与浮游生物的δ13C值相近,而与红树植物δ13C值差距较远,说明光背团水虱主要以浮游生物为食物来源。基于R语言稳定同位素混合模型（SIAR）计算结果显示,冬季和夏季各粒径级别浮游生物对不同生长阶段的光背团水虱的贡献率趋势基本一致,表现为1.2~25 μm粒级的浮游生物对光背团水虱平均贡献率最高,其次为25~50 μm粒级,粒径大于100 μm的浮游生物对体长小于5.5 mm的光背团水虱贡献率较低,对体长大于5.5 mm的光背团水虱的贡献率随着体长增大而相应增大,说明不同生长阶段的光背团水虱食性有差异。对光背团水虱食性分析的结果可为深入研究团水虱爆发的原因及危害红树林的作用机理提供基础资料。     

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.     

Contrasted sulphide chemistries in the environment of 13°N EPR vent fauna

To improve our understanding of the environmental constraints exerted on vent fauna, we investigated sulphide chemistry in the habitats of Riftia pachyptila and Alvinella pompejana, at the Genesis and Elsa EPR 13°N sites. Temperature, pH and sulphide measurement series were acquired in situ, around the organisms, from the submersible Nautile. Hot fluid samples were also collected to evaluate end-member composition at these sites. Under the assumption of conservative mixing, pH, total sulphide concentration and sulphide speciation gradients in relation to temperature were modelled. From the comparison of measured and calculated concentrations, deviation from conservative behaviour was highlighted for total sulphide versus temperature. While the observed sulphide depletion around tubeworms suggests significant subsurface removal or biological consumption, the apparent sulphide enrichment in the alvinellid environment may reveal either conductive cooling of diffusing fluids or a secondary sulphide source. The calculated sulphide speciation appears to be contrasted at the two sites studied. Because of the low iron content in Genesis fluid, iron sulphide would not constitute a dominant sulphide species and the toxic H₂S form would be predominant in the mixing zone. By contrast, iron is expected to play a dominant role in sulphide speciation at the Elsa site where the end-member is iron rich. With respect to sulphide, the conditions encountered in the different habitats considered in this study are strongly contrasted. A low fluid flux was observed in the R. pachyptila habitat, contrasting with previous ideas, and suggests that sulphide availability could be a major limiting factor. Particularly, the bioavailable HS⁻ form is expected to vary weakly along the mixing gradient. In contrast, sulphide in the A. pompejana environment is shown to be particularly high, about one order of magnitude higher than observed for other Eastern Pacific alvinellids. At Genesis, because of the acidic pH and low iron conditions encountered, exposure to high levels of toxic sulphide is expected. A. pompejana thus appears to be particularly tolerant to such toxic conditions, but, as previously suggested, less severe conditions may also be found when iron is rich enough in the medium to dominate sulphide chemistry.     

A Stable Carbon Isotope Study of the Food-web in a Freshwater-deprived South African Estuary, with Particular Emphasis on the Ichthyofauna

The importance of macrophytes as food sources for estuarine nekton is unclear. Previous carbon isotope investigations in the macrophyte-dominated, freshwater-deprived Kariega Estuary showed that the bivalveSolen cylindraceusdid not utilize the dominant estuarine macrophytes found within the estuary as a primary food source. This finding prompted questions as to what the nekton of this estuary utilize as primary energy sources. δ¹³C analyses of the principal autochthonous and allochthonous primary carbon sources, as well as the dominant invertebrate and fish species, indicate that there are two main carbon pathways within the Kariega Estuary. The littoral community, which incorporates the majority of crustaceans, gobies, mullet and a sparid, utilizes δ¹³C enriched primary food sources namelySpartina maritima,Zosteracapensis and epiphytes. The channel fauna, which includes the zooplankton, zooplanktivorous and piscivorous fish, utilizes a primary food source depleted in δ¹³C, which is most likely a mixture of phytoplankton, terrestrial plant debris and C₄macrophyte detritus. The C₃saltmarsh macrophytesSarcocornia perennisandChenolea diffusa, as well as benthic microalgae, appear to be less important as primary food sources to the nekton of the Kariega Estuary.     

Spatial variability in the trophic ecology and biology of the deep-sea shrimp Aristaeomorpha foliacea in the Mediterranean Sea

The trophic ecology, energy and reproductive states of the deep-water shrimp Aristaeomorpha foliacea, widely distributed along the slopes of the Mediterranean Sea Basins, were analysed in eight areas spread along ca. 3000 km in order to identify patterns in the habitat conditions supporting the species. From W to E the areas were situated between the north side of Eivissa (39°12′N, 1°20′E, in the Balearic Basin) and off Mersin, Turkey (36°15′N, 34°19′E, in the Levantine Sea). Trends identified mainly as a function of longitude from west to east were: (i) higher δ¹⁵N, parallel to δ¹⁵N shifts in the top 200 m of the water column for particulate organic N (Pantoja et al., 2002). The δ¹⁵N trend indicates that the deep trophic web, i.e. A. foliacea at 400–600 m, reflects the δ¹⁵N signal of the photic zone; (ii) a similar significant trend of δ¹³C, related with exploitation of pelagic versus benthic resources by A. foliacea in each area (i.e. by local variability of terrigenous inputs via submarine canyons). More depleted δ¹³C was found at mid-longitudes (Tyrrhenian Sea and Sicily Channel) linked to higher consumption of macroplankton prey (Pasiphaea spp., euphausiids and mesopelagic fishes). The feeding intensity (gut fullness, F) and prey diversity (J) of A. foliacea were related, according to generalized linear models, with the temperature and salinity of intermediate waters, variables in turn associated with latitude and longitude. Both F and J were higher in areas with greater shrimp density. The optimal ecological habitat of A. foliacea appears to be located in the Tyrrhenian Sea and the Sicily Channel, where we found the highest F, the greatest trophic diversity and A. foliacea in the best biological condition (i.e. with higher hepato-somatic index, HSI). These are also the areas with the highest densities of A. foliacea. In contrast, in the western Mediterranean Sea (Balearic Basin and the southern Balearic Islands), where A. foliacea has low densities, the shrimp showed generally lower values of trophic indicators and biological condition.     

环境饵料丰度的季节变化对筼筜湖3种大型底栖动物食性的影响——来自稳定同位素的证据

测定了不同时期筼筜湖3种大型底栖动物(沙筛贝Mytilopsis sallei、日本大螯蜚Grandidierella japonica和腺带刺沙蚕Neanthes japonica)及其潜在食源的稳定同位素组成(δ13C和δ15 N),研究环境饵料丰度的季节变化对筼筜湖3种大型底栖动物食性的影响。结果显示,筼筜湖悬浮颗粒有机物(Particulate organic matter∶POM)的δ13C和δ15 N存在明显的季节变化。3月,受到输入筼筜湖的西海域海水大量陆源有机碎屑以及湖区周围的生活污水以及餐饮业输入的影响,筼筜湖POM的δ13 C和δ15 N明显贫化;而在9月,POM中δ13 C和δ15 N明显富集的内源性浮游植物的贡献增加。沙筛贝是典型的底栖滤食者,主要以POM为食,但它比POM富集的δ13 C值(3月和9月二者之间的Δδ13 C分别为2.9‰和1.6‰)表明它还摄入其他δ13 C相对富集的食物来源:石莼来源的有机碎屑可能是3月份沙筛贝δ13C富集的原因,而9月份则是由于再悬浮的底栖微藻对沙筛贝食源的贡献引起的。食碎屑的腺带刺沙蚕和日本大螯蜚在3月以石莼及其表面的附生生物为食,而9月份底栖微藻和浮游植物来源的POM是它们食源的主要贡献者。本研究的结果显示,筼筜湖3种大型底栖动物的δ15 N都出现明显的季节变化(Δδ15 N介于2.2‰~4.3‰),这是由于它们食源稳定同位素的季节性波动及其食性的季节变化引起的,而消费者食性的季节性变化则受到不同时期环境食物可利用性的影响。     

Effect of body size and body mass on δC and δN in coastal fishes and cephalopods

Carbon and nitrogen isotopes have been widely used in the investigation of trophic relations, energy pathways, trophic levels and migrations, under the assumption that δ¹³C is independent of body size and that variation in δ¹⁵N occurs exclusively due to ontogenetic changes in diet and not body size increase per se. However, several studies have shown that these assumptions are uncertain. Data from food-webs containing an important number of species lack theoretical support on these assumptions because very few species have been tested for δ¹³C and δ¹⁵N variation in captivity. However, if sampling comprises a wide range of body sizes from various species, the variation of δ¹³C and δ¹⁵N with body size can be investigated. While correlation between body size and δ¹³C and δ¹⁵N can be due to ontogenetic diet shifts, stability in such values throughout the size spectrum can be considered an indication that δ¹³C and δ¹⁵N in muscle tissues of such species is independent of body size within that size range, and thus the basic assumptions can be applied in the interpretation of such food webs. The present study investigated the variation in muscle δ¹³C and δ¹⁵N with body size and body mass of coastal fishes and cephalopods. It was concluded that muscle δ¹³C and δ¹⁵N did not vary with body size or mass for all bony fishes with only one exception, the dragonet Callionymus lyra. Muscle δ¹³C and δ¹⁵N also did not vary with body size or mass in cartilaginous fishes and cephalopods, meaning that body size/mass per se have no effect on δ¹³C or δ¹⁵N, for most species analysed and within the size ranges sampled. The assumption that δ¹³C is independent of body size and that variation in δ¹⁵N is not affected by body size increase per se was upheld for most organisms and can be applied to the coastal food web studied taking into account that C. lyra is an exception.     

Environmental correlates of large-scale spatial variation in the δ13C of marine animals

Carbon stable isotopes can be used to trace the sources of energy supporting food chains and to estimate the contribution of different sources to a consumer's diet. However, the δ¹³C signature of a consumer is not sufficient to infer source without an appropriate isotopic baseline, because there is no way to determine if differences in consumer δ¹³C reflect source changes or baseline variation. Describing isotopic baselines is a considerable challenge when applying stable isotope techniques at large spatial scales and/or to interconnected food chains in open marine environments. One approach is to use filter-feeding consumers to integrate the high frequency and small-scale variation in the isotopic signature of phytoplankton and provide a surrogate baseline, but it can be difficult to sample a single consumer species at large spatial scales owing to rarity and/or discontinuous distribution. Here, we use the isotopic signature of a widely distributed filter-feeder (the queen scallop Aequipecten opercularis) in the north-eastern Atlantic to develop a model linking base δ¹³C to environmental variables. Remarkably, a single variable model based on bottom temperature has good predictive power and predicts scallop δ¹³C with mean error of only 0.6‰ (3%). When the model was used to predict an isotopic baseline in parts of the overall study region where scallop were not consistently sampled, the model accounted for 76% and 79% of the large-scale spatial variability (10¹–10⁴ km) of the δ¹³C of two fish species (dab Limanda limanda and whiting Merlangus merlangius) and 44% of the δ¹³C variability in a mixed fish community. The results show that source studies would be significantly biased if a single baseline were applied to food webs at larger scales. Further, when baseline δ¹³C cannot be directly measured, a calculated baseline value can eliminate a large proportion of the unexplained variation in δ¹³C at higher trophic levels.     

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.