Adapt or die—Response of large herbivores to environmental changes in Europe during the Holocene

Climate warming and human landscape transformation during the Holocene resulted in environmental changes for wild animals. The last remnants of the European Pleistocene megafauna that survived into the Holocene were particularly vulnerable to changes in habitat. To track the response of habitat use and foraging of large herbivores to natural and anthropogenic changes in environmental conditions during the Holocene, we investigated carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope composition in bone collagen of moose (Alces alces), European bison (Bison bonasus) and aurochs (Bos primigenius) in Central and Eastern Europe. We found strong variations in isotope compositions in the studied species throughout the Holocene and diverse responses to changing environmental conditions. All three species showed significant changes in their δ¹³C values reflecting a shift of foraging habitats from more open in the Early and pre‐Neolithic Holocene to more forest during the Neolithic and Late Holocene. This shift was strongest in European bison, suggesting higher plasticity, more limited in moose, and the least in aurochs. Significant increases of δ¹⁵N values in European bison and moose are evidence of a diet change towards more grazing, but may also reflect increased nitrogen in soils following deglaciation and global temperature increases. Among the factors explaining the observed isotope variations were time (age of samples), longitude and elevation in European bison, and time, longitude and forest cover in aurochs. None of the analysed factors explained isotope variations in moose. Our results demonstrate the strong influence of natural (forest expansion) and anthropogenic (deforestation and human pressure) changes on the foraging ecology of large herbivores, with forests playing a major role as a refugial habitat since the Neolithic, particularly for European bison and aurochs. We propose that high flexibility in foraging strategy was the key for survival of large herbivores in the changing environmental conditions of the Holocene.     

Stable isotope records of sei whale baleens from Chilean Patagonia as archives for feeding and migration behavior

Carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope variations in baleen plates of sei whales (Balaenoptera borealis) stranded after a mass mortality event in Chilean Patagonia were investigated to assess potential dietary and migratory patterns. Carbon and nitrogen isotope ratios of seven baleens from six individuals were analyzed. The δ¹³C values ranged from ? 19.1 to ? 15.9‰ and the δ¹⁵N values from 8.7 to 15.4‰. Variations of up to 2.9‰ for δ¹³C and 5.3‰ for δ¹⁵N were observed within one baleen. Carbon and nitrogen isotope records of each baleen were significantly correlated and showed recurring oscillations confirmed by wavelet analyses. Oscillations slightly differed in periodicity indicating variable baleen growth rates between 10.0 and 16.5 cm/year. Food sources of the whales are discussed in terms of available isotope data for potential prey taxa and potential migratory behavior on the basis of latitudinal isotope gradients of particulate organic matter. Cyclicity could be explained by regular migrations of the sei whales from subtropical calving areas to high‐latitude foraging grounds. δ¹⁵N records of baleens differed between individuals eventually pointing to diverse feeding and migratory preferences among sei whale individuals.     

Foraging habits in a generalist predator: Sex and age influence habitat selection and resource use among bottlenose dolphins (Tursiops truncatus)

This study examines resource use (diet, habitat use, and trophic level) within and among demographic groups (males, females, and juveniles) of bottlenose dolphins (Tursiops truncatus). We analyzed the δ¹³C and δ¹⁵N values of 15 prey species constituting 84% of the species found in stomach contents. We used these data to establish a trophic enrichment factor (TEF) to inform dietary analysis using a Bayesian isotope mixing model. We document a TEF of 0‰ and 2.0‰ for δ¹³C and δ¹⁵N, respectively. The dietary results showed that all demographic groups relied heavily on low trophic level seagrass‐associated prey. Bayesian standard ellipse areas (SEA_b) were calculated to assess diversity in resource use. The SEA_b of females was nearly four times larger than that of males indicating varied resource use, likely a consequence of small home ranges and habitat specialization. Juveniles possessed an intermediate SEA_b, generally feeding at a lower trophic level compared to females, potentially an effect of natal philopatry and immature foraging skills. The small SEA_b of males reflects a high degree of specialization on seagrass associated prey. Patterns in resource use by the demographic groups are likely linked to differences in the relative importance of social and ecological factors.     

Changes in diets of individual Baltic ringed seals (Phoca hispida botnica) during their breeding season inferred from stable isotope analysis of multiple tissues

The stable isotope ratios (δ¹³C and δ¹⁵N) of three tissues with different metabolic rates (plasma, liver, and muscle) were used to investigate temporal variation in diet among nine individual Baltic ringed seals (Phoca hispida botnica Gmelin) from the Bothnian Bay, northeast Baltic Sea. The isotope values from plasma should reflect the most recent diet, values from liver the diet of the past weeks prior to sampling, and values from muscle should integrate diet over almost the entire breeding season of the ringed seals. In general, δ¹³C values of liver were more enriched in ¹³C than were those of either muscle or plasma, suggesting that the diet of the seals may have included a higher proportion of ¹³C‐enriched benthic prey in April. Females showed more variable δ¹³C values than males, suggesting possible gender differences in diet or in foraging locations. The differences that were apparent between females possibly reflect individual variation in the onset and duration of parturition and lactation, both of which likely restrict female foraging. Previous data from parasite infections and from alimentary tract contents of the same seals were linked to the isotope data to assist in drawing inferences about changes in the diets of individual seals.     

Carbon isotopes in otolith amino acids identify residency of juvenile snapper (Family: Lutjanidae) in coastal nurseries

This study explored the potential for otolith geochemistry in snapper (Family: Lutjanidae) to identify residency in juvenile nursery habitats with distinctive carbon isotope values. Conventional bulk otolith and muscle stable isotope analyses (SIA) and essential amino acid (AA) SIA were conducted on snapper collected from seagrass beds, mangroves, and coral reefs in the Red Sea, Caribbean Sea, and Pacific coast of Panama. While bulk stable isotope values in otoliths showed regional differences, they failed to distinguish nursery residence on local scales. Essential AA δ¹³C values in otoliths, on the other hand, varied as a function of habitat type and provided a better tracer of residence in different juvenile nursery habitats than conventional bulk otolith SIA alone. A strong linear relationship was found between paired otolith and muscle essential AA δ¹³C values regardless of species, geographic region, or habitat type, indicating that otolith AAs recorded the same dietary information as muscle AAs. Juvenile snapper in the Red Sea sheltered in mangroves but fed in seagrass beds, while snapper from the Caribbean Sea and Pacific coast of Panama showed greater reliance on mangrove-derived carbon. Furthermore, compound-specific SIA revealed that microbially recycled detrital carbon, not water-column-based new phytoplankton carbon, was the primary carbon source supporting snapper production on coastal reefs of the Red Sea. This study presented robust tracers of juvenile nursery residence that will be crucial for reconstructing ontogenetic migration patterns of fishes among coastal wetlands and coral reefs. This information is key to determining the importance of nursery habitats to coral reef fish populations and will provide valuable scientific support for the design of networked marine-protected areas.     

Individual dietary specialization in a generalist predator: A stable isotope analysis of urban and rural red foxes

Some carnivores are known to survive well in urban habitats, yet the underlying behavioral tactics are poorly understood. One likely explanation for the success in urban habitats might be that carnivores are generalist consumers. However, urban populations of carnivores could as well consist of specialist feeders. Here, we compared the isotopic specialization of red foxes in urban and rural environments, using both a population and an individual level perspective. We measured stable isotope ratios in increments of red fox whiskers and potential food sources. Our results reveal that red foxes have a broad isotopic dietary niche and a large variation in resource use. Despite this large variation, we found significant differences between the variance of the urban and rural population for δ¹³C as well as δ¹⁵N values, suggesting a habitat‐specific foraging behavior. Although urban regions are more heterogeneous regarding land cover (based on the Shannon index) than rural regions, the dietary range of urban foxes was smaller compared with that of rural conspecifics. Moreover, the higher δ¹³C values and lower δ¹⁵N values of urban foxes suggest a relatively high input of anthropogenic food sources. The diet of most individuals remained largely constant over a longer period. The low intraindividual variability of urban and rural red foxes suggests a relatively constant proportion of food items consumed by individuals. Urban and rural foxes utilized a small proportion of the potentially available isotopic dietary niche as indicated by the low within‐individual variation compared to the between‐individual variation. We conclude that generalist fox populations consist of individual food specialists in urban and rural populations at least over those periods covered by our study.     

Is individual consistency in body mass and reproductive decisions linked to individual specialization in foraging behavior in a long‐lived seabird?

Individual specialization in diet or foraging behavior within apparently generalist populations has been described for many species, especially in polar and temperate marine environments, where resource distribution is relatively predictable. It is unclear, however, whether and how increased environmental variability – and thus reduced predictability of resources – due to global climate change will affect individual specialization. We determined the within‐ and among‐individual components of the trophic niche and the within‐individual repeatability of δ¹³C and δ¹⁵N in feathers and red blood cells of individual female southern rockhopper penguins (Eudyptes chrysocome) across 7 years. We also investigated the effect of environmental variables (Southern Annular Mode, Southern Oscillation Index, and local sea surface temperature anomaly) on the isotopic values, as well as the link between stable isotopes and female body mass, clutch initiation dates, and total clutch mass. We observed consistent red blood cell δ¹³C and δ¹⁵N values within individuals among years, suggesting a moderate degree of within‐individual specialization in C and N during the prebreeding period. However, the total niche width was reduced and individual specialization not present during the premolt period. Despite significant interannual differences in isotope values of C and N and environmental conditions, none of the environmental variables were linked to stable isotope values and thus able to explain phenotypic plasticity. Furthermore, neither the within‐individual nor among‐individual effects of stable isotopes were found to be related to female body mass, clutch initiation date, or total clutch mass. In conclusion, our results emphasize that the degree of specialization within generalist populations can vary over the course of 1 year, even when being consistent within the same season across years. We were unable to confirm that environmental variability counteracts individual specialization in foraging behavior, as phenotypic plasticity in δ¹³C and δ¹⁵N was not linked to any of the environmental variables studied.     

Isotopic fractionation with morphological change and sexual specificity in the lappet moth Euthrix potatoria

The δ¹⁵N values of adult holometabolous insects exceed those of larvae, but otherwise little information on terrestrial invertebrates has been obtained in food‐web analyses using stable isotope ratios (δ¹⁵N, δ¹³C). Changes in δ¹³C during metamorphosis and differences between males and females have not been examined. We collected the larvae and cocoons of Euthrix potatoria (L.) (Lepidoptera: Lasiocampidae) in the field and used them to assess the species’ isotopic fractionation. Each emerged moth was divided into five body parts. We conducted stable N and C isotope analyses for each body part, as well as for cocoons and exuviae, and also compared stable isotope ratios between sexes. We confirmed δ¹⁵N enrichment through metamorphosis and estimated that δ¹⁵N enrichment is accomplished by the relative concentration of ¹⁵N due to the excretion of copious meconium, which contains abundant ¹⁴N. We also observed changes in δ¹³C values through metamorphosis. Both isotope values tended to change more in males than in females. The proportion of the whole‐adult weight represented by meconium was higher in males than in females, suggesting that high meconium secretion in males contributes to the sexual difference in δ¹⁵N. These phenomena may be common in Holometabola, which require a pupal stage. For more accurate food‐web assessments, it is important to consider stable isotope changes during different life cycles, as well as sexual differences.     

Regional movement patterns of a small‐bodied shark revealed by stable‐isotope analysis

This study used stable‐isotope analysis to define the nearshore regional residency and movements of the small‐bodied Australian sharpnose shark Rhizoprionodon taylori. Plasma and muscle δ¹³C and δ¹⁵N of R. taylori were collected from across five embayments and compared with values of seagrass and plankton from each bay. Linear distances between adjacent bays ranged from 30 to 150 km. There was a positive geographic correlation between R. taylori tissue and environmental δ¹³C values. Populations with the highest tissue δ¹⁵N were collected from bays that had the highest environmental δ¹⁵N values. These results suggest that R. taylori did not forage more than 100 km away from their capture location within 6 months to 1 year. The successful application of isotope analysis to define R. taylori movement demonstrates that this technique may be used in addition to traditional methods to study the movement of sharks, even within similar habitats across regionally small spatial scales (<100 km).     

Temporal and spatial trends in marine carbon isotopes in the Arctic Ocean and implications for food web studies

The Arctic is undergoing unprecedented environmental change. Rapid warming, decline in sea ice extent, increase in riverine input, ocean acidification and changes in primary productivity are creating a crucible for multiple concurrent environmental stressors, with unknown consequences for the entire arctic ecosystem. Here, we synthesized 30 years of data on the stable carbon isotope (δ¹³C) signatures in dissolved inorganic carbon (δ¹³C‐DIC; 1977–2014), marine and riverine particulate organic carbon (δ¹³C‐POC; 1986–2013) and tissues of marine mammals in the Arctic. δ¹³C values in consumers can change as a result of environmentally driven variation in the δ¹³C values at the base of the food web or alteration in the trophic structure, thus providing a method to assess the sensitivity of food webs to environmental change. Our synthesis reveals a spatially heterogeneous and temporally evolving δ¹³C baseline, with spatial gradients in the δ¹³C‐POC values between arctic shelves and arctic basins likely driven by differences in productivity and riverine and coastal influence. We report a decline in δ¹³C‐DIC values (?0.011‰ per year) in the Arctic, reflecting increasing anthropogenic carbon dioxide (CO₂) in the Arctic Ocean (i.e. Suess effect), which is larger than predicted. The larger decline in δ¹³C‐POC values and δ¹³C in arctic marine mammals reflects the anthropogenic CO₂ signal as well as the influence of a changing arctic environment. Combining the influence of changing sea ice conditions and isotopic fractionation by phytoplankton, we explain the decadal decline in δ¹³C‐POC values in the Arctic Ocean and partially explain the δ¹³C values in marine mammals with consideration of time‐varying integration of δ¹³C values. The response of the arctic ecosystem to ongoing environmental change is stronger than we would predict theoretically, which has tremendous implications for the study of food webs in the rapidly changing Arctic Ocean.     

New insights into the trophic ecology of the scalloped hammerhead shark,Sphyrna lewini,in the eastern tropical Pacific Ocean

Scalloped hammerhead shark, Sphyrna lewini, is considered a generalist predator, showing individual specializations and different trophic levels. This species inhabits both oceanic islands and coastal nursery areas in the eastern tropical Pacific (ETP) Ocean. Existing trophic ecology studies on S. lewini have been focused on stomach contents. This study provides new insights into its foraging preferences, habitat use, and trophic position, using stable isotope analysis (SIA) of muscle tissue. Stable isotope signatures of δ¹³C and δ¹⁵N were determined in S. lewini muscles (n?=?29) from the Ecuadorian Pacific in 2013. Trophic position (TP), isotopic niches, and overlap of S. lewini were estimated by sex, age, and maturity stages to infer their dietary habits throughout life stages. SIA revealed complex movement patterns related to sex and age classes of S. lewini, highlighting high degrees of dietary plasticity and habitat use, with a stronger relation to coastal regions than previously reported. This study provides crucial information regarding essential areas and the related migration behavior of S. lewini, with important implications for their conservation and management in the ETP.

     

Caribbean octocorals record changing carbon and nitrogen sources from 1862 to 2005

During the last century, the global biogeochemical cycles of carbon (C) and nitrogen (N) have been drastically altered by human activities. A century of land‐clearing and biomass burning, followed by fossil fuel combustion have increased the concentration of atmospheric CO₂ by approximately 20%, and since the mid‐1900s, the use of agricultural fertilizers has been the primary driver of an approximate 90% increase in bioavailable N. Geochemical records obtained through stable isotope analysis of terrestrial and marine biota effectively illustrate rising anthropogenic C inputs. However, there are fewer records of anthropogenic N, despite the enormous magnitude of change and the known negative effects of N on ecosystem health. We used stable isotope values from independent octocorals (gorgonians) sampled across the Western Atlantic over the last 143 years to document human perturbations of the marine C and N pools. Here, we demonstrate that in sea plumes δ¹³C values and in both sea plumes and sea fans δ¹⁵N values declined significantly from 1862 to 2005. Sea plume δ ¹³C values were negatively correlated with increasing atmospheric CO₂ concentrations and corroborate known rates of change resulting from global fossil fuel combustion, known as the Suess effect. We suggest that widespread input of agricultural fertilizers to near‐shore coastal waters is the dominant driver for the decreasing δ ¹⁵N trend, though multiple anthropogenic sources are likely affecting this trend. Given the interest in using δ ¹⁵N as an indicator for N pollution in aquatic systems, we highlight the risk of underestimating contributions of pollutants as a result of source mixing as demonstrated by a simple isotope‐mixing model. We conclude that signals of major human‐induced perturbations of the C and N pools are detectable in specimens collected over wide geographic scales, and that archived materials are invaluable for establishing baselines against which we can assess environmental change.     

Evaluating marine diets through radiocarbon dating and stable isotope analysis of victims of the AD79 eruption of vesuvius

The stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope values of bone collagen are frequently used in paleodietary studies to assess the marine contribution to an individual's diet. Surprisingly, the relationship between stable isotope these values characteristics and the percentage of marine foods in diet has never been effectively demonstrated. To clarify this relationship, the stable isotope values and radiocarbon dates of nine humans and one sheep from Herculaneum, all who perished simultaneously during the AD 79 eruption of Vesuvius, were determined. Significant differences were found in the radiocarbon dates which are attributable to the incorporation of “old” carbon from the marine reservoir. The magnitude of the observed differences was linearly correlated with both δ¹³C and δ¹⁵N values allowing the response of each isotope to increasing marine carbon in collagen to be independently verified. Regression analyses showed that for every 1‰ enrichment in δ¹³C and δ¹⁵N, 56 years and 34 years were added to the radiocarbon age, respectively. Predictions of the maximum marine reservoir age differed considerably depending on which stable isotope was considered. This discrepancy is attributed to some degree of macronutrient scrambling whereby nitrogen from marine protein is preferentially incorporated in collagen over marine carbon. It is suggested that the macronutrient scrambling explains the observed relationship between δ¹³C and δ¹⁵N from Roman coastal sites and should be considered when interpreting any diet which is not dominated by protein. Nevertheless, without knowing the degree of macronutrient scrambling in different dietary scenarios, the accuracy of dietary reconstructions is severely compromised. Am J Phys Anthropol 152:345–352, 2013. © 2013 Wiley Periodicals, Inc.     

Stable isotope composition and parasitic infections of harbor seal young‐of‐the‐year used as prey‐based diet indicators

The stable isotope composition (δ¹³C and δ¹⁵N values) of harbor seals (Phoca vitulina) is influenced by their diet. Young‐of‐the‐year during lactation and postweaning fast are expected be enriched in ¹⁵N compared to foraging seals. We studied the temporal variation of stable isotope composition of young‐of‐the‐year and adults to determine from which point in time the young‐of‐the‐year tissues (i.e., muscles and vibrissae) are influenced by independent foraging only. These results were compared with the development of trophically transmitted parasitic infections. The δ¹⁵N values in young‐of‐the‐year muscles decreased from June (20.3‰ ± 0.5‰) to October (18.5‰ ± 0.4‰), while those of foraging seals were all year long below 19.2‰. This decrease coincides with the increase of parasitic infections in young‐of‐the‐year, reflecting a shift to fish diet. Together these results suggest that the muscles of the young‐of‐the‐year older than 5–6 mo reflect independent foraging and that they can therefore be used in community diet studies. The nursing signal in vibrissae was unclear, as the δ¹⁵N values of young‐of‐the‐year were stable over time, whereas those of adults varied seasonally. However, δ¹⁵N values of nursing pups were significantly higher than those of adults in May and June, maybe due to their reliance on milk.     

Thermal habitat segregation among morphotypes of whitefish (Coregonus lavaretus: Salmonidae) and invasive vendace (C. albula): a mechanism for co‐existence?

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Effect of nutritional condition on variation in δ13C and δ15N stable isotope values in Pumpkinseed sunfish (Lepomis gibbosus) fed different diets

Stable isotope analysis is frequently used to infer resource use in natural populations of fishes. Studies have examined factors, other than diet, that influence δ¹⁵N and δ¹³C including tissue-specific rates of equilibration and starvation. Most such studies completed under laboratory conditions tightly control food quantity and its isotopic composition, but it is also necessary to evaluate the influence of these factors under more natural conditions. Using pumpkinseed sunfish (Lepomis gibbosus) we evaluated whether restricted rations below minimum daily requirements affects tissue equilibration to a change in diet by holding fish on two treatments that often reflect divergent resource use in natural populations (pelagic zooplankton or littoral macroinvertebrates). Over 42 days, δ¹⁵N values increased while δ¹³C values did not change, additionally neither were related to diet treatment. Increased δ¹⁵N values were negatively related to body condition while δ¹³C values were not, indicating that stable isotope values were more affected by decreasing body condition than by diet. Additionally, δ¹⁵N values changed more in the blood and liver tissues than in white muscle tissue, indicating that restricting food availability had greater effects on tissues with greater metabolic activity. We hypothesize that stable isotope values of consumers are subject to a tissue-specific trade-off between sensitivity to changes in resource use and resistance to the effects of low resource availability. This trade-off may require consideration in stable isotope studies of wild populations facing periodic limitations of food availability.     

Ontogenetic variation in diet and habitat of Irrawaddy dolphins (Orcaella brevirostris) in the Gulf of Thailand and the Andaman Sea

Irrawaddy dolphins (Orcaella brevirostris) are threatened, little‐known cetaceans primarily jeopardized by fishing gear entanglement. Information regarding life history, foraging ecology, and movement are crucial to determine management units and conservation regions. This is the first analysis of the ontogenetic changes in diet and habitat of Irrawaddy dolphins from the Gulf of Thailand and the Andaman Sea using stable isotopes from skin and teeth. Results suggest limited age and sex structure indicated by weak ontogenetic and sexual variability in δ¹⁵N and δ¹³C in teeth. Significant variation in δ¹³C and δ¹⁵N values were observed between three Gulf regions and the Andaman Sea, indicating distinct regional groups. Isotope mixing models run on soft tissues from the eastern Gulf revealed a diet based primarily on fishes and secondarily on crustaceans, suggesting overlap with fishery targets. Conservation strategies that focus on reducing regional competition with humans and bycatch mortality are proposed. Fishing gear improvements and monitoring of fishing operations are a potential immediate response to reduce anthropogenic impact.     

Non-lethal dorsal fin sampling for stable isotope analysis in seahorses

Sampling collection for stable isotope analysis has traditionally involved the sacrifice of the animal. Seahorses (Hippocampus spp.) are listed as threatened by the Convention on International Trade in Endangered Species (http://www.cites.org) and consequently lethal sampling is undesirable. We evaluated the adequacy of dorsal fin tissue of adult seahorses Hippocampus guttulatus for stable isotope analysis as an alternative to lethal tissue sampling. Three seahorse tissues (dorsal fin, muscle, and liver) were analyzed for comparisons of δ¹⁵N and δ¹³C values. Similarities found between δ¹⁵N and δ¹³C values in dorsal fin and muscle tissue of H. guttulatus suggest that both tissues are adequate for stable isotope analysis to understand feeding ecology of seahorses. However, considering the threatened status of the species, dorsal fin tissue would be recommended in adult seahorses as a non-lethal sampling. The effect of lipid extraction on carbon and nitrogen stable isotope values was also evaluated in each seahorse tissues. Significant effects of lipids extraction did only occur for δ¹³C values in muscle and liver. It was found that lipid removal was not necessary to perform SIA in dorsal fin tissues. Due to the limited availability of fin tissue obtained from fin-clipping in seahorses, the relationship between the mass/surface of dorsal fin clip and stable isotope values was analyzed. δ¹⁵N and δ¹³C values in fin samples were found to be independent of the size of fin analyzed. According to our study, the use of fin-clipping sampling, with a minimum surface analyzed of 12.74?mm², was found to be an adequate method for SIA in seahorses.     

Stable isotope analyses on archived fish scales reveal the long‐term effect of nitrogen loads on carbon cycling in rivers

Stable isotope analysis of organic matter in sediment records has long been used to track historical changes in productivity and carbon cycling in marine and lacustrine ecosystems. While flow dynamics preclude stratigraphic measurements of riverine sediments, such retrospective analysis is important for understanding biogeochemical cycling in running waters. Unique collections of riverine fish scales were used to analyse δ¹⁵N and δ¹³C variations in the food web of two European rivers that experience different degrees of anthropogenic pressure. Over the past four decades, dissolved inorganic N loading remained low and constant in the Teno River (70°N, Finland); in contrast, N loading increased fourfold in the Scorff River (47°N, France) over the same period. Archived scales of Atlantic salmon parr, a riverine life‐stage that feeds on aquatic invertebrates, revealed high δ¹⁵N values in the Scorff River reflecting anthropogenic N inputs to that riverine environment. A strong correlation between dissolved inorganic N loads and δ¹³C values in fish scales was observed in the Scorff River, whereas no trend was found in the Teno River. This result suggests that anthropogenic N‐nutrients enhanced atmospheric C uptake by primary producers and its transfer to fish. Our results illustrate for the first time that, as for lakes and marine ecosystems, historical changes in anthropogenic N loading can affect C cycling in riverine food webs, and confirm the long‐term interactions between N and C biogeochemical cycles in running waters.     

Stable isotope enrichment in laboratory ant colonies: effects of colony age,metamorphosis, diet,and fat storage

Ecologists use stable isotopes to infer diets and trophic levels of animals in food webs, yet some assumptions underlying these inferences have not been thoroughly tested. We used laboratory‐reared colonies of Solenopsis invicta Buren (Hymenoptera: Formicidae: Solenopsidini) to test the effects of metamorphosis, diet, and lipid storage on carbon and nitrogen stable isotope ratios. Effects of metamorphosis were examined in ant colonies maintained on a control diet of domestic crickets and sucrose solution. Effects of a diet shift were evaluated by adding a tuna supplement to select colonies. Effects of lipid content on stable isotopes were tested by treating worker ants with polar and non‐polar solvents. δ¹³C and δ¹⁵N values of larvae, pupae, and workers were measured by mass spectrometry on whole‐animal preparations. We found a significant effect of colony age on δ¹³C, but not δ¹⁵N; larvae, pupae, and workers collected at 75 days were slightly depleted in ¹³C relative to collections at 15 days (Δδ¹³C = ?0.27‰). Metamorphosis had a significant effect on δ¹⁵N, but not δ¹³C; tissues of each successive developmental stage were increasingly enriched in ¹⁵N (pupae, +0.5‰; workers, +1.4‰). Availability of tuna resulted in further shifts of about +0.6‰ in isotope ratios for all developmental stages. Removing fat with organic solvents had no effect on δ¹³C, but treatment with a non‐polar solvent resulted in enriched δ¹⁵N values of +0.37‰. Identifying regular patterns of isotopic enrichment as described here should improve the utility of stable isotopes in diet studies of insects. Our study suggests that researchers using ¹⁵N enrichment to assess trophic levels of an organism at different sites need to take care not to standardize with immature insect herbivores or predators at one site and mature ones at another. Similar problems may also exist when standardizing with holometabolous insects at one site and spiders or hemimetabolous insects at another site.