Isotopic niche overlap of two planktivorous fish in southern China

The purpose of this study was to assess if there was trophic niche overlap of silver carp (Hypophthalmichthys molitrix) and bighead carp (H. nobilis) in four large freshwater ecosystems from southern China using stable carbon and nitrogen isotopes (δ¹³C and δ¹⁵N). Multivariate analysis of variance (MANOVA) on the δ¹³C and δ¹⁵N values measured from muscle tissue indicates trophic niche overlap in one unproductive and one highly productive large system and trophic niche segregation in two systems with moderate watershed size and productivity. For these two coexisting planktivorous fish, which were hitherto believed to occupy different trophic niches, this study demonstrated that the degree of their trophic niche overlap varied according to ecosystem properties.     

Fast carnivores and slow herbivores: differential foraging strategies among grizzly bears in the Canadian Arctic

Categorizing animal populations by diet can mask important intrapopulation variation, which is crucial to understanding a species’ trophic niche width. To test hypotheses related to intrapopulation variation in foraging or the presence of diet specialization, we conducted stable isotope analysis (δ¹³C, δ¹⁵N) on hair and claw samples from 51 grizzly bears (Ursus arctos) collected from 2003 to 2006 in the Mackenzie Delta region of the Canadian Arctic. We examined within-population differences in the foraging patterns of males and females and the relationship between trophic position (derived from δ¹⁵N measurements) and individual movement. The range of δ¹⁵N values in hair and claw (2.0–11.0‰) suggested a wide niche width and cluster analyses indicated the presence of three foraging groups within the population, ranging from near-complete herbivory to near-complete carnivory. We found no linear relationship between home range size and trophic position when the data were continuous or when grouped by foraging behavior. However, the movement rate of females increased linearly with trophic position. We used multisource dual-isotope mixing models to determine the relative contributions of seven prey sources within each foraging group for both males and females. The mean bear dietary endpoint across all foraging groups for each sex fell toward the center of the mixing polygon, which suggested relatively well-mixed diets. The primary dietary difference across foraging groups was the proportional contribution of herbaceous foods, which decreased for both males and females from 42–76 to 0–27% and 62–81 to 0–44%, respectively. Grizzlies of the Mackenzie Delta live in extremely harsh conditions and identifying within-population diet specialization has improved our understanding of varying habitat requirements within the population.     

Isotopic tracking of foraging and long-distance migration in northeastern Pacific pinnipeds

We investigated the impact of foraging location (nearshore vs offshore) and foraging latitude (high vs middle) on the carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope compositions of bone collagen of northern fur seals (Callorhinus ursinus), harbor seals (Phoca vitulina), California sea lions (Zalophus californianus), and northern elephant seals (Mirounga angustirostris). Nearshore-foraging harbor seals from California had δ¹³C values 2.0‰ higher than female northern elephant seals foraging offshore at similar latitudes. Likewise, nearshore-foraging harbor seals from Alaska had values 1.7‰ higher than male northern fur seals, which forage offshore at high latitudes. Middle-latitude pinnipeds foraging in either the nearshore or offshore were ¹³C enriched by ∼1.0‰ over similar populations from high latitudes. Male northern elephant seals migrate between middle and high latitudes, but they had δ¹³C values similar to high-latitude, nearshore foragers. Female northern fur seal δ¹³C values were intermediate between those of high- and middle-latitude offshore foragers, reflecting their migration between high- and middle-latitude waters. The δ¹³C values of California sea lions were intermediate between nearshore- and offshore-foraging pinnipeds at middle latitudes, yet there was no observational support for the suggestion that they use offshore food webs. We suggest that their “intermediate” values reflect migration between highly productive and less-productive, nearshore ecosystems on the Pacific coasts of California and Mexico. The relative uniformity among all of these pinnipeds in δ¹⁵N values, which are strongly sensitive to trophic level, reveals that the carbon isotope patterns result from differences in the δ¹³C of organic carbon at the base of the food web, rather than differences in trophic structure, among these regions. Finally, the magnitude and direction of the observed nearshore-offshore and high-to middle-latitude differences in δ¹³C values suggest that these gradients may chiefly reflect differences in rates and magnitudes of phytoplankton production as well as the δ¹³C value of inorganic carbon available for photosynthesis, rather than the input of ¹³C-enriched macroalgal carbon to nearshore food webs. Received: 8 September 1998 / Accepted: 24 February 1999     

Diet differences among age classes of Arctic seals: evidence from stable isotope and mercury biomarkers

A basic understanding of current food web dynamics and baseline data from which to measure future change is necessary to understand species re-distribution and altered competition for food with climate change. We use mercury (Hg) and carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope ratios as biomarkers to understand species diet differences and age class differences among ringed (Phoca hispida), bearded (Erignathus barbatus), and harbour (P. vitulina) seals in a subarctic marine ecosystem. Adult bearded seals had significantly lower δ¹⁵N and muscle Hg than bearded seal pups, whereas the opposite was observed in ringed seals where pups had lower δ¹⁵N than adults, suggesting age specific foraging differences in trophic food level for both species. For harbour seals, δ¹⁵N did not differ significantly among ages while Hg and δ¹³C did. The δ¹³C in muscle supports that bearded seals in this study are benthic feeders and are part of a separate food web from ringed seals and harbour seals. Harbour seals had the highest levels of mercury and δ¹⁵N, indicating they feed at a higher trophic level relative to the other two seal species. Carbon and nitrogen isotopic ratios and Hg levels illustrate how resources are partitioned among three seal species and offer evidence for separation based on life stages within species.     

Using stable-isotope analysis of feathers to distinguish moulting and breeding origins of seabirds

To determine whether stable isotope measurements of bird feathers can be used to identify moulting (interbreeding) foraging areas of adult seabirds, we examined the stable-carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopic composition of feathers of chicks and adults of black-browed albatrosses (Diomedea melanophrys) from Kerguelen Islands, southern Indian Ocean. Albatross chicks are fed primarily fish (75% by mass), the diet being dominated by various species of the family Nototheniidae and Channichthyidae which commonly occur in the shelf waters in the vicinity of the colony. δ¹³C and δ¹⁵N values in chick feathers, which are grown in summer in the breeding area, were lower than values in adult feathers, which are grown in winter (δ¹³C: –19.6‰ versus –17.6‰ and δ¹⁵N: 12.4‰ versus 15.7‰, respectively). No differences in δ¹³C and δ¹⁵N values were found in adult wing feathers moulted in 1993 and 1994 and in adult feathers formed at the beginning, middle and end of the 1994 moulting period. These data are consistent with adults moulting in the same area and feeding at the same trophic level from one year to the next and with no major changes in foraging ecology within a given moulting season; they suggest that foraging grounds were different in summer and winter and that these differed in their stable-isotope signature. Changes in both feather δ¹³C and δ¹⁵N values indicated feeding south of the Subtropical Front (STF) during chick rearing, which is in agreement with the known foraging ecology at this time and feeding north of the STF during moult. This, together with band recoveries from adult birds, indicates that black-browed albatrosses from Kerguelen Islands wintered in subtropical waters off southern Australia. The stable-isotope markers in feathers, therefore, have the potential for locating moulting areas of migratory seabird species moving between isotopically distinct regions and for investigating seabirds’ foraging ecology during the poorly known interbreeding period. Such information is needed for studies of year-round ecology of seabirds as well as for their conservation and the long-term monitoring of the pelagic environment. Received: 28 June 1999 / Accepted: 14 September 1999     

The Long and the Short of It: No Dietary Specialisation between Male and Female Western Sandpipers Despite Strong Bill Size Dimorphism

Many bird species show spatial or habitat segregation of the sexes during the non-breeding season. One potential ecological explanation is that differences in bill morphology favour foraging niche specialisation and segregation. Western sandpipers Calidris mauri have pronounced bill size dimorphism, with female bills averaging 15% longer than those of males. The sexes differ in foraging behaviour and exhibit partial latitudinal segregation during the non-breeding season, with males predominant in the north and females in the south. Niche specialisation at a local scale might account for this broad geographic pattern, and we investigated whether longer-billed females and shorter-billed males occupy different foraging niches at 16 sites across the non-breeding range. We used stable-nitrogen (δ¹⁵N) isotope analysis of whole blood to test for dietary specialisation according to bill length and sex. Stable-nitrogen isotope ratios increase with trophic level. We predicted that δ¹⁵N values would increase with bill length and would be higher for females, which use a greater proportion of foraging behaviour that targets higher-trophic level prey. We used stable-carbon (δ¹³C) isotope analysis to test for habitat segregation according to bill length and sex. Stable-carbon isotope ratios vary between marine- and freshwater-influenced habitats. We predicted that δ¹³C values would differ between males and females if the sexes segregate between habitat types. Using a model selection approach, we found little support for a relationship between δ¹⁵N and either bill length or sex. There was some indication, however, that more marine δ¹³C values occur with shorter bill lengths. Our findings provide little evidence that male and female western sandpipers exhibit dietary specialisation as a function of their bill size, but indicate that the sexes may segregate in different habitats according to bill length at some non-breeding sites. Potential ecological factors underlying habitat segregation between sexes include differences in preferred habitat type and predation risk.     

Evaluating the utility of stable isotope analyses of archived freshwater sample materials

We evaluated the potential utility of stable isotope analysis of tissues commonly archived by aquatic biologists. Previous studies with chemically preserved samples have shown contradictory results, which present an obstacle for the use of archived sample materials. We tested the effects of ethanol and formalin preservation on zooplankton and of ethanol on benthic macroinvertebrate δ¹³C and δ¹⁵N values. We found that neither formalin nor ethanol had a significant effect on δ¹³C and δ¹⁵N values of preserved zooplankton. Nor did ethanol significantly affect δ¹³C or δ¹⁵N values of macroinvertebrates. However, ethanol preservation slightly, but significantly decreased C:N ratios of both zooplankton and macroinvertebrates, probably reflecting some extraction of lipids. Overall, the effects of preservatives on δ¹³C and δ¹⁵N values that we observed were minor. We also compared δ¹³C and δ¹⁵N values analysed from roach scales and perch operculum bones with those analysed from muscle tissue. Decalcification of scales and operculum bones only slightly improved our comparison to muscle tissue δ¹³C and δ¹⁵N values. Decalcified scales had slightly higher δ¹³C and lower δ¹⁵N values. Similarly, decalcified operculum bones showed slightly increased δ¹³C and decreased δ¹⁵N values to those for fish muscle. Our results confirm that scales and operculum bones can provide a suitable proxy for fish muscle in isotope studies with minor correction. We conclude that various archived sample materials can indeed be used with confidence for historical reconstructions of freshwater food webs by stable isotope analysis. Handling editor: K. Martens     

Diet,individual specialisation and breeding of brown skuas (<Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>): an investigation using stable isotopes

The diet of brown skuas (Catharacta antarctica lonnbergi) on Bird Island, South Georgia was assessed using a combination of stable isotope analysis (SIA) and mixing model techniques. We found evidence that individual specialisation in diet of adult brown skuas was related to timing of breeding, which may reflect differences in intrinsic quality. Adults with more enriched ¹³C values hatched chicks earlier than those with depleted ¹³C values. Individuals with enriched ¹³C fed predominantly on Antarctic fur seal (Arctocephalus gazella) carrion and placenta while those with lower ratios appeared to rely more on burrowing petrels (e.g. Antarctic prions Pachyptila desolata). Individual foraging differences clearly influenced timing of breeding and potentially the reproductive output of breeding pairs. We confirmed that the main components of the diet of brown skuas during incubation are, in decreasing order of importance, Antarctic fur seal placenta, burrowing petrels and fur seal muscle. In addition, we identified fur seal faeces in the diet during this stage, which had not been detected previously by traditional sampling methods. Finally we identified a correlation in δ¹³C values between pair members, attributable to the influence of courtship feeding of females by males, or assortative mating according to foraging preference or intrinsic quality.     

Functional diversity in an Amazonian rainforest of French Guyana: a dual isotope approach (δ15N and δ13C)

Functional aspects of biodiversity were investigated in a lowland tropical rainforest in French Guyana (5°2′N, annual precipitation 2200 mm). We assessed leaf δ¹⁵N as a presumptive indicator of symbiotic N₂ fixation, and leaf and wood cellulose δ¹³C as an indicator of leaf intrinsic water-use efficiency (CO₂ assimilation rate/leaf conductance for water vapour) in dominant trees of 21 species selected for their representativeness in the forest cover, their ecological strategy (pioneers or late successional stage species, shade tolerance) or their potential ability for N₂ fixation. Similar measurements were made in trees of native species growing in a nearby plantation after severe perturbation (clear cutting, mechanical soil disturbance). Bulk soil δ¹⁵N was spatially quite uniform in the forest (range 3–5‰), whereas average leaf δ¹⁵N ranged from −0.3‰ to 3.5‰ in the different species. Three species only, Diplotropis purpurea, Recordoxylon speciosum (Fabaceae), and Sclerolobium melinonii (Caesalpiniaceae), had root bacterial nodules, which was also associated with leaf N concentrations higher than 20 mg g⁻¹. Although nodulated trees displayed significantly lower leaf δ¹⁵N values than non-nodulated trees, leaf δ¹⁵N did not prove a straightforward indicator of symbiotic fixation, since there was a clear overlap of δ¹⁵N values for nodulated and non-nodulated species at the lower end of the δ¹⁵N range. Perturbation did not markedly affect the difference δ¹⁵N_soil−δ¹⁵N_leaf, and thus the isotopic data provide no evidence of an alteration in the different N acquisition patterns. Extremely large interspecific differences in sunlit leaf δ¹³C were observed in the forest (average values from −31.4 to −26.7‰), corresponding to intrinsic water-use efficiencies (ratio CO₂ assimilation rate/leaf conductance for water vapour) varying over a threefold range. Wood cellulose δ¹³C was positively related to total leaf δ¹³C, the former values being 2–3‰ higher than the latter ones. Leaf δ¹³C was not related to leaf δ¹⁵N at either intraspecific or interspecific levels. δ¹³C of sunlit leaves was highest in shade hemitolerant emergent species and was lower in heliophilic, but also in shade-tolerant species. For a given species, leaf δ¹³C did not differ between the pristine forest and the disturbed plantation conditions. Our results are not in accord with the concept of existence of functional types of species characterized by common suites of traits underlying niche differentiation; rather, they support the hypothesis that each trait leads to a separate grouping of species. Received: 18 August 1997 / Accepted: 14 April 1998     

Trophic niches of thirteen damselfishes (Pomacentridae) at the Grand Récif of Toliara,Madagascar

The damselfishes, with more than 340 species, constitute one of the most important families that live in the coral reef environment. Most of our knowledge of reef-fish ecology is based on this family, but their trophic ecology is poorly understood. The aim of the present study was to determine the trophic niches of 13 sympatric species of damselfishes by combining stable isotope (δ¹⁵N and δ¹³C) and stomach content analyses. Isotopic signatures reveal three main groups according to their foraging strategies: pelagic feeders (Abudefduf sexfasciatus, A. sparoides, A. vaigiensis, Chromis ternatensis, C. dimidiata, Dascyllus trimaculatus and Pomacentrus caeruleus), benthic feeders (Chrysiptera unimaculata, Plectroglyphidodon lacrymatus and Amphiprion akallopisos) and an intermediate group (D. aruanus, P. baenschi and P. trilineatus). Stomach contents reveal that planktonic copepods and filamentous algae mainly represent the diets of pelagic feeders and benthic feeders, respectively. The intermediate position of the third group resulted from a partitioning of small planktonic prey, small vagile invertebrates and filamentous algae. In this last feeding group, the presence of a wide range of δ¹³C values in P. trilineatus suggests a larger trophic niche width, related to diet-switching over time. Some general considerations about the feeding habits of damselfishes reveal that their choice of habitat on the reef and their behavior appear to be good predictors of diet in this group. Benthic (algae and/or small invertebrates) feeders appear to be solitary and defend a small territory on the bottom; zooplankton feeders remain in groups just above the reef, in the water column.     

Does avian malaria infection affect feather stable isotope signatures?

It is widely accepted that stable isotope ratios in inert tissues such as feather keratin reflect the dietary isotopic signature at the time of the tissue synthesis. However, some elements such as stable nitrogen isotopes can be affected by individual physiological state and nutritional stress. Using malaria infection experiment protocols, we estimated the possible effect of malaria parasite infections on feather carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope signatures in juvenile common crossbills Loxia curvirostra. The birds were experimentally infected with Plasmodium relictum (lineage SGS1) and P. ashfordi (GRW2), two widespread parasites of passerines. Experimental birds developed heavy parasitemia of both parasites and maintained high levels throughout the experiment (33 days). We found no significant difference between experimental and control birds in both δ¹³C and δ¹⁵N values of feathers re-grown. The study shows that even heavy primary infections of malaria parasites do not affect feather δ¹³C and δ¹⁵N isotopic signatures. The results of this experiment demonstrate that feather isotope values of wild-caught birds accurately reflect the dietary isotopic sources at the time of tissue synthesis even when the animal’s immune system might be challenged due to parasitic infection.     

Dietary and isotopic specialization: the isotopic niche of three <Emphasis Type="Italic">Cinclodes</Emphasis> ovenbirds

By comparing the isotopic composition of tissues deposited at different times, we can identify individuals that shift diets over time and individuals with constant diets. We define an individual as an isotopic specialist if tissues deposited at different times have similar isotopic composition. If tissues deposited at different times differ in isotopic composition we define an individual as an isotopic generalist. Individuals can be dietary generalists but isotopic specialists if they feed on the same resource mixture at all times. We assessed the degree of isotopic and dietary specialization in three related Chilean bird species that occupy coastal and/or freshwater environments: Cinclodes oustaleti, Cinclodes patagonicus, and Cinclodes nigrofumosus. C. oustaleti individuals were both isotopic and dietary generalists. Tissues deposited in winter (liver and muscle) had distinct stable C (δ¹³C) and stable N isotope ratio (δ¹⁵N) values from tissues deposited in the summer (wing feathers) suggesting that birds changed the resources that they used seasonally from freshwater habitats in the summer to coastal habitats in the winter. Although the magnitude of seasonal isotopic change was high, the direction of isotopic change varied little among individuals. C. patagonicus included both isotopic specialists and generalists, as well as dietary specialists and generalists. The isotopic composition of the feathers and liver of some C. patagonicus individuals was similar, whereas that of others differed. In C. patagonicus, there were large inter-individual differences in the magnitude and the direction of seasonal isotopic change. All individuals of C. nigrofumosus were both isotopic and dietary specialists. The distribution of δ¹³C and δ¹⁵N values overlapped broadly among tissues and clustered in a small, and distinctly intertidal, region of δ space. Assessing individual specialization and unraveling the factors that influence it, have been key questions in animal ecology for decades. Stable isotope analyses of several tissues in appropriate study systems provide an unparalleled opportunity to answer them.     

Effects of nutritional restriction on nitrogen and carbon stable isotopes in growing seabirds

When using stable isotopes as dietary tracers it is essential to consider effects of nutritional state on isotopic fractionation. While starvation is known to induce enrichment of ¹⁵N in body tissues, effects of moderate food restriction on isotope signatures have rarely been tested. We conducted two experiments to investigate effects of a 50–55% reduction in food intake on δ¹⁵N and δ¹³C values in blood cells and whole blood of tufted puffin chicks, a species that exhibits a variety of adaptive responses to nutritional deficits. We found that blood from puffin chicks fed ad libitum became enriched in ¹⁵N and ¹³C compared to food-restricted chicks. Our results show that ¹⁵N enrichment is not always associated with food deprivation and argue effects of growth on diet–tissue fractionation of nitrogen stable isotopes (Δ¹⁵N) need to be considered in stable isotope studies. The decrease in δ¹³C of whole blood and blood cells in restricted birds is likely due to incorporation of carbon from ¹³C-depleted lipids into proteins. Effects of nutritional restriction on δ¹⁵N and δ¹³C values were relatively small in both experiments (δ¹⁵N: 0.77 and 0.41‰, δ¹³C: 0.20 and 0.25‰) compared to effects of ecological processes, indicating physiological effects do not preclude the use of carbon and nitrogen stable isotopes in studies of seabird ecology. Nevertheless, our results demonstrate that physiological processes affect nitrogen and carbon stable isotopes in growing birds and we caution isotope ecologists to consider these effects to avoid drawing spurious conclusions.     

Variability and directionality of temporal changes in δ<Superscript>13</Superscript>C and δ<Superscript>15</Superscript>N of aquatic invertebrate primary consumers

Seasonal oscillations in the carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope signatures of aquatic algae can cause seasonal enrichment–depletion cycles in the isotopic composition of planktonic invertebrates (e.g., copepods). Yet, there is growing evidence that seasonal enrichment–depletion cycles also occur in the isotope signatures of larger invertebrate consumers, taxa used to define reference points in isotope-based trophic models (e.g., trophic baselines). To evaluate the general assumption of temporal stability in non-zooplankton aquatic invertebrates, δ¹³C and δ¹⁵N time series data from the literature were analyzed for seasonality and the influence of biotic (feeding group) and abiotic (trophic state, climate regime) factors on isotope temporal patterns. The amplitude of δ¹³C and δ¹⁵N enrichment–depletion cycles was negatively related to body size, although all size-classes of invertebrates displayed a winter-to-summer enrichment in δ¹³C and depletion in δ¹⁵N. Among feeding groups, periphytic grazers were more variable and displayed larger temporal changes in δ¹³C than detritivores. For nitrogen, temporal variability and magnitude of directional change of δ¹⁵N was most strongly related to ecosystem trophic state (eutrophic > mesotrophic, oligotrophic). This study provides evidence of seasonality in the isotopic composition of aquatic invertebrates across very broad geographical and ecological gradients as well as identifying factors that are likely to modulate the strength and variability of seasonality. These results emphasize the need for researchers to recognize the likelihood of temporal changes in non-zooplankton aquatic invertebrate consumers at time scales relevant to seasonal studies and, if present, to account for temporal dynamics in isotope trophic models.     

Mechanism for the small-scale movement of carbon among estuarine habitats: organic matter transfer not crab movement

In theory, carbon is highly mobile in aquatic systems. Recent evidence from carbon stable isotopes of crabs (Parasesarma erythrodactyla and Australoplax tridentata), however, shows that in subtropical Australian waters, measurable carbon movement between adjacent mangrove and saltmarsh habitats is limited to no more than a few metres. We tested whether the pattern in crab δ¹³C values across mangrove and saltmarsh habitats was explained by crab movement, or the movement of particulate organic matter. We estimated crab movement in a mark–recapture program using an array of pitfall traps on 13 transects (a total of 65 traps) covering an area of 600 m² across the interface of these two habitats. Over a 19-day period, the majority of crabs (91% for P. erythrodactyla, 93% for A. tridentata) moved <2 m from the place of initial capture. Crab movement cannot, therefore, explain the patterns in δ¹³C values of crabs. δ¹³C values of detritus collected at 2-m intervals across the same habitat interface fitted a sigmoidal curve of a similar form to that fitting the δ¹³C values of crabs. δ¹³C values of detritus were 2–4‰ more depleted in saltmarsh (−18.5±0.6‰), and 4–7‰ more depleted in mangroves (−25.9±0.1‰) than δ¹³C values of crabs recorded previously in each habitat. Assimilation by crabs of very small detrital fragments or microphytobenthos, more enriched in ¹³C, may explain the disparity in δ¹³C values. Nevertheless, the pattern in δ¹³C values of detritus suggests that crabs obtain their carbon from up to several metres away, but without themselves foraging more then a metre or so from their burrow. Such detailed measurements of carbon movement in estuaries provide a spatially explicit understanding of the functioning of food webs in saltmarsh and mangrove habitats.     

Sexual Niche Segregation and Gender-Specific Individual Specialisation in a Highly Dimorphic Marine Mammal

While sexual segregation is expected in highly dimorphic species, the local environment is a major factor driving the degree of resource partitioning within a population. Sexual and individual niche segregation was investigated in the Australian fur seal (Arctocephalus pusillus doriferus), which is a benthic foraging species restricted to the shallow continental shelf region of south-eastern Australia. Tracking data and the isotopic values of plasma, red blood cells and whiskers were combined to document spatial and dietary niche segregation throughout the year. Tracking data indicated that, in winter, males and females overlapped in their foraging habitat. All individuals stayed within central Bass Strait, relatively close (< 220 km) to the breeding colony. Accordingly, both genders exhibited similar plasma and red cell δ¹³C values. However, males exhibited greater δ¹³C intra-individual variation along the length of their whisker than females. This suggests that males exploited a greater diversity of foraging habitats throughout the year than their female counterparts, which are restricted in their foraging grounds by the need to regularly return to the breeding colony to suckle their pup. The degree of dietary sexual segregation was also surprisingly low, both sexes exhibiting a great overlap in their δ¹⁵N values. Yet, males displayed higher δ¹⁵N values than females, suggesting they fed upon a higher proportion of higher trophic level prey. Given that males and females exploit different resources (mainly foraging habitats), the degree of individual specialisation might differ between the sexes. Higher degrees of individual specialisation would be expected in males which exploit a greater range of resources. However, comparable levels of inter-individual variation in δ¹⁵N whisker values were found in the sampled males and females, and, surprisingly, all males exhibited similar seasonal and inter-annual variation in their δ¹³C whisker values, suggesting they all followed the same general dispersion pattern throughout the year.     

Contributions of Different Organic Carbon Sources to <Emphasis Type="Italic">Daphnia</Emphasis> in the Pelagic Foodweb of a Small Polyhumic Lake: Results from Mesocosm DI<Superscript>13</Superscript>C-Additions

Abstract Freshwater ecosystems derive organic carbon from both allochthonous and autochthonous sources. We studied the relative contributions of different carbon sources to zooplankton in a small, polyhumic, steeply stratified lake, using six replicate surface-to-sediment enclosures established during summer and autumn 2004. We added ¹³C-enriched bicarbonate to the epilimnion of half the enclosures for three weeks during each season and monitored carbon stable isotope ratios of DIC, DOC, POC and Daphnia, along with physical, chemical and biological variables. During summer, ¹³C-enriched DIC (δ¹³C up to 44 ± 7.2‰) was soon taken up by phytoplankton (δ¹³C up to −5.1 ± 13.6‰) and was transmitted to Daphnia (δ¹³C up to −1.7 ± 7.2‰), demonstrating consumption of phytoplankton. In contrast, during autumn, ¹³C-enriched DIC (δ¹³C up to 56.3 ± 9.8‰) was not transmitted to Daphnia, whose δ¹³C became progressively lower (δ¹³C down to −45.6 ± 3.3‰) concomitant with decreasing methane concentration. Outputs from a model suggested phytoplankton contributed 64–84% of Daphnia diet during summer, whereas a calculated pelagic carbon mass balance indicated only 30–40% could have come from phytoplankton. Although autumn primary production was negligible, zooplankton biomass persisted at the summer level. The model suggested methanotrophic bacteria contributed 64–87% of Daphnia diet during autumn, although the calculated carbon mass balance indicated a contribution of 37–112%. Thus methanotrophic bacteria could supply virtually all the carbon requirement of Daphnia during autumn in this lake. The strongly ¹³C-depleted Daphnia values, together with the outputs from the models and the calculated carbon mass balance showed that methanotrophic bacteria can be a greater carbon source for Daphnia in lakes than previously suspected.     

Foraging areas of Wilson’s storm-petrel Oceanites oceanicus in the breeding and inter-breeding period determined by stable isotope analysis

To understand the year-round ecology of seabirds it is necessary not only to study the birds in their breeding grounds, but also to gain information about their movements during the inter-breeding period. Especially for the smaller procellariiform species, such studies are still scarce, mainly due to methodological problems. The recovery rates of banded birds are low and satellite tracking devices still far too heavy to equip these small birds. Here, we present data on foraging areas of Wilson’s storm-petrel Oceanites oceanicus inferred from stable isotope analysis. We compared ratios of δ¹³C and δ¹⁵N between different life-history stages and between the breeding and inter-breeding period. Samples of adult and chick feathers, chick down and egg-white were taken between 1996 and 2005 on King-George-Island, South Shetland Islands. δ¹³C values can be clearly distinguished between the breeding and inter-breeding period. During the inter-breeding period, most pre-breeders foraged in the same area as breeders, but four pre-breeders were found to forage in latitudes north of the Subtropical Front. In the 2002 inter-breeding period adult birds wintered further north than in 2003, which is in line with the different locations of food rich frontal systems in these years. We show that isotope ratios of both δ¹³C and δ¹⁵N increase from egg white, over chick down to chick feathers. We suggest that this isotopic change, due to a change in both foraging location and diet between egg production and chick feeding, may be used to trace the shift from the use of maternal resources from the egg to the uptake of nutrients from the diet.     

Temporal and spatial variability in stable isotope compositions of a freshwater mussel: implications for biomonitoring and ecological studies

Stable isotopes can be used to elucidate ecological relationships in community and trophic studies. Findings are calibrated against baselines, e.g. from a producer or primary consumer, assumed to act as a reference to the isotopic context created by spatio-temporal attributes such as geography, climate, nutrient, and energy sources. The ability of an organism to accurately represent a community base depends on how, and over what time-scale, it assimilates ambient materials. Freshwater mussels have served as references for trophic studies of freshwater communities and as indicators of change in nutrient pollution load or source. Their suitability as reference animals has not yet been fully explored, however. We conducted a series of studies examining the suitability of freshwater mussels as isotopic baselines, using their ability to reflect variation in ambient nutrient loads as a case scenario. (1) We analyzed bivalve foot tissue δ¹⁵N and δ¹³C from 22 stream reaches in the Piedmont region of North Carolina, USA to show that compositions varied substantially among locations. Site mean bivalve δ¹³C values correlated with site ambient particulate organic matter (POM) δ¹³C values, and site mean bivalve δ¹⁵N values correlated with site ambient water dissolved δ¹⁵N-NO₃ values. (2) Similarity of results among sample types demonstrated that the minimally invasive hemolymph sample is a suitable substitute for foot tissue in δ¹⁵N analyses, and that small sample sizes generate means representative of a larger population. Both findings can help minimize the impact of sampling on imperiled freshwater mussel populations. (3) In a bivalve transplantation study we showed that hemolymph δ¹⁵N compositions responded to a shift in ambient dissolved δ¹⁵N-NO₃, although slowly. The tissue turnover time for bivalve hemolymph was 113 days. We conclude that bivalves serve best as biomonitors of chronic, rather than acute, fluctuations in stream nutrient loads, and provide initial evidence of their suitability as time-integrated isotopic baselines for community studies.     

The importance of quantifying inherent variability when interpreting stable isotope field data

Stable isotope data are often used to assess diet, trophic level, trophic niche width and the extent of omnivory. Notwithstanding ongoing discussions about the value of these approaches, variations in isotopic signatures among individuals depend on inherent variability as well as differences in feeding habitats. Remarkably, the relative contributions of diet variation and inherent variability to differences in δ¹⁵N and δ¹³C among individuals have not been quantified for the same species at the same life history stages, and inherent variability has been ignored or assumed. We quantified inherent variability in δ¹³C and δ¹⁵N among individuals of a marine fish (the European sea bass, Dicentrarchus labrax) reared in a controlled environment on a diet of constant isotopic composition and compared it with variability in δ¹³C and δ¹⁵N among individuals from wild bass populations. The analysis showed that inherent variability among reared individuals on a controlled diet was equivalent to a large proportion of the observed variability among wild individuals and, therefore, that inherent variability should be measured to establish baseline variability in wild populations before any assumptions are made about the influence of diet. Given that inherent variability is known to be dependent on species, life history stage and the environment, our results show that it should be quantified on a case-by-case basis if diet studies are intended to provide absolute assessments of dietary habits.