Moulting patterns drive within-individual variations of stable isotopes and mercury in seabird body feathers: implications for monitoring of the marine environment

One major limitation in the use of body feathers of seabirds as a monitoring tool of the trophic structure and contamination levels of marine ecosystems is the degree of heterogeneity in feather chemical composition within individuals. Here, we tested the hypothesis that moulting patterns drive body feather heterogeneity, with synchronous moult minimizing within-individual variations, in contrast to asynchronous feather growth. Chicks of white-chinned petrels Procellaria aequinoctialis (representative of bird chicks) and adults of king penguins Aptenodytes patagonicus (representative of adult penguins) that moult their body feathers synchronously showed very low within-individual variations in their feather δ¹³C and δ¹⁵N values and mercury (Hg) concentrations. By contrast, body feathers of adults of Antarctic prions Pachyptila desolata (representative of adult seabirds with asynchronous feather growth during a protracted moult) presented much higher within-individual variances for the three parameters. These findings have three important implications for birds presenting a synchronous body moult. (1) They suggest that all body feathers from the same individual have identical δ¹³C and δ¹⁵N values and Hg content. (2) They predict negligible within-individual variations in the body feather values of other useful stable isotopes, such as δ²H and δ³⁴S, as well as in the concentrations of other compounds that are deposited in the keratin structure. (3) Analysis of one or any number of pooled body feathers is equally representative of the individual. In conclusion, we recommend that long-term routine monitoring investigations focus on birds presenting synchronous rather than asynchronous moult of body feathers both in marine and terrestrial environments. This means targeting chicks rather than adults and, for seabirds, penguins rather than adults of flying species.     

Plasticity versus repeatability in seabird migratory behaviour

Pelagic seabird populations can use several discrete wintering areas, but it is unknown if individuals use the same wintering area year after year. This would have consequences for their population genetic structure and conservation. We here study the faithfulness of individuals to a moulting area within and among years in a small pelagic seabird, the Thin-billed prion, which moult their primary feathers during the early part of the non-breeding period. According to stable carbon isotope ratios (δ¹³C) of these feathers, 90% of Thin-billed prions moult in Antarctic and 10% in South American waters. Repeated samples from individuals in 2 or 3 years indicated that several birds changed between Antarctic and South American moulting areas or vice versa. However, individuals moulting in an area in one year were more likely to do so again. Four out of five adults maintained highly conserved δ¹³C over the extended moulting period. One bird, however, had systematic changes in δ¹³C indicating latitudinal movements between the two areas during moult. Thus, the present data show that this seabird species has a highly flexible migratory strategy, not only at the population level, but also at the individual level, enabling these seabirds to exploit a highly unpredictable environment.     

Stable isotopes identify age- and sex-specific dietary partitioning and foraging habitat segregation in southern giant petrels breeding in Antarctica and southern Patagonia

We examined the isotopic signatures (δ¹³C, δ¹⁵N) of adult body feathers from southern giant petrels Macronectes giganteus collected at two breeding colonies in Antarctica (Potter Peninsula and Cape Geddes) and one in southern Patagonia (Observatorio Island), as well as in whole blood collected from adults of both sexes at each Antarctic colonies and from chicks at Potter Peninsula. As body feather moult is a continuous process in giant petrels, feathers provide an integrated annual signal of an adult’s diets and foraging habitats. In contrast, the stable isotope values of adult and chick blood are reflective of their diets during the breeding season. We found that sex-specific dietary segregation in adults breeding in Antarctica was notable during the breeding season (blood samples) but absent when examined across the entire year (feather samples). In addition, blood stable isotope values differed between chicks and adults, indicating that adults provision their offspring with a relatively higher amount of penguin and seal prey that what they consume themselves. This finding confirms previous work that suggests that chicks are preferentially fed with prey of presumably higher nutritional value such as carrion. Finally, based on isotopic differences between major oceanographic zones in the Southern Ocean, our data indicate population-specific differences in foraging distribution, with Antarctic populations move seasonally between Antarctic and subantarctic zones, while Patagonian populations likely forage in subtropical waters and in continental shelf habitats year-round.     

Short- and long-term consistency in the foraging niche of wandering albatrosses

The wandering albatross (Diomedea exulans) is regarded as a generalist predator, but can it be consistent in its foraging niche at an individual level? This study tested short- and long-term consistency in the foraging niche in terms of habitat use, trophic level and, by inference, prey selection. Fieldwork was carried out at Bird Island, South Georgia, in May–October 2009, during the chick-rearing period. Blood (plasma and cells) and feathers for stable isotope analyses (δ¹³C and δ¹⁵N) were sampled from 35 adults on their return from a foraging trip during which they carried stomach temperature, activity and global positioning system loggers. Results suggest short-term consistency in foraging niche in relation to both oceanic water mass and trophic level, and long-term consistency in use of habitat. Consistent differences between individuals partly reflected sex-specific habitat preferences. The proportion of consistent individuals (i.e., with a narrow foraging niche) was estimated at c. 40?% for short-term habitat and trophic level (prey) preferences and 29?% for longer-term habitat preference, suggesting this is an important characteristic of this population and potentially of pelagic seabirds in general. Foraging consistency was not related to body condition or level of breeding experience; instead, it may reduce intraspecific competition.     

Sexual segregation in distribution,diet and trophic level of seabirds: insights from stable isotope analysis

Considerable attention has focused on inter- and intraspecific variation in trophic niches of marine predators. Although this has revealed evidence for sexual segregation in distribution in some species, few studies have been able to address sex-related dietary specialisation. Stable isotope analysis of blood cells collected from albatrosses and petrels at South Georgia during chick-rearing indicated a difference in δ¹³C, suggesting that females fed to the north of males, only in two species with male-biased sexual size dimorphism; in no species did sexes differ in trophic level (δ¹⁵N). Based on a wider review, significant differences between sexes in isotope signatures were much more common in seabirds during the pre-laying or breeding than the nonbreeding period, presumably reflecting greater between-sex partitioning of resources when foraging ranges are more constrained and competition is greater. Sex differences, or their absence, were usually consistent across successive stages during the pre-laying and breeding periods, but not necessarily year-round nor between populations. Significant differences in isotope signatures between males and females were extremely rare in monomorphic species, suggesting a link between sexual size dimorphism and segregation in diet or distribution. Among the Southern Ocean albatrosses, sex differences in δ¹³C suggested the underlying mechanism was related to habitat specialisation, whereas in other size-dimorphic taxa (both male- and female-biased), sex differences were more common in δ¹⁵N than δ¹³C and therefore more consistent with size-mediated competitive exclusion or dietary specialisation.     

The role of stable isotopes and mercury concentrations to describe seabird foraging ecology in tropical environments

Nitrogen (δ¹⁵N) and carbon (δ¹³C) stable isotopes and contaminants, such as mercury, have been widely used to characterise foraging ecology of temperate and polar seabirds. In this study, for the first time, we used isotopic signatures and mercury levels of feathers and blood of eight tropical seabird species, that forage in a range-gradient between inshore and offshore areas, to describe the foraging habits of a large tropical seabird community (from two neighboring islands of the Seychelles archipelago, western Indian Ocean) during both the breeding and inter-breeding periods. Overall, we found a high overlap in both δ¹⁵N and δ¹³C signatures among species. The high inter-specific overlap in δ¹⁵N values was expected, given the similarities in the diet of the species from this community. However, several unexpected results, such as (1) the consistently higher δ¹⁵N signatures of white terns (Gygis alba), (2) the large variation in inter-specific differences in δ¹⁵N signatures among the sampling groups (season, age, island and tissue) and (3) the consistent low δ¹⁵N values of breeding birds during the northwest monsoon (austral summer), suggest that δ¹⁵N signatures cannot be used as indicators of seabird trophic levels in this community. The high inter-specific overlap in δ¹³C signatures and the absence, during the breeding season, of a δ¹³C gradient that follows the inshore-offshore foraging gradient within the community can be explained by the habitat homogeneity of the Seychelles continental shelf and suggest that birds forage mostly within the limits of this “plateau”. On the other hand, the similarities in δ¹³C values between the breeding and inter-breeding periods in species that are known to show post-breeding dispersal, strongly support the hypothesis of a lack of latitudinal variation in δ¹³C signatures of POM in the central Indian Ocean, and the consequent inaccuracy of δ¹³C values to track seabird movements within this geographic area. Inter-specific differences in mercury levels seem to be related to prey size, while consistent higher mercury concentrations in one of the studied islands suggest different island mercury-backgrounds and possible segregation in foraging areas between the seabirds of the two islands. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A three-isotope approach to disentangling the diet of a generalist consumer: the yellow-legged gull in northwest Spain

The widespread omnivory of consumers and the trophic complexity of marine ecosystems make it difficult to infer the feeding ecology of species. The use of stable isotopic analysis plays a crucial role in elucidating trophic interactions. Here we analysed δ¹⁵N, δ¹³C and δ³⁴S in chick feathers, and we used a Bayesian triple-isotope mixing model to reconstruct the diet of a generalist predator, the yellow-legged gull (Larus michahellis) that breeds in the coastal upwelling area off northwest mainland Spain. The mixing model indicated that although chicks from all colonies were fed with a high percentage of fish, there are geographical differences in their diets. While chicks from northern colonies consume higher percentages of earthworms, refuse constitutes a more important source in the diet of chicks from western colonies. The three-isotope mixing model revealed a heterogeneity in foraging habitats that would not have been apparent if only two stable isotopes had been analysed. Moreover, our work highlights the potential of adding δ³⁴S for distinguishing not only between terrestrial and marine prey, but also between different marine species such as fish, crabs and mussels.     

Differences in δ13C and δ15N values between feathers and blood of seabird chicks: implications for non-invasive isotopic investigations

Blood and feathers are the most targeted tissues for isotopic investigations in avian ecology, primarily because they can be easily and non-destructively sampled on live individuals. Comparing blood and feather isotopic ratios can provide valuable information on dietary shifts, trophic specialization and migration patterns, but it requires a good knowledge of the isotopic differences between the two tissues. Here, δ¹³C and δ¹⁵N values of whole blood (in blood cells of a few species) and simultaneously grown body feathers were measured in seabird chicks to quantify the tissue-related isotopic differences. Seabirds include 27 populations of 22 wild species that were sampled in 2000–2008, and a review of the literature added 8 groups (including adult birds) to the analysis. The use of a large data set that overall encompasses wide δ¹³C and δ¹⁵N ranges allowed us to depict for the first time accurate relationships between blood and feather isotopic ratios across avian taxa. Blood was impoverished in ¹³C and generally in ¹⁵N compared with feathers. Both mean δ¹³C and δ¹⁵N values of feathers and blood were highly positively and linearly related [feather δ¹³C = 0.972 (±0.020) blood δ¹³C + 0.962 (±0.414), and feather δ¹⁵N = 1.014 (±0.056) blood + 0.447 (±0.665), respectively; both P < 0.0001]. The regressions should be applied to mathematically correct feather or whole blood δ¹³C and δ¹⁵N values when comparing isotopic ratios within and between ecological studies on birds.     

Isotopic evidence of distinct feeding ecologies and movement patterns in two migratory predators (yellowfin tuna and swordfish) of the western Indian Ocean

Ecologists primarily use δ¹⁵N values to estimate the trophic level of organisms, while δ¹³C, and even recently δ¹⁵N, are utilized to delineate feeding habitats. However, many factors can influence the stable isotopic composition of consumers, e.g. age, starvation or isotopic signature of primary producers. Such sources of variability make the interpretation of stable isotope data rather complex. To examine these potential sources of variability, muscle tissues of yellowfin tuna (Thunnus albacares) and swordfish (Xiphias gladius) of various body lengths were sampled between 2001 and 2004 in the western Indian Ocean during different seasons and along a latitudinal gradient (23°S to 5°N). Body length and latitude effects on δ¹⁵N and δ¹³C were investigated using linear models. Both latitude and body length significantly affect the stable isotope values of the studied species but variations were much more pronounced for δ¹⁵N. We explain the latitudinal effect by differences in nitrogen dynamics existing at the base of the food web and propagating along the food chain up to top predators. This spatial pattern suggests that yellowfin and swordfish populations exhibit a relatively unexpected resident behaviour at the temporal scale of their muscle tissue turnover. The body length effect is significant for both species but this effect is more pronounced in swordfish as a consequence of their different feeding strategies, reflecting specific physiological abilities. Swordfish adults are able to reach very deep water and have access to a larger size range of prey than yellowfin tuna. In contrast, yellowfin juveniles and adults spend most of their time in the surface waters and large yellowfin tuna continue to prey on small organisms. Consequently, nitrogen isotopic signatures of swordfish tissues are higher than those of yellowfin tuna and provide evidence for different trophic levels between these species. Thus, in contrast to δ¹³C, δ¹⁵N analyses of tropical Indian Ocean marine predators allow the investigation of complex vertical and spatial segregation, both within and between species, even in the case of highly opportunistic feeding behaviours. The linear models developed in this study allow us to make predictions of δ¹⁵N values and to correct for any body length or latitude differences in future food web studies.     

Isotopic niches of emperor and Adélie penguins in Adélie Land,Antarctica

The emperor and Adélie penguins are the only two species of penguins that co-occur at high-Antarctic latitudes. We first measured and compared their isotopic niches on the same year in Adélie Land in spring, when the two species co-exist. Emperor and Adélie penguins segregated by their blood isotopic signatures, with adult δ¹³C values (−24.5 ± 0.2 and −25.4 ± 0.2‰, respectively) suggesting that emperor penguins foraged in more neritic waters than Adélie penguins in spring. At that time, difference in their δ¹⁵N values (4.1‰, 12.0 ± 0.4 vs. 7.9 ± 0.1‰) encompassed more than one trophic level, indicating that emperor penguins preyed mainly upon fish (and squids), while Adélie penguins fed exclusively on euphausiids. Second, we compared the food of breeding adults and chicks. The isotopic signatures of adults and chicks of emperor penguins were not statistically different, but δ¹⁵N value of Adélie penguin chicks was higher than that of adults (10.2 ± 0.8 vs. 9.0 ± 0.2‰). The difference showed that adult Adélie penguins captured higher trophic level prey, i.e. higher-quality food, for their chicks. Third, the isotopic signatures of Adélie penguins breeding in Adélie Land showed that adults fed on Antarctic krill in oceanic waters in spring and shifted to neritic waters in summer where they preyed upon ice krill for themselves and upon fish and euphausiids for their chicks. A comparison of isotopic niches revealed large overlaps in both blood δ¹³C and δ¹⁵N values within the community of Antarctic seabirds and pinnipeds. The continuum in δ¹⁵N values nevertheless encompassed more than one trophic level (5.2‰) from Adélie penguin and crabeater seal to the Weddell seal. Such a broad continuum emphasizes the fact that all Antarctic seabirds and marine mammals feed on varying proportions of a few crustacean (euphausiids) and fish (Antarctic silverfish) species that dominate the intermediate trophic levels of the pelagic neritic and oceanic ecosystems.     

Utilization of nitrogen derived from seabird guano by terrestrial and marine plants at St. Paul, Pribilof Islands, Bering Sea, Alaska

Dense, conspicuous colonies of seabirds and pinnipeds breed on ocean islands throughout the world. Such colonies have been shown to have local impacts on prey populations, but whether or not they affect nutrient cycling has been debated. We determined the natural abundance levels of the stable isotopes (C and N) of primary producers, seabirds and other consumers at and near St. Paul Island, Pribilof Islands, Bering Sea, in summer 1993. Marine primary producers (phytoplankton, as particulate organic matter, and kelp) collected near seabird colonies were ca. 6.5‰ enriched in both ¹⁵N and ¹³C relative to those collected further from shore. Terrestrial plants collected near the seabird colonies were enriched in ¹⁵N (δ¹⁵N ca. 22‰) compared with conspecifics collected away from the colonies (δ¹⁵N ca. 11‰). The trend towards higher δ¹⁵N values in marine and terrestrial plants near bird colonies is consistent with their uptake of ornithogenic N. This ¹⁵N-enrichment of plants using ornithogenic N can be attributed to a combination of two processes: trophic enrichment, and volatilization of ammonia produced during degradation of terrestrially deposited guano. Seabird breeding colonies at St. Paul Island appear to be significant sources of recycled nitrogen for terrestrial plants in the vicinity of colonies and for phytoplankton in the nearshore zone. Received: 4 July 1997 / Accepted: 20 November 1997     

Analysis of the cyanotoxins anatoxin-a and microcystins in Lesser Flamingo feathers†

Feathers from carcasses of the Lesser Flamingo (Phoeniconaias minor), which had died after ingesting cyanobacterial toxins (cyanotoxins) contained between 0.02 and 30.0?µg microcystin-LR equivalents per gram of feather according to HPLC and ELISA analysis of feather extracts. Anatoxin-a was detected less frequently in the Lesser Flamingo feathers, up to 0.8?µg anatoxin-a per gram of feather being recorded. When feathers from different body regions were analysed and compared for microcystins and anatoxin-a, wing feathers were found to contain the highest concentrations of these cyanotoxins, the order of concentration and frequency of analytical detection being wing?>?breast?>?head. Consistent with the presence of the microcystins and anatoxin-a in gut contents and the livers of the dead birds and negligible in vitro adsorption to feathers, the cyanotoxins associated with the feathers of the dead wild flamingos are inferred to be primarily of dietary origin.     

Use of radar detectors to track attendance of albatrosses at fishing vessels

Despite international waters covering over 60% of the world's oceans, understanding of how fisheries in these regions shape ecosystem processes is surprisingly poor. Seabirds forage at fishing vessels, which has potentially deleterious effects for their population, but the extent of overlap and behavior in relation to ships is poorly known. Using novel biologging devices, which detect radar emissions and record the position of boats and seabirds, we measured the true extent of the overlap between seabirds and fishing vessels and generated estimates of the intensity of fishing and distribution of vessels in international waters. During breeding, wandering albatrosses (Diomedea exulans) from the Crozet Islands patrolled an area of over 10 million km² at distances up to 2500 km from the colony. Up to 79.5% of loggers attached to birds detected vessels. The extent of overlap between albatrosses and fisheries has widespread implications for bycatch risk in seabirds and reveals the areas of intense fishing throughout the ocean. We suggest that seabirds equipped with radar detectors are excellent monitors of the presence of vessels in the Southern Ocean and offer a new way to monitor the presence of illegal fisheries and to better understand the impact of fisheries on seabirds.     

Contribution of fishery discards to the diet of the Black-browed albatross (Thalassarche melanophris) during the non-breeding season: an assessment through stable isotope analysis

Black-browed albatrosses (Thalassarche melanophris) disperse over the Argentinean Continental Shelf and neighboring waters during their non-breeding season. It is one of the most frequent seabirds attending fishing vessels and also the most common Procellariform in the bycatch of longliners and trawlers in the area. Understanding the use of fishery discards by this species is an important issue when assessing the potential effect of strategic discard management in decreasing the abundance, interactions, and mitigating mortality. In the present study, we analyzed carbon and nitrogen stable isotope compositions in the blood of Black-browed albatrosses to assess the relative contribution of discards from different fisheries to the diet of this species in winter. Samples were obtained in winter 2011 from fishing vessels operating between 41–43°S and 57–59°W. No sex differences in δ¹³C and δ¹⁵N were observed. Results indicate that during their non-breeding season, isotopic signatures of Black-browed albatrosses are closer to discards and offal generated by fisheries and in particular by trawlers. The large fishing effort of trawl fisheries in Argentina highlights the urgency of an exhaustive analysis to find practical and effective ways to reduce the number of seabirds attending trawlers.     

Inter-colony movements,at-sea behaviour and foraging in an immature seabird: results from GPS-PPT tracking,radio-tracking and stable isotope analysis

Seabird populations contain large numbers of immatures––in some instances comprising >50% of the fully grown adults in the population. These birds are significant components of marine food webs and may contribute to compensatory recruitment and dispersal, but remain severely understudied. Here, we use GPS-PTTs, radio-tracking and analysis of stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes to investigate the movements and foraging ecology of immature seabirds. Our study focussed on immature northern gannets Morus bassanus aged 2–4 attending non-breeding aggregations alongside a large breeding colony. GPS-PTT tracking of five birds revealed that immatures have the ability to disperse widely during the breeding season, with some individuals potentially prospecting at other colonies. Overall, however, immatures were faithful to the colony of capture. During returns to the focal colony, immatures acted as central place foragers, conducted looping and commuting flights, and analysis of the variance in first-passage time revealed evidence of area-restricted search (ARS) behaviour. In addition, stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope analyses indicate that immatures were isotopically segregated from breeders. Our findings provide insights into the foraging, prospecting and dispersal behaviour of immature seabirds, which may have important implications for understanding seabird ecology and conservation.     

Distribution of foraging habitats of male loggerhead turtles (Caretta caretta) as revealed by stable isotopes and satellite telemetry

Most studies on the foraging ecology of loggerhead turtles (Caretta caretta) have focused on adult females and juveniles. Little is known about the foraging patterns of adult male loggerheads. We analyzed tissues for carbon and nitrogen stable isotopes (δ¹³C and δ¹⁵N) from 29 adult male loggerheads tracked with satellite transmitters from one breeding area in Florida, USA, to evaluate their foraging habitats in the Northwest Atlantic (NWA). Our study revealed large variations in δ¹³C and δ¹⁵N and a correlation between both δ¹³C and δ¹⁵N and the latitude to which the loggerheads traveled after the mating season, thus reflecting a geographic pattern in the isotopic signatures. Variation in δ¹³C and δ¹⁵N can be explained by differences in food web baseline isotopic signatures rather than differences in loggerhead trophic levels. Stable isotope analysis may help elucidate residency and migration patterns and identify foraging sea turtle subpopulations in the NWA due to the isotopically distinct habitats used by these highly migratory organisms.     

Differences in diet composition and foraging patterns between sexes of the Magellanic penguin (Spheniscus magellanicus) during the non-breeding period as revealed by δ13C and δ15N values in feathers and bone

In diving seabirds, sexual dimorphism in size often results in sex-related differences of foraging patterns. Previous research on Magellanic penguins, conducted during the breeding season, failed to reveal consistent differences between the sexes on foraging behavior, despite sexual dimorphism. In this paper, we tested the hypothesis that male and female Magellanic penguins differ in diet and foraging patterns during the non-breeding period when the constraints imposed by chick rearing activities vanish. We used stable isotope ratios of carbon and nitrogen in feather and bone to characterize the diet and foraging patterns of male and female penguins in the South Atlantic at the beginning of the 2009–2010 and 2010–2011 post-breeding seasons (feathers) and over several consecutive breeding and migratory seasons (bone). The mean δ¹³C and δ¹⁵N values of feathers showed no differences between the sexes in any of the three regions considered or in the diet composition between the sexes from identical breeding regions; however, Bayesian ellipses showed a higher isotopic niche width in males at the beginning of the post-breeding season. Stable isotope ratios in bone revealed the enrichment of males with δ¹³C compared with females across the three regions considered. Furthermore, the Bayesian ellipses were larger for males and encompassed those of females in two of the three regions analyzed. These results suggest a differential use of winter resources between the sexes, with males typically showing a larger diversity of foraging/migratory strategies. The results also show that dietary differences between male and female Magellanic penguins may occur once the constraints imposed by chick rearing activities cease at the beginning of the post-breeding season.     

Isotopic shifts with size, culture habitat, and enrichment between the diet and tissues of the Japanese scallop Mizuhopecten yessoensis (Jay, 1857)

Use of stable isotope signatures to trace diet patterns in cultured marine bivalves, particularly when changing culture habitat, requires knowledge of the isotopic shift and enrichment between diet and consumer’s tissues. The aim of this study was to determine the patterns of isotope change and the variability of enrichment values (∆δ¹³C and ∆δ¹⁵N) in different tissues (muscle, gonad, digestive gland) of the Japanese scallop (Mizuhopecten yessoensis). It was hypothesized that the isotopic signatures of a consumer’s tissues changed during settlement and that the changes were related to variations in the isotopic signatures of food sources and gut contents. Particular attention was paid to the isotope enrichment between the diet and a consumer’s tissues using isotope analysis of gut content. Muscle δ¹⁵N values decreased significantly 3–5 months post-settlement in a nearshore seabed, concomitant with the ingestion of lower δ¹⁵N food. For juvenile scallops, sinking particles (SP) were considered a more important dietary source than suspended particulate organic matter (SPOM), based on the correspondence between SP and gut contents δ¹³C. Enrichment values (∆δ¹³C and ∆δ¹⁵N) varied with tissue and season. ∆δ¹⁵N was 2.4‰ in muscle, 1.2‰ in gonad, and 0.7‰ in the digestive gland. ∆δ¹³C was 3.2‰ in muscle, 2.3‰ in gonad, and −0.5‰ in the digestive gland. ∆δ¹⁵N was the lowest in summer (0.3‰), and ∆δ¹³C was the highest in autumn (2.8‰). ∆δ¹⁵N values were significantly influenced by age, but not ∆δ¹³C. Patterns of isotope ratios and enrichment values may be related to physiological attributes and differences in diet. This is the first study to demonstrate isotopic shift and enrichment encountered in different tissues of a cultured scallop when changing culture habitat.     

Carbon- and oxygen-isotope composition of the cuttlebone of Sepia officinalis: a tool for predicting ecological information?

Analysis of the isotope composition of calcareous structures of marine organisms has proved useful in providing biological data. The present study constitutes the first detailed work undertaken on the isotope composition of coleoid cephalopods. We analysed the carbon- and oxygen-isotope composition [δ¹³C (CO²⁻ ₃) and δ¹⁸O (CO²⁻ ₃), respectively] of the cuttlebone aragonite of wild and cultivated specimens of Sepia officinalis Linnaeus, 1758. δ¹³C (CO²⁻ ₃) ranged from −2.94 to 1.00‰, δ¹⁸O (CO²⁻ ₃) from −0.18 to 2.08‰. The carbon-isotope composition is not in equilibrium with the carbon species of the ambient seawater, and does not reflect the deposition of CaCO₃ in seawater. The potential influence of environmental factors and biological processes on the carbon-isotope composition of the cuttlebone is discussed. In contrast to δ¹³C, the oxygen-isotope composition of cuttlebone aragonite appears to be in isotopic equilibrium with the ambient seawater. Seasonal changes in isotopic temperature revealed by our analyses agreed with changes in the temperature of the ambient seawater. CaCO₃ was deposited all year round. A maximum life span of 2 yr, a year-round spawning season, and variable growth rates among and within individuals have been inferred from the isotopic temperatures. Received: 14 April 1998 / Accepted: 26 November 1998     

Stable isotopes document winter trophic ecology and maternal investment of adult female southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis>) breeding at the Kerguelen Islands

Individual specialisation is widespread and can affect a population’s ecological and evolutionary dynamics. Whether intra-specific niche differences can influence reproductive investment was examined in a marine mammal, the southern elephant seal (Mirounga leonina), whose females were known to forage in two different areas during the austral winter. The study was conducted at Kerguelen Islands (49°21′S, 70°18′E), southern Indian Ocean, in late winter–early spring 2006. Pups were used as proxies of their mothers’ biology and combined information on their weaning mass (a proxy of females’ foraging success and short-term fitness) together with their blood δ¹³C value (a proxy of female foraging zone). First, the use of isotopic signature of pups was validated to study the female foraging ecology during their pre-breeding trip by demonstrating that δ¹³C and δ¹⁵N values of pups and their mothers were positively and linearly correlated. Then, blood samples were taken from a large number of newly-weaned pups, which were also weighed, to provide information at the population level. Estimated δ¹³C values of female seals encompassed a large range of values (from −23.7 to −19.1‰) with an unimodal frequency distribution, suggesting no contrasted foraging areas within the population. No significant relationship was found between pup weaning mass and their carbon signature, indicating no link between female foraging areas and maternal foraging success and investment. Finally, blood δ¹³C and δ¹⁵N values gave new insights into southern elephant seal ecology, suggesting that females mainly foraged north of the Polar Front where they preyed upon myctophid fish in late winter.