Stable isotope analysis in two sympatric populations of bottlenose dolphins <Emphasis Type="Italic">Tursiops truncatus</Emphasis>: evidence of resource partitioning?

Skin and muscle from 43 bottlenose dolphins (38 juveniles/adults, 5 calves) stranded in NW Spain were analysed to determine whether stable isotope ratios (δ¹³C and δ¹⁵N) could be used to assess dietary variation, habitat segregation and population substructure. Results were compared with published stomach contents data. Stable isotope ratios from 17 known prey species were also determined. Isotope ratios of the main prey (blue whiting, hake) varied significantly in relation to fish body size. Dolphin calves showed significant heavy isotope enrichments compared to adult females. Excluding calves, δ¹⁵N decreased with increasing dolphin body size, probably related to an ontogenetic shift in diet towards species at lower trophic levels, e.g. on blue whiting as suggested by stomach content results. Bottlenose dolphins were divided into two putative populations (North, South) based on previous genetic studies, and values of δ¹³C and δ¹⁵N differed significantly between these two groups, confirming the existence of population structuring.     

The trophic link between squid and the emperor penguin <Emphasis Type="Italic">Aptenodytes forsteri</Emphasis> at Pointe Géologie,Antarctica

Cephalopod beaks retrieved from stomachs of dead emperor penguin chicks at Pointe Géologie, Terre Adélie, provide information on taxonomic and size composition of the penguin’s squid diet, on the trophic range of the squid species preyed upon and on the fractional trophic impact of the penguin on the whole food web. Emperor penguins prey upon four squid species (Psychroteuthis glacialis, Kondakovia longimana, Gonatus antarcticus, Alluroteuthis antarcticus) and do not take squid larger than 480 mm mantle length. Larger squid live either below the penguin’s diving range or are beyond its handling capacity. Nitrogen stable isotope ratios indicate that squids cover a range of about two trophic levels (2.5–8‰ δ¹⁵N). The impact of the emperor penguin, however, concentrates on the upper part of this range, about 68% of its squid prey being >6‰ δ¹⁵N. The principal components of the emperor’s diet, fish, krill and squid, differ distinctly in average trophic level. Consequently the trophic position of the emperor penguin changes accordingly with diet composition and may differ by almost one trophic level between different emperor penguin colonies.     

δ<Superscript>13</Superscript>C and δ<Superscript>15</Superscript>N values in reef corals <Emphasis Type="Italic">Porites lutea</Emphasis> and <Emphasis Type="Italic">P. cylindrica</Emphasis> and in their epilithic and endolithic algae

In summer 1998, shallow water corals at Sesoko Island, Japan (26°38′N, 127°52′E) were damaged by bleaching. In August 2003, partially damaged colonies of the massive Porites lutea and the branching P. cylindrica were collected at depths of 1.0–2.5 m. The species composition of epilithic algal communities on dead skeletal surfaces of the colonies (‘red turfs’, ‘green turfs’, ‘red crusts’) and the endolithic algae (living in coral skeletons) growing close to and away from living coral polyps was determined. Carbon and nitrogen stable isotope values of organic matter (δ¹³C and δ¹⁵N) from all six of these biological entities were determined. There were no significant differences in the isotope composition of coral tissues of the two corals, with P. lutea having δ¹³C of −15.3 to −9.6‰ and δ¹⁵N of 4.7–6.1‰ and P. cylindrica having similar values. Polyps in both species living close to an interface with epilithic algae had similar isotope values to polyps distant from such an interface. Despite differences in the relative abundance of the algal species in red turfs and crusts, their δ¹³C and δ¹⁵N values were not significantly different from each other (−18.2 to −13.9, −20.6 to −16.2, 1.1–4.3, and 3.3 to 4.9‰, respectively). The green algal turf had significantly higher δ¹³C values (−14.9 to −9.3‰) than that of red turfs and crusts but similar δ¹⁵N (1.2–4.1‰) to the red algae. The data do not suggest that adjoining associations of epilithic algae and coral polyps exchange carbon- and nitrogen-containing metabolites to a significant extent. The endolithic algae in the coral skeletons had δ¹³C values of −14.8 to −12.3‰ and δ¹⁵N of 4.0–5.4‰. Thus they did not differ significantly from the coral polyps in their carbon and nitrogen isotope values. The similarity in carbon isotope values between the coral polyps and endolithic algae may be attributed to a common source of CO₂ for zooxanthellae and endolithic algae, namely, from respiration by the coral host. While it is difficult to fully interpret similarity in the nitrogen isotope composition of coral tissue and of green endolithic algae and the difference in δ¹⁵N between green epilithic and endolithic algae, the data are consistent with nitrogen-containing metabolites from the scleractinian coral serving as a significant source of nitrogen for the endolithic algae.     

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.     

Diet of the social groups of long-finned pilot whales (<Emphasis Type="Italic">Globicephala melas</Emphasis>) in the Strait of Gibraltar

The Strait of Gibraltar is inhabited throughout the year by a group of pilot whales (Globicephala melas), but their spatial distribution varies between Summer and Autumn. In this paper, we have used carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) stable isotope signatures to investigate the differences in diet amongst seasons, sex and stable social units. Skin samples were collected from 56 individually photo-identified pilot whales during Autumn 2005 and Summer 2006. These individuals were genetically sexed and their isotopic signature determined. The level of inter-individual association both within and between stable social units were compared to Euclidean distances between individual isotopes signatures. No differences in either δ¹⁵N or δ¹³C were found according to the sex of individuals, but significant seasonal differences were found in δ¹⁵N, although not in the δ¹³C values. This suggests that pilot whales are resident year round in the Strait, a finding supported by independent photo-identification. The variation in δ¹⁵N could reflect a shift in pilot whale diet through the year, with pilot whales feeding at a higher trophic level in Autumn compared to Summer. This could also represent a change in the diet of pilot whale prey species. The δ¹³C values were significantly different amongst the four stable social units sampled and individual δ¹³C values were significantly related to the level of inter-individual association, while no relationship was found for δ¹⁵N. These results suggest that within the same general area (i.e. the Strait of Gibraltar), there is some level of specialisation in habitat or prey choice between pilot whales social units.     

Spatio-temporal variations in the diet and stable isotope composition of the Argentine hake <Emphasis Type="Italic">Merluccius hubbsi</Emphasis> Marini, 1933 of the continental shelf of southeastern Brazil

The feeding ecology of Merluccius hubbsi was investigated in 2 regions of SE Brazil. The major food sources for the hakes were fish, crustaceans, and squid. In the upwelling system of Cabo Frio, the diet was very similar in the summers of 2001/2002 and spring 2002; fish were the most important prey followed by crustaceans. In Ubatuba, euphausiids were an important prey during the winter 2001 (100 m), while in the summer 2002, fish and amphipods predominated in the diet in the shallower site (40 m) and squid in the deeper site (100 m). The hakes showed temporal differences in stable isotope signatures in both regions, while C:N ratios varied only in Cabo Frio. δ¹⁵N and δ¹³C (bulk and corrected for lipid content) increased with fish length, which seems to be related to the increasing importance of fish and decreasing importance of euphausiids and amphipods in the diet of larger hakes. The mean trophic level of 3.7 for M. hubbsi was estimated using δ¹⁵N of bivalves as baseline and the fractionation of 3.4‰ between trophic levels.     

Influence of nutrients in the feeding ecology of seagrass (Posidonia oceanica L.) consumers: a stable isotopes approach

Nitrogen inputs to coastal environments can considerably alter the abundance of primary producers. However, how herbivores modify their trophic signatures and adjust to changes in food resource conditions remains controversial. Here, we assess the effect of nutrient availability on the diet shifts of the two main Mediterranean herbivores, the Sparid fish Sarpa salpa L. and the sea urchin Paracentrotus lividus (Lmk.) that feed mostly on the seagrass Posidonia oceanica L. (Delile), epiphytes and benthic macroalgae. To do this, we (1) investigate the patterns of isotopic composition (δ¹³C and δ¹⁵N signatures) of the two herbivores and their potential food sources in three areas of contrasting nutrient conditions and, (2) we assess the diet shift along this nutrient gradient by estimating the isotopic nutrient enrichment (i.e., the contribution of δ¹³C and δ¹⁵N signatures in consumers’ tissues relative to potential food sources). Food web signatures of δ¹³C were similar among the three study sites, and no patterns of δ¹³C shift were observed in their diets. In contrast, there was a consistent increase in N contents and δ¹⁵N along the nutrient gradient for all primary producers and their consumers. The rate of δ¹⁵N enrichment was also clearly distinctive between the two herbivores: in P. lividus it increased by 61% along the nutrient gradient, while in S. salpa it remained constant. Our results suggest that sea urchins behave as facultative omnivores and feed on vegetable or mixed diets depending on the trophic status of the system. It is unclear, however, if this modification is behavioral or the consequence of mere changes in the availability of food items, as animal epiphytes (e.g., hydrozoans, bryozoans and ascidians) can also became more abundant on seagrass leaves under increased nutrient conditions. In contrast, adult fish appear to feed on vegetal material independent of nutrient availability in the ecosystem.     

Erratum to: Prey selection by resident common bottlenose dolphins (Tursiops truncatus) in Sarasota Bay, Florida

Prey selection was investigated in wild, resident common bottlenose dolphins, Tursiops truncatus, during the summer months in Sarasota Bay, Florida, USA. Stomach content analyses of 15 dolphins with extensive sighting histories and well-documented distributions were used to determine prey use. Prey availability was assessed by purse seine surveys. We compared the relative abundances of prey available to estimates of prey use at closely matching spatial and temporal scales. G-tests determined that dolphins in this study significantly selected for prey at the species, family, and soniferous/non-soniferous prey levels (G _adj  = 753.98–1,775.93, df = 1–21, p ≤ 0.01). While comprising only 6.3% of the total available prey, soniferous fishes accounted for 51.9% of the total prey consumed. Manly’s standardized forage ratios and 95% Bonferroni confidence intervals determined significant positive selection for soniferous prey and against non-soniferous prey (β^S = 0.9461 vs. β^NS = 0.0539). Dolphins selected against Gerridae, Clupeidae, and Sparidae (β ≤ 0.0014), as well as against all the species within those families (β ≤ 0.0190). It is likely that passive listening for soniferous prey provides an ecological or energetic advantage to cetaceans utilizing this specific foraging technique.     

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.     

Feeding ecology of the grooved tiger shrimp <Emphasis Type="Italic">Penaeus semisulcatus</Emphasis> De Haan (Decapoda: Penaeidae) in inshore waters of Qatar,Arabian Gulf

The feeding ecology of the green tiger shrimp Penaeus semisulcatus was studied in inshore fishing grounds off Doha, Qatar, using a combination of stable isotope (δ¹³C and δ¹⁵N) analysis and gut contents examination. Samples of post-larvae, juvenile and adult shrimp and other organisms were collected from intertidal and subtidal zones during the spawning season (January–June). Shrimp collected from shallow water seagrass beds were mostly post-larvae and juveniles and were significantly smaller than the older juveniles and adults caught in deeper macroalgal beds. Gut content examination indicated that post-larvae and juvenile shrimp in seagrass beds fed mainly on benthos such as Foraminifera, polychaetes, benthic diatoms and small benthic crustaceans (amphipods, isopods and ostracoda), whereas larger shrimp in the macroalgal beds fed mainly on bivalve molluscs and to a lesser extent polychaetes. In shrimp from both seagrass and algal beds, unidentifiable detritus was also present in the gut (18, 32%). δ¹³C values for shrimp muscle tissue ranged from −9.5 ± 0.26 to −12.7 ± 0.05‰, and δ¹⁵N values increased with increasing shrimp size, ranging from 4.1 ± 0.03 to 7.7 ± 0.11‰. Both δ¹⁵N values and δ¹³C values for shrimp tissue were consistent with the dietary sources indicated by gut contents and the δ¹³C and δ¹⁵N values for primary producers and prey species. The combination of gut content and stable isotope data demonstrates that seagrass beds are important habitats for post-larvae and juvenile P. semisulcatus, while the transition to deeper water habitats in older shrimp involves a change in diet and source of carbon and nitrogen that is reflected in shrimp tissue stable isotope ratios. The results of the study confirm the linkage between sensitive shallow water habitats and the key life stages of an important commercially-exploited species and indicate the need for suitable assessment of the potential indirect impacts of coastal developments involving dredging and land reclamation.     

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.     

Alimentary niche partitioning in the Galapagos sea lion, Zalophus wollebaeki

Sea lions are generally considered opportunistic feeders. However, studies from different areas suggest their diet consists mostly of four to five types of prey. Previous studies in Galapagos sea lions have identified at least three feeding strategies for this species, suggesting diversification of their diet. Diet diversification is favored in organisms with relatively high trophic position and subject to high intra-specific and low inter-specific competition. Zalophus wollebaeki meet these criteria as the only pinniped on San Cristobal Island, where three sea lion rookeries are located within 11 km: a distance considerably shorter than their 41 km foraging range. To measure the degree of diet diversification, we used scats and stable isotope analyses. A total of 270 scat samples from lactating females and 142 fur samples from sea lion pups were collected during the breeding season 2006. The scat analysis identified distinct diets among rookeries, with minimal trophic overlap (Cλ = 0.19), a trophic level TL = 4.5 (secondary–tertiary carnivore), and trophic breadth of a specialist predator (B _i  = 0.37). The mean δ¹⁵N and δ¹³C values were 13.07 ± 0.52 and −16.34 ± 0.37, respectively. No significant difference was found in the δ¹⁵N values from the sea lion rookeries, but differences were found inter- and intra-population in δ¹³C values for pups from different groups (ANOVA P < 0.05). Our results indicate that diet diversification is present in the Galapagos sea lion and may play important role to the survival of the species in a habitat where pinniped populations are limited.     

Variability of albacore (<Emphasis Type="Italic">Thunnus alalunga</Emphasis>) diet in the Northeast Atlantic and Mediterranean Sea

This study aims to describe the variability of albacore (Thunnus alalunga) diet in the Northeast Atlantic and Mediterranean Sea and to identify possible relationships between this variability and the features of different feeding areas, the behavior, and the energetic needs of albacore. Stomach contents from albacore caught in five zones of the Bay of Biscay and surrounding waters (n = 654) and three zones of the Mediterranean Sea (n = 152) were analyzed in terms of diet composition and stomach fullness. Carbon and nitrogen stable isotope and C/N ratios were measured for white muscle and liver from albacore in the Bay of Biscay (n = 41) and Mediterranean Sea (n = 60). Our results showed a spatial, seasonal, inter-annual, and size-related variability in the diet of albacore. Albacore diet varied by location in the Mediterranean Sea, with a particularly high proportion of cephalopods, and low δ¹⁵N values in the Tyrrhenian Sea. In the Northeast Atlantic, albacore consumed a higher proportion of crustaceans and a lower proportion of fishes in the most offshore sampling zone than inshore. The digestion states of the major prey reflected a diurnal feeding activity, indicative of feeding in deeper waters offshore, whereas on the continental slope, feeding probably occurred in surface waters at night. Important seasonal and inter-annual diet variability was observed in the southeast of the Bay of Biscay, where preferred albacore prey appeared to be anchovy (Engraulis encrasicolus). Stomach fullness was inversely related to body size, probably reflecting higher energetic needs for smaller individuals. Albacore from the Bay of Biscay had significantly lower δ¹³C and higher δ¹⁵N values compared with albacore from the Mediterranean Sea, indicative of regional baseline shifts, and trophic position and muscle lipid stores in albacore increased with body size.     

A rapid ontogenetic shift in the diet of juvenile yellowfin tuna from Hawaii

Within the tropical and subtropical oceans, tuna forage opportunistically on a wide variety of prey. However, little is known about the trophic ecology of the smallest size classes which play an important role in stock assessments and fisheries management. The foraging behavior of yellowfin tuna, Thunnus albacares (23.5–154.0 cm FL), collected from nearshore Fish Aggregating Devices (FADs) around Oahu was studied using stable isotope and stomach contents analyses. Emphasis was placed on small juveniles. Yellowfin tuna changed their diets significantly between 45 and 50 cm forklength (ca. 1.5 kg). Smallest size classes fed on planktonic organisms inhabiting the shallow mixed layer, primarily larval stomatopod and decapod crustaceans, whereas larger tuna fed on teleosts and adult Oplophorus gracilirostris, a vertically migrating mesopelagic species of shrimp. When interpreting the variation in prey δ ¹⁵N values, we considered both their relative trophic position and δ ¹⁵N values of the nitrogen at the base of the food web. Based on the distinct diet shift of the yellowfin tuna, demonstrated by both isotope and stomach content analyses, we propose a critical mass threshold was reached at about 45 cm FL that enabled sufficient endothermic capability to allow tuna to access prey dwelling in deeper, colder water. These ontogenetic changes in foraging range and commensurate shift in diet of small tunas would affect their vulnerability to fishing pressure.     

Comparative foraging ecology and ecological niche of a superabundant tropical seabird: the sooty tern Sterna fuscata in the southwest Indian Ocean

Over 6-million pairs of sooty terns Sterna fuscata breed once a year in the southwest Indian Ocean, mostly on three islands of the Mozambique Channel (Europa, Juan de Nova and Glorieuses) and in the Seychelles region. Seasonal reproduction in either winter or summer is the dominant strategy in the area, but non-seasonal reproduction also occurred in some places like at Glorieuses Archipelago. The feeding ecology of the sooty tern was investigated during the breeding seasons to determine whether terns showed significant differences in their trophic ecology between locations. Regurgitations were analyzed to describe the diet of individuals when breeding, and stable isotopes and mercury concentrations were used to temporally integrate over the medium-term of the trophic ecology of both adults and chicks. Overall, the diet was composed of fish, flying squid and fish larvae in different proportions. At Europa and Aride in the Seychelles, where winter reproduction occurs, large epipelagic prey like flying fish or squid dominated the diet. At Juan de Nova, sooty terns reproduce in summer and rely mostly on fish larvae. At Glorieuses (non-seasonal breeding), the diet was intermediate with fish larvae and flying squid being important prey items. The stable-carbon and nitrogen isotope values in blood confirm the differences observed in dietary analysis, and demonstrate different feeding strategies between colonies. δ¹³C values of feathers showed spatial segregation between birds from the Mozambique Channel and the Seychelles region. Terns from the Seychelles had also higher δ¹⁵N values. Feather δ¹³C values also suggest a significant shift from summer to wintering habitat for birds from Juan de Nova. This study emphasizes the high phenotypic plasticity of the species, which may explain its numerical dominance in all tropical waters of the World’s Ocean.     

Foraging ecology of leatherback sea turtles in the Western North Atlantic determined through multi-tissue stable isotope analyses

Leatherback turtles, Dermochelys coriacea, are highly migratory, spending most of their lives submerged or offshore where their feeding habits are difficult to observe. In order to elucidate the foraging ecology of leatherbacks off Massachusetts, USA, stable isotope analyses were performed on leatherback tissues and prey collected from 2005 to 2009. Stable isotope ratios of nitrogen and carbon were determined in whole blood, red blood cells, blood plasma, muscle, liver, and skin from adult male, female, and subadult leatherbacks. Isotopic values were analyzed by body size (curved carapace length) and grouped by sex, and groups were tested for dietary differences. Gelatinous zooplankton samples were collected from leatherback foraging grounds using surface dip nets and stratified net tows, and prey contribution to leatherback diet was estimated using a two-isotope Bayesian mixing model. Skin and whole blood δ¹³C values and red blood cell δ¹⁵N values were correlated with body size, while δ¹³C values of red blood cells, whole blood, and blood plasma differed by sex. Mixing model results suggest that leatherbacks foraging off Massachusetts primarily consume the scyphozoan jellyfishes, Cyanea capillata and Chrysaora quinquecirrha, and ctenophores, while a smaller proportion of their diet comes from holoplanktonic salps and sea butterflies (Cymbuliidae). Our results are consistent with historical observations of leatherback turtles feeding on scyphozoan prey in this region and offer new insight into size- and sex-related differences in leatherback diet.     

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.     

Central place foraging by breeding Cook’s petrel Pterodroma cookii: foraging duration reflects range, diet and chick meal mass

Pelagic seabirds are central place foragers during breeding and variation in foraging trip duration and range reflect differences in diet and chick provisioning, through the exploitation of divergent habitats of varying productivity. We tested whether these relationships hold in small procellarriids by equipping chick-rearing Cook’s petrel Pterodroma cookii (200 g) with geolocation-immersion loggers, conducting isotope analysis of blood and measuring chick meal mass following foraging trips of varying duration. Cook’s petrel tracked during chick rearing from Little Barrier Island (LBI) and Codfish Island (CDF), New Zealand had larger maximum ranges during longer foraging trips. Blood nitrogen isotope signatures (δ¹⁵N) of adults were significantly higher after foraging trips of longer duration, but not of greater maximum range. There was no significant relationship between blood carbon isotope signatures (δ¹³C) and foraging trip characteristics. Proportion of time spent on the sea surface and the mass of the meal brought back to chicks were consistently greater for Cook’s petrel with larger maximum ranges, which in the case of birds from CDF coincided with productive subtropical convergence zone habitats. As predicted, trip duration reflected divergent foraging behaviours in Cook’s petrel during breeding. We suggest that the availability of different prey is a key factor governing at-sea distributions and dietary composition of this species.     

Delayed ontogenic dietary shift and high levels of omnivory in green turtles (<Emphasis Type="Italic">Chelonia mydas</Emphasis>) from the NW coast of Africa

Young green turtles (Chelonia mydas) spend their early lives as oceanic omnivores with a prevalence of animal prey. Once they settle into neritic habitats (recruitment), they are thought to shift rapidly to an herbivorous diet, as revealed by studies in the Greater Caribbean. However, the precise timing of the ontogenic dietary shift and the actual relevance of animal prey in the diet of neritic green turtles are poorly known elsewhere. Stable isotopes of carbon, sulfur and nitrogen in the carapace scutes of 19 green turtles from Mauritania (NW Africa), ranging from 26 to 102 cm in curved carapace length (CCLmin), were analyzed to test the hypothesis of a rapid dietary shift after recruitment. Although the length of residence time in neritic habitats increased with turtle length, as revealed by a significant correlation between turtle length and the δ¹³C and the δ³⁴S of the scutes, comparison of the δ¹⁵N of the innermost and outermost layers of carapace scutes demonstrated that consumption of macrophytes did not always start immediately after recruitment, and turtles often resumed an animal-based diet after starting to graze on seagrasses. As a consequence, seagrass consumption did not increase gradually with turtle size and animal prey largely contributed to the diet of turtles within the range 29–59 cm CCLmin (76–99% of assimilated nutrients). Seagrass consumption by turtles larger than 59 cm CCLmin was higher, but they still relied largely on animal prey (53–76% of assimilated nutrients). Thus, throughout most of their neritic juvenile life, green turtles from NW Africa would be better classified as omnivores rather than herbivores. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.