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.     

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.     

Fatty acid signatures of stomach contents reflect inter- and intra-annual changes in diet of a small pelagic seabird,the Thin-billed prion <Emphasis Type="Italic">Pachyptila belcheri</Emphasis>

In diet analyses of seabirds, fatty acid signatures (FAS) can be used to overcome biases due to differential digestion of prey and enable the analysis of very digested diet samples. We applied FAS analysis to stomach contents of a small sub-Antarctic seabird, the Thin-billed Prion Pachyptila belcheri, which feeds mainly on squid during incubation and on crustacea during chick rearing. This seasonal dietary switch of Thin-billed prions was reflected in differences in FAS in regurgitates, as were inter-annual differences in diet composition. A discriminant function analysis correctly classified 93.4% of cases with respect to year (2006–2008) and stage of the breeding cycle (incubation versus chick rearing). The dominant types of crustacea in the diet of Thin-billed prions (amphipods Themisto gaudichaudii, euphausiids, decapods Munida gregaria, and calanoid copepods) were distinguished well by characteristic FAS patterns. However, the FAS of the two main types of prey by volume, amphipods T. gaudichaudii and squid Gonatus antarcticus, were similar to each other. Although FAS were successfully applied in the analysis of prey in stomach contents of prions, FAS of some prey species were similar and may not be distinguishable from each other if used in quantitative diet analyses.     

Food of emperor penguins (Aptenodytes forsteri) in the western Ross Sea, Antarctica

The diet of the emperor penguin Aptenodytes forsteri in the western Ross Sea during spring was investigated by analysis of stomach contents sampled at three different localities. At Cape Washington, emperor penguins feeding chicks consistently preyed on fishes (89 to 95% by mass) and crustaceans (5 to 11%) over the four spring seasons examined. By far the commonest prey was the Antarctic silverfish Pleuragramma antarcticum (89% of the fish prey); the remainder of fish prey were mainly unidentified juveniles of different species of channichthyid fishes. Three species dominated the crustacean part of the diet, i.e. the gammarid amphipods Abyssorchomene rossi/plebs (30% of the crustacean prey) and Eusirus microps (22%), together with the euphausiid Euphausia crystallorophias (24%). At Coulman Island and Cape Roget, fishes, mainly P. antarcticum, formed the bulk of the food (88 and 93% by mass, respectively), crustaceans were minor prey (2.5 and 0.4%), and the squid Psychroteuthis glacialis accounted for a small but significant part of the food (3.5 and 0.8%). This study emphasizes the importance of the small, shoaling pelagic fish Pleuragramma antarcticum as a key link between zooplankton and top predators, including seabirds, in the food web and marine ecosystem of the Ross Sea. Received: 20 May 1997 / Accepted: 8 October 1997     

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.     

Diet of the northern fulmar Fulmarus glacialis: reliance on commercial fisheries?

The massive expansion in breeding numbers and range of the northern fulmar Fulmarus glacialis over the last two centuries is generally attributed to an increased availability of fish offal and discarded fishes from commercial fisheries. This implies that discards should be a major component in fulmar diets in the more recently colonised areas in the south of their range. This paper examines the contemporary diet of the northern fulmar at three major breeding sites, Fair Isle (Shetland, UK), Iceland and Disko Fjord (western Greenland). At Fair Isle, 89% of regurgitates contained fishes, with sandeels (Ammodytidae) recorded in 37%; 32% contained crustaceans (mainly decapods); 8% contained squid. There was also a temporal trend; sandeels declining and crustaceans increasing in the diet from July to August. In Iceland, fishes were also the main prey (47 to 93% of wet mass), with sandeels common in the south and west, and capelin (Mallotus villosus) in the north and east. Other fish species were mainly discards, and together with discarded decapods and fish offal formed 5 to 72% of the diet, depending on the sector. Euphausiids, amphipods and copepods constituted 3 to 13% of the diet. At Disko, 39% of samples contained fishes, 64% contained crustaceans (mainly amphipods), 22% contained squid, and 16% contained pteropods. There was a clear temporal trend, with the bulk of the samples made up of crustaceans in mid-June, capelin from late June to late July, and crustaceans and pteropods from late July to late August. An extensive review of published studies was also carried out. The general pattern was for birds in more southerly populations to consume more discarded fishes, fish offal and benthic invertebrates. However, a considerable proportion of their diet also consisted of juvenile gadids, sandeels, capelin and pelagic zooplankton, which fulmars catch for themselves, and we suggest that breeding adults in the south are less dependent on fishing waste than is generally assumed. Received: 25 September 1998 / Accepted: 7 May 1999     

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.     

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.     

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.     

Inter-annual changes in diet and foraging trip lengths in a small pelagic seabird, the thin-billed prion Pachyptila belcheri

Central place foragers are constrained in their foraging distribution by the necessity to return to their nest site at regular intervals. In many petrels that feed on patchily distributed prey from the sea surface over large foraging areas, alternating long and short foraging trips are used to balance the demands of the chick with the requirements of maintaining adult body condition. When the local conditions are favourable for prey density and quality, adults should be able to reduce the number of long foraging trips. We studied the flexibility in foraging trip lengths of a small pelagic petrel, the thin-billed prion Pachyptila belcheri, over three breeding seasons with increasingly favourable, cold-water conditions. During a warm-water influx in February 2006, chicks were fed less frequently and adults carried out foraging trips of up to 8 days. When conditions became more favourable with colder water temperatures in 2007 and 2008, thin-billed prions decreased trip lengths, more often attended their chick every day, and long foraging trips of six to eight days were not registered during 2008. Chick growth rates mirrored this, as chicks grew poorly during 2006, intermediate during 2007 and best during 2008. Thin-billed prions preyed mainly on squid during incubation and mainly on amphipods and euphausiids during chick-rearing. In the poorest season only, the diet was substantially supplemented with very small copepods. Together, the present results indicate that during warm-water conditions, thin-billed prions had difficulties in finding sufficient squid, amphipods or euphausiids and were forced to switch to lower trophic level prey, which they had to search for over large ocean areas.     

Feeding habits of the albacore tuna <Emphasis Type="Italic">Thunnus alalunga</Emphasis> (Perciformes,Scombridae) from central Mediterranean Sea

In order to assess feeding habits and trophic level of this species, we analysed the stomach content of 189 specimens collected monthly in the southern Tyrrhenian Sea (central Mediterranean Sea). The results showed that this species is a pelagic predator focusing on paralepidid fish and, more in detail, on two species such as Paralepis speciosa and Paralepis coregonoides. Among cephalopods, Heteroteuthis dispar was the most abundant prey item in the stomach contents, whereas among crustaceans the hyperidean amphipods Phrosina semilunata and Brachyscelus crusculum were the most representative species. In the present study, the diet of the albacore did not vary significantly throughout the period of study focusing, in the different season of catch, on the same prey species. Not even a size-related diet shift was observed, with a partitioning of the albacore sample into five size groups. Levin’s standardized index showed a low value indicating a limited trophic niche width, with a diet dominated by a limited number of taxa and then a specialist behaviour for this predator. The estimated TROPH value for T. alalunga resulted very similar to that calculated in Mediterranean Sea for other species as well as Xiphias gladius, Thunnus thynnus, Euthynnus alletteratus and Sarda sarda. Then, the albacore tuna can be considered as a top carnivore.     

Use of multiple tools to assess the feeding preference of coastal dolphins

The feeding preferences of the coastal dolphins Pontoporia blainvillei and Sotalia guianensis in south-eastern Brazil (21o18′S–22o25′S) were assessed through the prey’s index of relative importance (IRI), total mercury concentration (Hg_tot), and stable isotopic (δ¹⁵N and δ¹³C) to compare their efficiency in the discrimination of prey contribution to the predators’ diet. The IRI was the best tool to describe the dolphins’ preference, while Hg_tot and δ¹⁵N seemed to be efficient as a trophic marker when the diet is made up of prey of varying sizes, as observed in S. guianensis. Both dolphins presented lighter δ¹⁵N than their prey species, which is an unusual pattern. However, as the sample size to isotope ratios analysis was small, especially to the dolphins, the results should be considered with caution, and further studies are necessary to corroborate these findings. The δ¹³C values characterized a typical coastal food chain, confirming the preferential area of these species.     

Feeding habits of the Madeira rockfish <Emphasis Type="Italic">Scorpaena maderensis</Emphasis> from central Mediterranean Sea

In order to assess diet composition and niche breadth of this species, we analysed the stomach content of 182 specimens collected monthly along the eastern coast of Sicily (Central Mediterranean Sea). Overall, 50 prey taxa belonging to five major groups (algae, gastropods, crustaceans, polychaetes, fishes) were identified in 102 full stomachs. Benthic or epibenthic crustaceans, such as decapods, amphipods and isopods were the most important prey, whereas algae, gastropods, polychaetes and fishes were only occasionally ingested. In terms of composition by species, the diet of Scorpaena maderensis was characterized by a variety of rare or unimportant prey, which was consumed by few individuals only, although sometimes in large amount. As a result, S. maderensis can be considered a generalized and opportunistic feeder. The feeding intensity followed roughly a seasonal trend, with a minimum food intake in summer. The individual fish size was the most important factor affecting diet. According to the observed ontogenetic shift, small-sized individuals fed primarily on small crustaceans (i.e. amphipods and isopods), whereas large-sized specimens consumed preferably bigger and more vagile prey, such as walking and swimming decapods. No significant differences in diet were observed in relation to sex of predator and sampling season.     

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.     

Trophic relations of the cephalopod Martialia hyadesi (Teuthoidea: Ommastrephidae) at the Antarctic Polar Front,Scotia Sea

Samples of the squid Martialia hyadesi were collected aboard two Japanese squid-jigging vessels carrying out commercial fishing trials at the Antarctic Polar Frontal Zone, north Scotia Sea, in February 1989. The dissected stomachs of 61 specimens were classified according to fullness and the contents were examined visually. Identifiable food items included fish sagittal otoliths, crustacean eyes, the lappets on euphausiid first antennule segments and cephalopod sucker rings. The most frequent items in the squid's diet were the myctophid fishes Krefftichthys anderssoni and Electrona carlsbergi, the euphausiid Euphausia superba and a hyperiid amphipod, probably Themisto gaudichaudi. A small proportion of the sample had been feeding cannibalistically. Total lengths of the fish prey were estimated from sagittal otolith size using published relationships. All fish were relatively small; 7 to 35% of squid mantle-length. However, it is possible that some heads of larger fish are discarded by the squid and so are not represented by otoliths in the stomach contents. Over the size range of squid in the sample there was no relationship between size of fish prey and size of squid. Similarly, when the squid sample was divided into groups according to prey categories: crustaceans, crustaceans+fish, fish, cephalopod, there was no evidence that dietary preference was related to squid size. The prevalence of copepod-feeding myctophids in the diet of this squid, which is itself a major prey item of some higher predators in the Scotia Sea, suggests that a previously unrecognised food chain: copepod-myctophid-M. hyadesi-higher predator, may be an important component of the Antarctic oceanic ecosystem.     

Stable isotopic evidence for trophic groupings and bio-regionalization of predators and their prey in oceanic waters off eastern Australia

Muscle tissue was collected for stable isotope analysis (SIA) from the main fish predators and their fish and cephalopod prey from oceanic waters off eastern Australia between 2004 and 2006. SIA of δ¹⁵N and δ¹³C revealed that the species examined could be divided into three main trophic groups. A “top predator” group consisted mainly of large billfish (Xiphias gladius and Tetrapturus audax), yellowfin (Thunnus albacares), bigeye (T. obesus) and southern bluefin (T. maccoyii) tunas and sharks; with mako (Isurus oxyrinchus) the highest. Below this tier was a second group composed of mid-trophic level fishes including albacore tuna (Thunnus alalunga), lancet fish (Alepisaurus ferox), mahi mahi (Coryphaena hippuris) and ommastrephid squid. Underlying both groups was a grouping of small fishes including myctophids, small scombrids and nomeids as well as surface fishes including macrorhamphosids. These groupings were based largely on mean animal size which showed a positive linear relation to δ¹⁵N (r ² = 0.58). Some species showed significant ontogenetic variation in either δ¹⁵N (swordfish, lancet fish, yellowfin and albacore tuna) or δ¹³C (mako shark). We also noted a consistent latitudinal change in δ¹⁵N and δ¹³C at ~28°S for the top predator species, particularly albacore and yellowfin tuna. The differences were consistent with a change from oligotrophic Coral Sea to nutrient rich Tasman Sea waters. These differences suggest that predatory fishes may have extended residence time in distinct regions off eastern Australia.     

Contrasting C and N isotope ratios from sperm whale skin and squid between the Gulf of Mexico and Gulf of California: effect of habitat

To investigate feeding variation between populations of an apex oceanic predator, stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) have been compared in skin of female and immature male sperm whales (Physeter macrocephalus) from the northern Gulf of Mexico (GoM) and the Gulf of California (GC). Whale sexes were determined genetically. The δ¹³C and δ¹⁵N values from squid muscle were used from the GC, and from inshore and offshore sites in the GoM. We documented contrastingly lower δ¹³C and δ¹⁵N from whales and squid of the GoM compared with those from the GC. While this difference may be associated with variation in trophic position, geographic variation in biochemical cycling influenced significantly the contrasting isotope values between gulfs. Within the northern GoM, the highly distinct δ¹⁵N values of neritic squid versus mesopelagic squid provide further evidence of habitat specificity in δ¹⁵N.     

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.     

Seasonal changes in the diet of a critically endangered seabird and the importance of trawling discards

Pelagic seabirds obtain food from oceans where the availability of their prey changes rapidly both seasonally and spatially. Here, we investigated changes in the trophic habits of the critically endangered Balearic shearwater (Puffinus mauretanicus) through the breeding season and tested for dietary differences between sexes and age classes. We analysed δ¹⁵N and δ¹³C values in blood of adults during the pre-incubation, incubation and chick-rearing periods and of their chicks. Using a two-isotope mixing model, we estimated dietary contributions based on isotope values from potential prey species which included small pelagic species available naturally and demersal fish species available only from trawling discards. Balearic shearwaters showed clear isotopic and dietary variation through the breeding season. During pre-incubation, breeding adults appeared to exploit demersal fish, whereas during the incubation and chick-rearing period, they fed mainly on pelagic anchovies (Engraulis encrasicolus) and pilchards (Sardina pilchardus). Similarly, chicks were fed mainly with anchovies, a resource with a high energetic value. This variation in the dietary habits of adult shearwaters during the breeding season was probably related to both natural and fishery-induced seasonal changes in the availability of potential prey species within their main feeding grounds. However, changes in the nutritional requirements of the shearwaters could also play an important role. Indeed, diet differed between sexes during pre-incubation: females fed less on trawling discards and more on small pelagic fish than males. This sexual segregation in diet could be the consequence of higher nutritional requirements of females during this period. Our study reveals the differential importance of both trawling discards and small pelagic fish species for a pelagic seabird depending on the breeding period and illustrates the importance of considering the entire breeding season when making inferences about the importance of specific prey in seabird dietary studies.     

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.