Flexible foraging strategies of gentoo penguins <Emphasis Type="Italic">Pygoscelis papua</Emphasis> over 5 years in the South Shetland Islands,Antarctica

Gentoo penguins Pygoscelis papua show considerable plasticity in their diet, diving, and foraging behaviors among colonies; we expected that they might exhibit similar variability over time, at a single site, since flexible foraging habits would provide a buffer against changes in prey availability. We examined interannual changes in the foraging strategies and diet of gentoo penguins in the South Shetland Islands, Antarctica, over 5 years with variable prey abundance. Antarctic krill Euphausia superba was the primary diet item, and fish the secondary, though the importance of these items varied among years. Diving behavior also varied over time: different dive depth distributions were observed among years. Nonetheless, chick-rearing success remained relatively constant, indicating that gentoo penguins were able to maintain chick provisioning by altering their foraging strategy among years. Variable abundance of krill in the region did not have observable impacts on the diet, foraging behaviors or chick-rearing success of gentoo penguins. We suggest that foraging plasticity may be one reason that gentoo penguin populations have remained stable in the region, while their congeners (P. antarctica and P. adeliae) with less flexible foraging strategies have declined.     

Are penguins and seals in competition for Antarctic krill at South Georgia?

The Antarctic fur seal (Arctocephalus gazella) and macaroni penguin (Eudyptes chrysolophus) are sympatric top predators that occur in the Southern Ocean around South Georgia where they are, respectively, the main mammal and bird consumers of Antarctic krill (Euphausia superba). In recent years the population of fur seals has increased, whereas that of macaroni penguins has declined. Both species feed on krill of similar size ranges, dive to similar depths and are restricted in their foraging range at least while provisioning their offspring. In this study we test the hypothesis that the increased fur seal population at South Georgia may have resulted in greater competition for the prey of macaroni penguins, leading to the decline in their population. We used: (1) satellite-tracking data to investigate the spatial separation of the Bird Island populations of these two species whilst at sea during the breeding seasons of 1999 and 2000 and (2) diet data to assess potential changes in their trophic niches between 1989 and 2000. Foraging ranges of the two species showed considerable overlap in both years, but the concentrations of foraging activity were significantly segregated spatially. The size of krill taken by both species was very similar, but over the last 12 years the prevalence of krill in their diets has diverged, with nowadays less krill in the diet of macaroni penguins than in that of Antarctic fur seals. Despite a significant degree of segregation in spatial resource use by the study populations, it is likely that the South Georgia populations of Antarctic fur seal and macaroni penguin exploit the same krill population during their breeding season. For explaining the opposing population trends of the two species, the relative contributions of independent differential response to interannual variation in krill availability and of interspecies competition cannot be resolved with available evidence. The likely competitive advantage of Antarctic fur seals will be enhanced as their population continues to increase, particularly in years of krill scarcity.     

Chinstrap penguins alter foraging and diving behavior in response to the size of their principle prey, Antarctic krill

Penguins may exhibit plasticity in their diving and foraging behaviors in response to changes in prey availability. Chinstrap penguins are dependent predators of Antarctic krill in the Scotia Sea region, but krill populations have fluctuated in recent years. We examined the diet of chinstrap penguins at Livingston Island, South Shetland Islands, in relation to their diving and foraging behavior using time-depth recorders over six breeding seasons: 2002–2007. When krill were smaller, more chinstrap penguins consumed fish. In these years, chinstrap penguins often exhibited a shift to deep dives after sundown, and then resumed a shallower pattern at sunrise. These night dives were unexpectedly deep (up to 110 m) and mean night dive depths sometimes exceeded those from the daytime. The average size of krill in each year was negatively correlated to mean night dive depths and the proportion of foraging trips taken overnight. Based on these patterns, we suggest that when krill were small, penguins increasingly targeted myctophid fish. The average krill size was negatively correlated to the time chinstrap penguins spent foraging which suggests that foraging on smaller krill and fish incurred a cost: more time was spent at sea foraging.     

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     

Modelling predation by transient leopard seals for an ecosystem-based management of Southern Ocean fisheries

Correctly quantifying the impacts of rare apex marine predators is essential to ecosystem-based approaches to fisheries management, where harvesting must be sustainable for targeted species and their dependent predators. This requires modelling the uncertainty in such processes as predator life history, seasonal abundance and movement, size-based predation, energetic requirements, and prey vulnerability. We combined these uncertainties to evaluate the predatory impact of transient leopard seals on a community of mesopredators (seals and penguins) and their prey at South Georgia, and assess the implications for an ecosystem-based management. The mesopredators are highly dependent on Antarctic krill and icefish, which are targeted by regional fisheries. We used a state-space formulation to combine (1) a mark-recapture open-population model and individual identification data to assess seasonally variable leopard seal arrival and departure dates, numbers, and residency times; (2) a size-based bioenergetic model; and (3) a size-based prey choice model from a diet analysis. Our models indicated that prey choice and consumption reflected seasonal changes in leopard seal population size and structure, size-selective predation and prey vulnerability. A population of 104 (90–125) leopard seals, of which 64% were juveniles, consumed less than 2% of the Antarctic fur seal pup production of the area (50% of total ingested energy, IE), but ca. 12–16% of the local gentoo penguin population (20% IE). Antarctic krill (28% IE) were the only observed food of leopard seal pups and supplemented the diet of older individuals. Direct impacts on krill and fish were negligible, but the “escapement” due to leopard seal predation on fur seal pups and penguins could be significant for the mackerel icefish fishery at South Georgia. These results suggest that: (1) rare apex predators like leopard seals may control, and may depend on, populations of mesopredators dependent on prey species targeted by fisheries; and (2) predatory impacts and community control may vary throughout the predator's geographic range, and differ across ecosystems and management areas, depending on the seasonal abundance of the prey and the predator's dispersal movements. This understanding is important to integrate the predator needs as natural mortality of its prey in models to set prey catch limits for fisheries. Reliable estimates of the variability of these needs are essential for a precautionary interpretation in the context of an ecosystem-based management.     

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.     

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.     

Temporal variation in Adélie penguin diet at Béchervaise Island,east Antarctica and its relationship to reproductive performance

Diet, and in particular, food quality and quantity can influence the reproductive performance of marine predators. Also, the diet of specialist predators is often monitored in programmes that model and manage ecosystems. We examined the diet of Adélie penguins (Pygoscelis adeliae), an important consumer of Southern Ocean living resources, at Béchervaise Island, east Antarctica, during the chick-rearing periods for 11 years between 1991–1992 and 2002–2003. We also investigated the relationship between diet and annual reproductive performance. Substantial inter- and intra-annual variation in both meal mass and composition was evident: adults generally returned with larger food loads during the crèche compared with the guard stages, and diet composition was dominated by two prey types, krill and fish, which combined contributed to >90% of the diet by mass in 7 out of 11 years. Females generally brought back meals dominated by krill; males generally consumed fish-dominated meals. However, both sexes returned with a high proportion of krill when annual mean meal mass was also high, suggesting that more food was available in high krill years. There was also evidence that years of high reproductive performance were positively correlated with years of both high meal and krill mass. We believe that our results indicate that there is significant long-term inter- and intra-annual variability in the amount of food available to Adélie penguins and that this was reflected in their diet and measures of reproductive performance. Coupled with the observation that penguins did not switch prey, this indicates that Adélie penguins from Béchervaise Island are dependent predators of krill. This contrasts with populations in other locations but supports the notion that Adélie penguins are an informative species to monitor the management of Southern Ocean marine living resources in this region.     

Diet of the Humboldt penguin (<Emphasis Type="Italic">Spheniscus humboldti</Emphasis>) in northern and southern Chile

The diet of the Humboldt penguin (Spheniscus humboldti) was examined and compared in two colonies in Chile. Field work was conducted on Pan de Azúcar Island in northern Chile in the breeding season 1998/1999 and on the Puñihuil Islands in southern Chile over two successive breeding seasons during 1997/1998 and 1998/1999. Penguin diet was studied by stomach-pumping birds and analysed by species composition, size and mass of prey. Fish were the dominant prey item at both sites, the contribution of cephalopods and crustaceans varying between sites. The fish prey consisted predominantly of school fish, but there were clear latitudinal differences in fish prey taken. Penguins in the northern colony consumed primarily garfish (Scomberesox saurus), while birds at the southern colony of Puñihuil fed primarily on anchovy (Engraulis ringens), Araucanian herring (Strangomera bentincki) and silverside (Odontesthes regia). The results showed significant differences in terms of numbers of fish taken between the two breeding seasons at Puñihuil. In 1997/1998 penguins consumed almost exclusively anchovy, while they fed primarily on silversides in the successive year. Almost all prey, except stomatopods, were characterised as being pelagic species that occur in relatively inshore water, consistent with the foraging behaviour of Humboldt penguins. The dependence of Humboldt penguins on commercially exploited, schooling prey species makes the species particularly susceptible to changes in prey stocks, due to non-sustainable fisheries management.Communicated by O. Kinne, Oldendorf/Luhe     

Geospatial variability of krill and top predators within an Antarctic submarine canyon system

The spatial distribution patterns of krill, seabirds (penguin, petrel and albatross), fur seals and baleen whales were mapped in nearshore waters (<50 km from land) to investigate their habitat selection within two adjacent submarine canyons near Livingston Island, Antarctica. Three shipboard surveys were conducted (February 2005–2007), and an echosounder was used to measure the distribution and abundance of krill while simultaneously conducting visual surveys to map seabird and marine mammals. Using a multispecies approach, we test the hypothesis that spatial organization of krill and top predators co-vary according to fine-scale changes in bathymetry in the nearshore marine environment. GAMs are used to examine the effect of sea depth, slope and distance to isobaths on the spatial distribution and abundance of krill and predators. Spatial distribution patterns of krill and predators relate to fine-scale (1–10 km) changes in bathymetry and exhibit cross-shelf gradients in abundance. Krill were concentrated along the shelf-break and abundant within both submarine canyons. Predators exhibited different preferences for locations within the submarine canyon system that relates to their foraging behavior. Penguins concentrated closer to shore and within the head of the east submarine canyon immediately adjacent to a breeding colony. Whales were also concentrated over the head of the east canyon (overlapping with penguins), whereas albatrosses and fur seals were concentrated in the west canyon. Fur seals also showed preference for steep slopes and were concentrated along the shelf-break. Petrels exhibited peaks in abundance throughout both submarine canyons. Owing to their orientation, size and proximity to the coastline, submarine canyons provide important habitat heterogeneity for krill and a variety of predators. This study highlights the multispecies approach for studying spatial ecology of top predators and krill and has implications for marine spatial management of the Scotia Sea.     

Depth utilisation by breeding Adélie penguins,Pygoscelis adeliae,at Esperanza Bay,Antarctica

Miniature depth gauges were attached in December 1987 and January 1988 to Adélie penguins,Pygoscelis adeliae, breeding at Esperanza on the Antarctic Peninsula. Results from 34 birds showed that foraging penguins with eggs and with brooded and crèching chicks spent mean periods away from the nest of 96, 36 and 21 h, respectively, during which time means of 29.0 h (30%), 11.2 h (31%) and 2.7 h (13%), respectively, were spent under water at depths > 5 m. Time under water was positively correlated with time absent from nest. Maximum depth reached was 170 m but overall birds spent most time at shallower depths. Birds foraging for brooded chicks dived deeper than birds foraging for crèching chicks. Stomach-pumping indicated that the principal prey caught at this time was krill,Euphausia superba. Mean mass changes of adults during single foraging trips indicated that krill were caught at a mean rate of 7.2 g min^–1 spent under water.     

Exploring the effects of reductions in krill biomass in the Southern Ocean on blue whales using a state-dependent foraging model

Many species of baleen whales were hunted to near extinction in the Southern Hemisphere. The recovery of these populations will be affected by the availability of krill, a major dietary component, in the Southern Ocean. We combine a novel energetics model for baleen whales with a state dependent foraging model to explore the impacts of an expanding krill fishery on baleen whales. We parameterize the model for blue whales, but with simple modifications it could be applied to most baleen whales. We predict that an expanding fishery will have a small but significant impact on the blue whale population through decreased birth rates. However, spreading the catch limit throughout the range of krill can reduce these effects. In addition, whales may be able to reduce these impacts through adaptive changes in foraging behavior. The relationship between krill abundance and blue whale foraging and reproductive success is nonlinear, such that larger reductions in krill biomass, potentially following a loss of sea ice due to climate change, could have a much larger negative impact on the recovery of blue whales.     

Higher trophic level prey does not represent a higher quality diet in a threatened seabird: implications for relating population dynamics to diet shifts inferred from stable isotopes

Diet quality is a key determinant of population dynamics. If a higher trophic level, more fish-based diet is of higher quality for marine predators, then individuals with a higher trophic level diet should have a greater body mass than those feeding at a lower trophic level. We examined this hypothesis using stable isotope analysis to infer dietary trophic level and foraging habitat over three years in eastern rockhopper penguins Eudyptes chrysocome filholi on sub-Antarctic Campbell Island, New Zealand. Rockhopper penguins are ‘Vulnerable’ to extinction because of widespread and dramatic population declines, perhaps related to nutritional stress caused by a climate-induced shift to a lower trophic level, lower quality diet. We related the stable nitrogen (δ¹⁵N) and carbon (δ¹³C) isotope values of blood from 70 chicks, 55 adult females, and 55 adult males to their body masses in the 2010, 2011, and 2012 breeding seasons and examined year, stage, age, and sex differences. Opposite to predictions, heavier males consumed a lower trophic level diet during incubation in 2011, and average chick mass was heavier in 2011 when chicks were fed a more zooplankton-based, pelagic/offshore diet than in 2012. Contrary to the suggested importance of a fish-based diet, our results support the alternative hypothesis that rockhopper penguin populations are likely to be most successful when abundant zooplankton prey are available. We caution that historic shifts to lower trophic level prey should not be assumed to reflect nutritional stress and a cause of population declines.     

Linking predator and prey behaviour: contrasts between Antarctic fur seals and macaroni penguins at South Georgia

Antarctic fur seals Arctocephalus gazella and macaroni penguins Eudyptes chrysolophus are the two main land-based krill Euphausia superba consumers in the northern Scotia Sea. Using a combination of concurrent at-sea (predator observations, net hauls and multi-frequency acoustics), and land-based (animal tracking and diet analysis) techniques, we examined variability in the foraging ecology of these sympatric top predators during the austral summer and autumn of 2004. Krill availability derived from acoustic surveys was low during summer, increasing in autumn. During the breeding season, krill occurred in 80% of fur seal diet samples, with fish remains in 37% of samples. Penguin diets contained the highest proportion of fish in over 20 years of routine monitoring (46% by mass; particularly the myctophid Electrona antarctica), with krill (33%) and amphipods (Themisto gaudichaudii; 21%) also occurring. When constrained by the need to return and feed their offspring both predator species foraged to the northwest of South Georgia, consistent with an area of high macrozooplankton biomass, but fur seals were apparently more successful at exploiting krill. When unconstrained by chick-rearing (during March) penguins foraged close to the Shag Rocks shelf-break, probably exploiting the high daytime biomass of fish in this area. Penguins and seals are able to respond differently to periods of reduced krill abundance (in terms of variability in diet and foraging behaviour), without detriment to the breeding success of either species. This highlights the importance of myctophid fish as an alternative trophic pathway for land-based predators in the Scotia Sea ecosystem.     

How age and sex drive the foraging behaviour in the king penguin

As predicted by life history theory, once recruited into the breeding population and with increasing age, long-lived animals should be able to manage more efficiently the conflict between self-maintenance and reproduction. Consequently, breeding performances should improve with age before stabilizing at a certain level. Using temperature–depth recorders and isotopic analysis, we tested how age affects the foraging behaviour of king penguin Aptenodytes patagonicus during one trip in the chick-rearing phase. Depending on sex and age, king penguins expressed two different foraging strategies. Older birds gained more daily mass per unit body mass than younger ones. Older females conducted shorter trips, dived deeper and performed more prey pursuits. They also had higher blood levels of δ¹⁵N than younger individuals and males indicating sex- and age-specific dietary regimes. However, we found no differences in carbon isotopic signature, suggesting that individuals exploited the same foraging areas independently of sex and age. Our results suggest that king penguins are able to increase the quantity of energy extracted with increasing age and that such a strategy is sex-related. Our study is the first to reveal of an interaction between age and sex in determining foraging efficiency in king penguins.     

Evaluating the relative importance of intrinsic and extrinsic factors on the foraging activity of top predators: a case study on female little penguins

A complex interaction of biotic and abiotic factors influences animal foraging activity. It is often difficult to understand which factors may affect animals’ foraging and how it is affected. For instance, whereas the effect of sexual dimorphism on foraging activity has been reported in several species, little is known of the complex interactions between variables acting at a finer scale, e.g. the variability of body mass within a sex. Evaluating the importance of these finer scale factors is also essential to the understanding of foraging behaviour. We propose here a simple approach by applying principal component analysis (PCA) in a novel way to examine relationships between biotic and abiotic factors affecting foraging behaviour of top predators. We studied female little penguins (Eudyptula minor) of known age, carrying miniature accelerometers during the guard stage of breeding. Surprisingly, the body mass of the females did not influence any of the foraging parameters, but females foraging later in the breeding season dived shallower and more often, showing a strong correlation with laying date. Similarly, the diving effort of females was greater with increasing chick age within the same breeding stage. These results indicate that for female little penguin, the relationship between changes in prey availability and hunting effort can change at a fine scale, within a breeding stage. Therefore, any analysis of little penguin foraging behaviour during breeding should consider the timing in relation to the breeding season. We encourage researchers to develop the use of this PCA approach as it could help clarify the complexity of the underlying mechanisms determining foraging activity and we propose that it should be used as a first step of foraging behaviour analysis, before examining a particular relationship.     

Comparison of diving behavior and foraging habitat use between chinstrap and gentoo penguins breeding in the South Shetland Islands, Antarctica

Chinstrap, Pygoscelis antarctica, and gentoo, P. papua, penguins are sympatric species that inhabit the Antarctic Peninsula. To evaluate differences in the foraging habitat of these two species, we recorded their foraging locations and diving behavior using recently developed GPS-depth data loggers. The study was conducted on King George Island, Antarctica during the chick-guarding period of both species, from December 2006 to January 2007. The area used for foraging, estimated as the 95% kernel density of dive (>5 m) locations, overlapped partially between the two species (26.4 and 68.5% of the area overlapped for chinstrap and gentoo penguins, respectively). However, the core foraging area, estimated as the 50% kernel density, was mostly separate (12.8 and 25.0% of the area overlapped for chinstrap and gentoo penguins, respectively). Chinstrap penguins tended to use off-shelf (water depth > 200 m) regions (77% of the locations for dives >5 m), whereas gentoo penguins mainly used on-shelf (water depth < 200 m) areas (71% of dive locations). The data on foraging locations, diving behavior, and bathymetry indicated that gentoo penguins often performed benthic dives (28% of dives >5 m), whereas chinstrap penguins almost always used the epipelagic/mid-water layer (96% of dives >5 m). Diving parameters such as diving bottom duration or diving efficiency differed between the species, reflecting differences in the use of foraging habitat. The diving parameters also suggested that the on-shelf benthic layer was profitable foraging habitat for gentoo penguins. Conversely, the relationship between trip duration, date, and stomach content mass suggested that the chinstrap penguins went further from the colony to forage as the season progressed, possibly reflecting a reduction in prey availability near the colony. Our results suggest that chinstrap and gentoo penguins segregated their foraging habitat in the Antarctic coastal marine environment, possibly due to inter- and intra-specific competition for common prey resources.     

Fatty acid evidence for the importance of myctophid fishes in the diet of king penguins, Aptenodytes patagonicus

We tested the usefulness of the fatty acid signature-method in investigating the diet of seabirds in conjunction with the conventional technique of stomach-content analysis. We compared the fatty acid composition of subcutaneous white adipose tissue (SWAT) of king penguin chicks (Aptenodytes patagonicus) during fattening periods to that of total lipids from their food. In both spring and autumn, the fatty acid composition of chick SWAT was identical to that of the dietary lipids. Because the diet of adult king penguins feeding for self-maintenance (i.e. not for their chicks) was essentially unknown, we subsequently analysed their SWAT fatty acid patterns after premolting and prebreeding foraging trips (during which they build up large energy reserves). The fatty acid composition of SWAT from adults was identical to that of chick adipose tissue and food. King penguin diet and SWAT were characterized by high levels of very long-chain mono-unsaturated fatty acids (20 to 24 carbon atoms, 16 to 23% by mass) and (n-3) poly-unsaturated fatty acids (19 to 27%); these consisted mainly of 20:1n-9 (5 to 8%) and 22:1n-11 (5 to 8%), and 22:6n-3 (10 to 13%) and 20:5n-3 (3 to 9%), respectively. Prey items identified from chick stomach contents indicated that the bulk of the food was oceanic myctophid fishes, mainly Electrona carlbergi, Krefftichthys anderssoni and Protomyctophum tenisoni. The fatty acid composition of four other species of myctophid fishes was similar to that of penguin diet and SWAT, but markedly different from that measured for a squid species and that reported for crustaceans. These findings indicate that adult king penguins prey on myctophid fish not only to feed their chicks but also for their own nutrition. The fatty acid signature-technique is therefore a reliable method to gain information on the food and feeding ecology of seabirds when more conventional techniques are of limited value. Such information is important to the understanding of trophic relationships between key species of the ecosystems, and also to provide insight into the nature of avian adaptations to the marine environment. Received: 11 December 1997 / Accepted: 25 July 1998     

A Bayesian hierarchical model of Antarctic fur seal foraging and pup growth related to sea ice and prey abundance

We created a Bayesian hierarchical model (BHM) to investigate ecosystem relationships between the physical ecosystem (sea ice extent), a prey measure (krill density), predator behaviors (diving and foraging effort of female Antarctic fur seals, Arctocephalus gazella, with pups) and predator characteristics (mass of maternal fur seals and pups). We collected data on Antarctic fur seals from 1987/1988 to 1994/1995 at Seal Island, Antarctica. The BHM allowed us to link together predators and prey into a model that uses all the data efficiently and accounts for major sources of uncertainty. Based on the literature, we made hypotheses about the relationships in the model, which we compared with the model outcome after fitting the BHM. For each BHM parameter, we calculated the mean of the posterior density and the 95% credible interval. Our model confirmed others' findings that increased sea ice was related to increased krill density. Higher krill density led to reduced dive intensity of maternal fur seals, as measured by dive depth and duration, and to less time spent foraging by maternal fur seals. Heavier maternal fur seals and lower maternal foraging effort resulted in heavier pups at 22 d. No relationship was found between krill density and maternal mass, or between maternal mass and foraging effort on pup growth rates between 22 and 85 days of age. Maternal mass may have reflected environmental conditions prior to the pup provisioning season, rather than summer prey densities. Maternal mass and foraging effort were not related to pup growth rates between 22 and 85 d, possibly indicating that food was not limiting, food sources other than krill were being used, or differences occurred before pups reached age 22 d.     

Diet of young Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) in eastern and western Atlantic foraging grounds

Atlantic bluefin tuna (Thunnus thynnus) are highly migratory predators whose abundance, distribution, and somatic condition have changed over the past decades. Prey community composition and abundance have also varied in several foraging grounds. To better understand underlying food webs and regional energy sources, we performed stomach content and stable isotope analyses on mainly juvenile (60–150 cm curved fork length) bluefin tuna captured in foraging grounds in the western (Mid-Atlantic Bight) and eastern (Bay of Biscay) Atlantic Ocean. In the Mid-Atlantic Bight, bluefin tuna diet was mainly sand lance (Ammodytes spp., 29% prey weight), consistent with historic findings. In the Bay of Biscay, krill (Meganyctiphanes norvegica) and anchovy (Engraulis encrasicolus) made up 39% prey weight, with relative consumption of each reflecting annual changes in prey abundance. Consumption of anchovies apparently declined after the local collapse of this prey resource. In both regions, stable isotope analysis results showed that juvenile bluefin tuna fed at a lower trophic position than indicated by stomach content analysis. In the Mid-Atlantic Bight, stable isotope analyses suggested that >30% of the diet was prey from lower trophic levels that composed <10% of the prey weights based upon traditional stomach content analyses. Trophic position was similar to juvenile fish sampled in the NW Atlantic but lower than juveniles sampled in the Mediterranean Sea in previous studies. Our findings indicate that juvenile bluefin tuna targeted a relatively small range of prey species and regional foraging patterns remained consistent over time in the Mid-Atlantic Bight but changed in relation to local prey availability in the Bay of Biscay.