Dietary segregation of krill-eating South Georgia seabirds

The diets of six of the main seabird species (two petrels, two albatrosses, two penguins) breeding at Bird Island, South Georgia were studied simultaneously during the chick-rearing period in 1986. For five species, Antarctic krill Euphausia superba was the main food (39–98% by mass); grey-headed albatrosses took mainly the ommastrephid squid Martialia hyadesi (71%) and only 16% krill. The size of the krill taken was similar between seabird species, although there were small but significant differences between penguins and the other species. Sex and reproductive status of krill, however, was different between all seabird species, reflecting some combination of differences in foraging ranges, selectivity by predators, or differences in escape responses of krill. For the krill-eating species, the rest of the diet varied substantially between species, comprising Martialia and nototheniid fish (blackbrowed albatross and, along with lanternfish, white-chinned petrel), lanternfish and amphipods (Antarctic prion and macaroni penguin), and icefish (gentoo penguin). Long-term data on breeding success and information on diet in 5–10 other years suggest that in 1986 seabird diet and reproductive performance was indicative of a year of good availability of krill around South Georgia. In such circumstances, ecological segregation between krill-eating species appears to be maintained chiefly by differences in foraging range and feeding methods, which are reviewed. This situation is rather different from the few studies of seabird communities elsewhere, where prey type and size are believed to be the main mechanisms of dietary segregation.     

THE FOOD AND FEEDING ECOLOGY OF GREY-HEADED ALBATROSS DIOMEDEA CHRYSOSTOMA AND BLACK-BROWED ALBATROSS D. MELANOPHRIS

The food and feeding ecology of Black-browed Albatrosses and Grey-headed Albatrosses was studied from 1975 to 1978 at Bird Island, South Georgia. Two hundred and seventy food samples (averaging 75–85 % by weight of the mean chick feed) were collected from adults of each species in February and March. Chicks of both species received meals of the same size, of which half consisted of liquid. The three major components of the solid diet (krill, squid and fish) were similar for both albatrosses. By weight, fish represented about 35 % of the diet of both species; squid predominated (50 %) in the diet of Grey-headed Albatross, and krill (40 %) in the diet of Black-browed Albatross. Lampreys were confined to the Grey-headed Albatross and, although squid of similar sizes were taken by both species, Black-browed Albatrosses took a much greater diversity of squid. Each major prey type was associated with a characteristic amount of liquid in the complete samples and only in the case of krill and lamprey was this lipid-rich.
As these two albatrosses are of similar size, breed over the same period and feed meals of equivalent weight to their chicks at similar intervals, the difference in the composition of the diet is possibly the most significant mechanism of ecological segregation (in the breeding season).
Evidence of the effect of krill shortage in drastically reducing Black-browed Albatross breeding success is presented to support this. The two species have largely non-overlapping winter oceanic ranges which are probably also correlated with the distribution of preferred prey.     

A comparison of Antarctic petrel (Thalassoica antarctica) diets with net samples of Antarctic krill (Euphausia superba) taken from the Prydz Bay region

Samples of the stomach contents of Antarctic petrels (Thalassoica antarctica) were obtained on board ship in the Prydz Bay region of Antarctica from birds which had spontaneously regurgitated. The weight of food and the species composition of the stomach contents were measured. Antarctic krill,Euphausia superba, the sole prey item taken, were compared to krill obtained by nets in the same region as part of a large-scale krill survey. Krill from petrel stomach samples were larger in mean size than those sampled by nets. This may be attributed to selection of the larger sized krill by the petrels, it may be caused by the nets sampling different populations of krill or it may be due to net avoidance by the larger krill.     

Food, feeding ecology and ecological segregation of seabirds at South Georgia

At the sub-Antarctic island of South Georgia 25 of the 29 breeding species are seabirds. Fifteen of these have recently been studied in some detail. By examining the timing of their breeding seasons and their diet and feeding ecology (especially feeding techniques and potential foraging ranges), the nature of their ecological isolating mechanisms, and in particular the way in which they partition the resources of the marine environment, are reviewed.
Although breeding season adaptations occur (winter breeding in Wandering Albatross and King Penguin; out of phase breeding in two species-pairs of small petrels) these are less important than dillerences in food and feeding ecology. There is a fundamental distinction between the niche of pursuit-diving species (mainly penguins) and the remainder which are basically surface-feeders. The two abundant krill-eating penguins show clear differences in feeding zones. Three albatrosses and a petrel feed mainly on squid and there are differences in both the species and size of the prey of each. The remaining seabirds chiefly take krill (although the giant petrels are extensive scavengers and some smaller petrels specialize on copepods) and utilize different feeding methods and areas to do so.
Various adaptations related to inshore and offshore feeding zones are discussed. Although most species possess a combination of ecological isolating mechanisms additional evidence for the particular importance of dietary differences is presented.     

Diet and feeding ecology of saddle-back (Saguinus fuscicollis) and moustached (S. mystax) tamarins in an Amazonian terra firme forest

The feeding ecology of two small-bodied primate species—saddle-back ( Saguinus fuscicollis auilapiresi ) and moustached tamarins ( S. mystax pileatus )—occurring in stable, mixed-species groups was studied in a terra firme forest site in the upper Urucu River, Amazonas, Brazil. Ecological data are based primarily on one mixed-species group of 5–8 saddle-back and 8–10 moustached tamarins. The overall vegetative and animal-prey components of each tamarin species' diet, their selection of food species, and the seasonal variation in their use of plant resources are described, and compared to those of callitrichids elsewhere. The extremely diverse diet of tamarins included at least 136 tree, 33 vine and liana, 12 epiphyte and nine shrub species, as well as a wide range of prey items. They fed primarily on ripe fruit pulp of most of these species for most of the year, but shifted to floral nectar and plant exudates of a few key plant species during the dry season. Taxonomic overlap in plant diet was nearly complete between the two tamarin species, but they diverged considerably in their prey capture techniques. Saddle-backs used the low forest understorey, and manipulatively searched for sedentary prey concealed within discrete, usually rigid, microhabitats, whereas moustached tamarins used the midstorey where they visually searched for mobile prey well exposed on foliage. These and other feeding and foraging patterns are discussed in the light of other callitrichid species studied to date.     

Leopard seals (Hydrurga leptonyx) use suction and filter feeding when hunting small prey underwater

Leopard seals (Hydrurga leptonyx) are unusual among apex predators in that they feed at both the top and near the bottom of marine food webs; they capture and consume marine amniotes (seals and penguins) as well as krill. This is thought to be achieved with their unusual dentition: rostral caniniform teeth function to grip large prey and tricuspate postcanines function to sieve krill. The use of canine teeth is known, yet until now, the function of the postcanines has never been documented. Here, we present the first direct observations of filter feeding in leopard seals. Suction was used to draw small prey into the mouth followed by expulsion of ingested seawater through the sieve formed by postcanine teeth. Individuals show abrasive wear on canines and incisors, but not postcanines. This suggests that postcanines are not systematically used for piercing prey during macrophagous feeding, confirming that the postcanines primarily serve a sieving function. Rather than being less efficient at feeding as a result of its polarized diet, the leopard seal is well adapted towards two disparate feeding modes.     

Harp seal foraging behaviour during summer around Svalbard in the northern Barents Sea: diet composition and the selection of prey

The harp seal Pagophilus groenlandicus is a major high trophic level predator in the Barents Sea, and to better understand their function in the Barents Sea ecosystem, we need to understand their foraging behaviour during their most intensive feeding period. We analysed the diet composition and prey preference of 184 harp seals and 94 faeces samples, sampled in the northern Barents Sea (around Svalbard) during the period May–August in 1996, 1997, and 2004–2006. Concurrent with the sampling of seals, prey availability was assessed in one area in 1996 and 1997 and in two areas in 2006 using standard acoustic methods. Our study showed that harp seal diet composition varied significantly both in time (year) and space, and that their diets appeared to be size dependent. Both subadult (<150 cm) and adult seals were associated with pelagic crustaceans (particularly krill), whereas primarily adult seals were associated with fish (capelin, gadoids and flatfish). Krill was the most important prey group (63 %) followed by polar cod (16 %) and other fish species (10 %). The prey preference of harp seals varied in time and space; polar cod was often preferred by the seals whereas krill was commonly consumed in lower proportion than observed in the survey area. Gadoids and capelin had either been exploited in the same or less proportion as observed in the survey sea. This study emphasises the ecological significance of krill as prime food for harp seals during their intensive feeding period in summer.     

Distribution and diet of 0-group cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) and haddock (<Emphasis Type="Italic">Melanogrammus aeglefinus</Emphasis>) in the Barents Sea in relation to food availability and temperature

Distribution of 0-group cod (Gadus morhua) and haddock (Melanogrammus aeglefinus) in August–September 2005 and 2006 was mainly restricted to the Atlantic waters of the western and central areas of the Barents Sea. The main distribution of 0-group fish overlapped largely with areas of high biomass (>7 gm⁻² dry weight) of zooplankton. The copepod Calanus finmarchicus and krill Thysanoessa inermis, which are dominant zooplankton species in both Atlantic and boreal waters of the Barents Sea, were the main prey of 0-group cod and haddock. The main distribution, feeding areas and prey of 0-group cod and haddock overlapped, implying that competition for food may occur between the two species. However, though their diet coincided to a certain degree, haddock seems to prefer smaller and less mobile prey, such as Limacina and appendicularians. As 0-group fish increased in size, there seems to be a shift in diet, from small copepods and towards larger prey such as krill and fish. Overall, a largely pelagic feeding behaviour of 0-group cod and haddock was evident from this study.     

Success and its limits among structural models of complex food webs

1. Following the development of the relatively successful niche model, several other simple structural food web models have been proposed. These models predict the detailed structure of complex food webs given only two input parameters, the numbers of species and the number of feeding links among them. 2. The models claim different degrees of success but have not been compared consistently with each other or with the empirical data. We compared the performance of five structural models rigorously against 10 empirical food webs from a variety of aquatic and terrestrial habitats containing 25-92 species and 68-997 links. 3. All models include near-hierarchical ordering of species' consumption and have identical distributions of the number of prey of each consumer species, but differ in the extent to which species' diets are required to be contiguous and the rules used to assign feeding links. 4. The models perform similarly on a range of food-web properties, including the fraction of top, intermediate and basal species, the standard deviations of generality and connectivity and the fraction of herbivores and omnivores. 5. For other properties, including the standard deviation of vulnerability, the fraction of cannibals and species in loops, mean trophic level, path length, clustering coefficient, maximum similarity and diet discontinuity, there are significant differences in the performance of the different models. 6. While the empirical data do not support the niche model's assumption of diet contiguity, models which relax this assumption all have worse overall performance than the niche model. All the models under-estimate severely the fraction of species that are herbivores and exhibit other important failures that need to be addressed in future research.     

The food and feeding ecology of the giant petrels Macronectes halli and M. giganteus at South Georgia

Giant petrels are the chief scavenging seabirds in the Southern Ocean. Quantitatively analysed regurgitations from chicks of both Macronectes halli and M. giganteus at Bird Island, South Georgia throughout the chick rearing period consisted predominantly of adult Macaroni penguins Eudyptes chrysolophus. Significant quantities of burrowing petrels and krill were taken but the amount varied in different years. Only male M. halli fed regularly at fur seal carcases. Although 10 species were found in regurgitations, squid were unimportant in the diet. Fish are possibly more important during the winter.
Intersexual differences in diet were of greater significance than interspecific ones with males taking more carrion and females more crustaceans. Chicks were fed during both day and night and in both species males received significantly more food than females during the fledging period. M. giganteus chicks received more than M. halli at all ages.
Despite their extensive reliance on carrion both species take a variety of prey and this may be especially important during the winter when penguin and seal carrion is scarce.     

Influence of the diet on the bioaccumulation of heavy metals in zooplankton-eating petrels at Kerguelen archipelago,Southern Indian Ocean

Concentrations of cadmium, mercury, copper and zinc were measured in muscle, kidney and liver tissues and in the main prey of five species of zooplankton-eating petrels: blue petrel (Halobaena caerulea), thin-billed (Pachyptila belcheri) and Antarctic (P. desolata) prions, and South Georgian (Pelecanoides georgicus) and common (Pelecanoides urinatrix) diving petrels. Since some of these species are closely related species with respect to body size, timing of moult and life span, their diet has been examined to evaluate its influence on heavy-metal bioaccumulation. Inter-specific differences were significant for Hg concentrations in the liver and for Zn concentrations in both liver and kidney tissues. Blue petrels exhibited the highest Hg concentrations in the liver (3.9±2.02 µg.g^–1 wet weight). No significant differences were found in Cd concentrations between species. Exposure to heavy metals through the most important prey species in the diet during the breeding period was evaluated. The most evident result was the influence of fish prey on Hg levels. Although crustacean species exhibit different cadmium concentrations, the diet composition does not appear to be discriminant for Cd bioaccumulation within the small petrel community at Kerguelen.     

Effects of habitat fragmentation on provisioning rates, diet and breeding success in two species of tit (great tit and blue tit)

The aim of this study was to examine the effects of forest fragmentation on the ability of parent birds to provide their young with an adequate food supply. To examine whether prey population densities of the great tit (Parus major L.) and the blue tit (P. caeruleus L.) vary between study areas in different forest size classes we compared provisioning rates and chick diet and related these parameters to breeding success. We filmed 217 nests over two breeding seasons and collected data on frass fall as a general estimate of caterpillar availability. Nests which were attended by none or one parent only during filming (n = 46) were excluded from the analyses. In both years and for both species feeding rates were highest in the smallest fragments and lowest in the large forest. There was also a suggestion that differences in feeding rates between areas vary between years. We found no consistent tendency for prey size to change with forest size, although both species brought slightly smaller prey items to the nest in the smallest forest fragments and feeding rates correlated negatively with prey size. Caterpillars were the main item fed to nestlings, in both species. We found no evidence to suggest that either frass fall or the proportion of caterpillars in the diet varied with forest size. There was also no correlation between mean frass fall and the total number of caterpillars brought to the nests, in either species. Breeding success, as measured by clutch size, brood size, fledging weight and fledging success, did not differ between the small fragments and the large forest, in either species. There was also no relationship between provisioning rate (as concerns volume of prey fed to nestlings and the quality of chick diet) and breeding success parameters. In conclusion, this study does not suggest suboptimal foraging or breeding conditions in small fragments compared to a nearby large forest, for either species. Received: 23 June 1997 / Accepted: 29 December 1997     

Limitation of similarity and feeding habits in three syntopic species of newts (Triturus, Amphibia)

Differences in body size between both species and sex were observed to be close to Hutchinsonian ratios in an assemblage of three species of newts (Triturus italicus, T alpestris and T carnifex) in southern Italy According to feeding, differences in body size were not related with a strict food niche segregation The diet of larger newt included indeed all the prey eaten by smaller newts but showed new prey types Differences between species were more marked than between sexes of the same species, despite a strong sexual dimorphism in body size Larger newts foraged in a wider range of microhabitats and exhibited a greater diversity of foraging tactics Triturus italicus fed essentially on plankton and chironomids in shallow water at the banks of the pond Triturus alpestris foraged also in deeper water, particularly on lsopods in vegetation Prey fallen on the water surface constituted the main part of T carnifex diet According to recent data on newt evolution, speciation consisted in the reduction of body size Our data show that this process has been related with a specialization of the diet and of habitat use     

The food and chick feeding of blackbellied stormpetrel (Fregetta tropica) at King George Island, South Shetlands

Food and chick feeding of the blackbellied stormpetrel were investigated at King George Island, South Shetland Islands, from December 1995 till March 1996. Seventy-five regurgitates and 174 samples from stomach flushing were analysed. Provisioning rate and meal size of the chicks were estimated by daily and three-hourly chick weighing. The mean weight of regurgitates of adult birds was 1.63 g. Fish (53.3% of occurrence) and crustaceans (50%) formed the main part of the diet. The frequencies of crustaceans in the diet were 8.3% amphipods, 21.7% euphausiids, 1.7% mysids and 18.3% unidentified crustaceans. Cephalopods represented only a small part in the diet (1.7%). Diet composition did not change throughout the season. The feeding frequencies of the chicks were 0.98 feedings per day on average. The mean meal size ranged from 9.2 to 11.0 g. Feeding and meal size data were calculated by different methods and compared with those of other petrel species. Received: 26 August 1997 / Accepted: 16 December 1997     

DNA as a Dietary Biomarker in Antarctic Krill, Euphausia superba

The diet of Antarctic krill (Euphausia superba) has been studied using a variety of techniques, but current methods still suffer from problems that are difficult to solve. This study examined an alternative approach utilizing DNA as a prey biomarker. Methods were developed for the preservation, extraction, and identification of prey DNA from krill collected in the field. Group-specific polymerase chain reaction (PCR) was used to amplify diatom prey (Phylum: Bacillariophyta) and the results from DNA clone libraries were compared with microscopic diet analysis. DNA analysis was superior to microscopy for prey detection. However, differences in prey relative abundance estimates between the two techniques suggested some bias in the DNA-based estimates. Quantification showed that large amounts of prey DNA had been successfully preserved and extracted. Overall the results suggest that the application of DNA-based diet analysis to krill warrants further investigation, particularly for prey that are difficult to study using other methods.     

Chick provisioning and nest attendance of male and female Wilson’s storm petrels <Emphasis Type="Italic">Oceanites oceanicus</Emphasis>

Seabirds show a range of patterns of sexual size dimorphism and sex-specific parental investment, but the underlying causes remain poorly understood. The aim of the present study was to test two longstanding hypotheses of parental investment in a sexually monomorphic species, Wilson’s storm petrel Oceanites oceanicus, namely that males attend chicks more frequently and females deliver larger meals (Beck and Brown in Br Antarct Surv Sci Rep 69:1–54, 1972). We recorded in eight seasons, both during incubation and chick rearing, which adult was caught first in a nest and found no difference in the probability of catching a male or a female first in any year. Additionally, in five seasons we employed a miniature video camera to record nest attendance during chick rearing and found no significant difference except for 2006, a year with very low krill availability, where females visited the nest less often than males. We then combined video observations with periodic weighing of chicks to estimate mean daily feeding mass (g/day) of males and females and found no difference in the amount of food delivered per day between the sexes. However, in years with low krill availability, males and females tended to use different strategies to achieve the same feeding rates, with females undertaking longer foraging trips and delivering heavier meals. Thus, our results do not support the hypothesis of a general sex-specific parental investment in Wilson’s storm petrels, but a tendency for a context-dependent sex-specific investment in the years of food shortage.     

Common Guillemot Uria aalge parents adjust provisioning rates to compensate for low food quality

The quantity and quality of food available within the foraging area set important constraints for chick‐rearing birds, but responses to low quality are not well understood. This study explored the potential for parent birds to adjust quantity (feeding rate) and quality (energy content) in chick provisioning, by studying Common Guillemots Uria aalge on Stora Karlsö, Baltic Sea, predominantly utilizing Sprat Sprattus sprattus, during conditions of high food quantity but reduced food quality. Quality is central to reproductive success in this single‐prey loader. From the chick's perspective, provisioning rates should be increased to compensate for low food quality and to fulfil its growing needs with increasing age. However, the high energy cost of flying in Guillemots makes it important for parent birds to minimize commutes to feeding areas. Provisioning parameters were recorded during three dawn‐to‐dusk watches each breeding season from 2005 to 2013, when clupeids, presumably Sprat, constituted 98% of chick diet. Generalized additive mixed models showed that both feeding rate and size of clupeids (a proxy for energy content) varied between years and changed non‐linearly with chick age, but that there was no change within breeding seasons. Chick age and year explained 36% of the variation in feeding rate but only 2% of the variation in the size of clupeids in chick diets. We conclude that parent birds tried to adjust both feeding rate and prey size, but were less successful with the latter. A strong negative correlation was found between annual feeding rates and size of clupeids, evaluated as the differences relative to the baseline year, and adjusted for the effects of chick age. Although the differences between years were small, the relationship indicates a compensation mechanism that does not seem to impact adult survival, and by which increased feeding rates can partly counteract reduced chick energy intake when food quality is low.     

Diet and foraging opportunism of the Guiana dolphin (Sotalia guianensis) in the Abrolhos Bank,Brazil

Diet analysis allows exploring how coastal dolphins interact with the environment and their role in the marine food webs. We studied the diet and feeding ecology of the Guiana dolphin, Sotalia guianensis, through analysis of stomach content from 42 animals stranded on the eastern coast of Brazil. A total of 1,336 semidigested prey items (fish, otoliths, cephalopod beaks, and crustaceans) were identified. Teleost fish comprised the most frequent food item (92% of the total), followed by cephalopods, and crustaceans. Prey belonged to 34 taxa and richness in individual stomachs varied from 1 to 15 prey taxa. Prey were generally small, but showed a significant trend to increase in size with dolphin length. The main prey of Guiana dolphins were demersal, estuarine, and sound-making fish such as catfish and sciaenids. No sex-related differences in diet were found. Diet composition varied seasonally and occurrence of prey items was coherent with breeding or high abundance periods of some fish species and squids. Our study shows the importance of demersal prey from estuarine and soft-sediment habitats to Guiana dolphin in the Abrolhos Bank and reveals that feeding habits are generalist and opportunistic, with diet reflecting the seasonal abundance and availability of prey.     

Winter diet and foraging range of gentoo penguins (<Emphasis Type="Italic">Pygoscelis papua</Emphasis>) from Kidney Cove,Falkland Islands

The winter diet and foraging range of gentoo penguins, Pygoscelis papua, were studied at Kidney Cove, Falkland Islands. The mean wet mass of the 56 stomach-content samples collected from May to October was 32.7ᇃ.4 g. The diet consisted generally of cephalopods, crustaceans and fish, as well as two other molluscs. Among the six cephalopod species identified, the commercially fished Patagonian squid, Loligo gahi, had the highest abundance and was also the main prey by reconstituted mass (53% of the total reconstituted mass). Lobster krill, Munida gregaria, one of five species of crustaceans, was the most abundant prey species by frequency of occurrence and by number (68% and 60%, respectively). Rock cod, Patagonotothen ramsayi, accounted for the majority of the fish diet with 34% of the total reconstituted mass. Most prey species identified in the winter diet were also abundant diet components during the breeding season. However, the known biology of the prey species and their rate of digestion indicated that, in winter, adults may forage further offshore than during the breeding season. This assumption was supported by the results obtained from two birds satellite-tracked during the study period. Both birds remained mainly in inshore waters and returned frequently ashore, but one penguin foraged up to 276 km from the coast. The differences in the foraging behaviour of the two birds were reflected in significant differences with regard to time spent underwater, distance travelled per day and calculated travelling speed. Furthermore, the progress of cohorts of L. gahi over the winter is consistent with results from life-cycle studies in this region and suggests that birds have been foraging in the feeding grounds of L. gahi.     

Feeding Ecology of Northeast Atlantic Mackerel,Norwegian Spring-Spawning Herring and Blue Whiting in the Norwegian Sea

The Norwegian spring-spawning (NSS) herring (Clupea harengus), blue whiting (Micromesistius poutassou) and Northeast Atlantic (NEA) mackerel (Scomber scombrus) are extremely abundant pelagic planktivores that feed in the Norwegian Sea (NS) during spring and summer. This study investigated the feeding ecology and diet composition of these commercially important fish stocks on the basis of biological data, including an extensive set of stomach samples in combination with hydrographical data, zooplankton samples and acoustic abundance data from 12 stock monitoring surveys carried out in 2005–2010. Mackerel were absent during the spring, but had generally high feeding overlap with herring in the summer, with a diet mainly based on calanoid copepods, especially Calanus finmarchicus, as well as a similar diet width. Stomach fullness in herring diminished from spring to summer and feeding incidence was lower than that of mackerel in summer. However, stomach fullness did not differ between the two species, indicating that herring maintain an equally efficient pattern of feeding as mackerel in summer, but on a diet that is less dominated by copepods and is more reliant on larger prey. Blue whiting tended to have a low dietary overlap with mackerel and herring, with larger prey such as euphausiids and amphipods dominating, and stomach fullness and feeding incidence increasing with length. For all the species, feeding incidence increased with decreasing temperature, and for mackerel so did stomach fullness, indicating that feeding activity is highest in areas associated with colder water masses. Significant annual effects on diet composition and feeding-related variables suggested that the three species are able to adapt to different food and environmental conditions. These annual effects are likely to have an important impact on the predation pressure on different plankton groups and the carrying capacity of individual systems, and emphasise the importance of regular monitoring of pelagic fish diets.