Breeding success and chick provisioning in Wilson’s storm-petrels <Emphasis Type="Italic">Oceanites oceanicus</Emphasis> over seven years: frequent failures due to food shortage and entombment

We examined the natural variation in parameters of the breeding success of Wilsons storm-petrels on King George Island, South Shetland Islands, during seven breeding seasons, and determined the relative influence of krill abundance and climatic conditions. Feeding frequencies, feeding rates and meal sizes varied between years, and feeding rates were correlated with data of the Antarctic krill Euphausia superba abundance in the Elephant Island region. The overall breeding success of breeding pairs was low, ranging from 0 to 34%. Snow storms and wind conditions depressed breeding success even in years of good food availability, and snow storms were the most important determinant of the breeding success over the seven seasons. Snow storms during the breeding season occurred during periods of prevailing easterly winds. In addition, the distribution of wind directions was found to strongly influence the krill availability to storm-petrels.     

Resource allocation in Wilson's storm-petrels<Emphasis Type="Italic"> Oceanites oceanicus</Emphasis> determined by measurement of glucocorticoid excretion

When resources are limited, life-history theory predicts that long-lived animals should allocate available resources to body maintenance rather than to reproduction in order to maximise their lifetime reproductive success. In the present study, we estimated physiological stress in a small procellariiform seabird, the Wilson's storm-petrel Oceanites oceanicus, as a means of understanding how limited resources are partitioned between provisioning parents and their chicks. We analysed adrenocortical activity of Wilson's storm-petrels during the breeding season by measuring glucocorticoid (GC) excretion, using an enzyme immunoassay measuring tetrahydrocorticosterone concentrations in extracts of faeces and urine of chicks and adults. Faecal GC measures were negatively correlated with chick body condition, suggesting that measures of tetrahydrocorticosterone in faeces and urine can be used to assess adrenal activity characteristic for physiological stress in Wilson's storm-petrels. In the breeding season of 1999, the colony was subject to low food availability, and the faecal and urine GC levels of chicks were elevated during these months of chronic starvation. In contrast, adults did not show elevated GC levels. The data thus suggest that Wilson's storm-petrels respond to unfavourable conditions by maintaining their own body condition and reducing provisioning of food to their chicks. Communicated by R.F. Oliviera     

The diet of the South Georgia shag Phalacrocorax georgianus at South Orkney Islands in five consecutive years

The diet of the South Georgia shag Phalacrocorax georgianus at Laurie Island, South Orkney Islands, was studied by the analysis of 420 pellets (regurgitated casts) collected from the 1995 to 1999 breeding seasons. Demersal-benthic fish were by far the main prey, followed by molluscs (mainly octopods and bivalves) and polychaetes. Harpagifer antarcticus (in 1995, 1998, 1999) and Lepidonotothen nudifrons (1996, 1997) were the most frequent prey, followed by Gobionotothen gibberifrons or Trematomus newnesi. Harpagifer antarcticus, followed by L. nudifrons or T. newnesi, was the most numerous prey in all of the seasons and predominated in mass in 1995 and 1996, followed by Notothenia coriiceps and L. nudifrons. In 1997 and 1999, G. gibberifrons and N. coriiceps were the species that most contributed to the diet whereas N. coriiceps and H. antarcticus contributed most in 1998. Our results differ from those reported for the South Orkney Islands in previous studies. These differences could be due to the use of different diet-analysis methods and to shags-related and/or fisheries-related changes in the food availability around the colonies. These alternative explanations are analysed and discussed.     

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.     

Comparative study of the effects of the late pack-ice break-off on chinstrap and Adélie penguins' diet and reproductive success at Laurie Island,South Orkney Islands,Antarctica

The effects of ice cover on diet and breeding chronology of Adélie (Pygoscelis adeliae) and chinstrap penguins (P. antarctica) were studied in the colonies located at Point Martin, South Orkney Islands. Reproductive success and diet parameters were measured as described in the CCAMLR Environmental Monitoring Program protocols. It was found that late pack-ice break-off in 1998 resulted in lower reproductive success than in 1999 for chinstrap penguins, while Adélie penguin parameters were rather insensitive. This species has shown to be able to cope with this environmental factor, as suggested by its circumpolar and more southerly range. Significant differences in stomach contents' weights and in the proportion of whole krill were found between ice-covered and open-sea periods within and between years. Foraging trips of both species became longer, resulting in underfeeding of chicks and nest desertion by chinstrap-penguin parents. Simultaneous analysis of reproductive parameters of both species and their diets have proven to be useful in discriminating between variation in reproductive success parameters due to prey accessibility and prey abundance.     

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.     

Chick provisioning and breeding success of Adélie penguins at Béchervaise Island over eight successive seasons

Data on foraging trip durations and meal sizes were collected by means of an automated monitoring system from breeding Adélie penguins (Pygoscelis adeliae) at Béchervaise Island over eight breeding seasons, commencing in 1991-92. Significant interannual differences in duration of foraging trips, meal sizes and diet composition were observed. These parameters were correlated with breeding success and fledging masses, and provide useful indicators of food availability during the breeding season. Chicks were particularly vulnerable to a decreased availability of food during the first 3 weeks after hatching.     

Nocturnal foraging by great skuas Stercorarius skua: implications for conservation of storm-petrel populations

At St Kilda, Outer Hebrides, a large colony of great skuas Stercorarius skua feed extensively on one of the largest colonies of Leach’s storm-petrels Oceanodroma leucorhoa in Europe, but little is known about the dynamics of this predator–prey system. Recently published population estimates of storm-petrels make it possible to estimate the impact of skua predation for the first time. Although skuas in the southern hemisphere catch petrels attending breeding colonies at night, it is not known whether congeners in the northern hemisphere also forage during the hours of darkness. We found (using radio-transmitters) that skuas regularly forage at night and (using light intensifying equipment) observed them catching storm-petrels at night. However, skuas also foraged during daylight hours, and it is unknown whether they might also catch storm-petrels at sea. Data on diet composition reveals that the proportion of storm-petrels in skua diet declined between 1996 and 1997, but remained constant thereafter. Although a large proportion of the storm-petrel prey is likely to consist of non-breeders, numbers consumed suggest that breeders and an unknown quantity of transients may also been eaten. The numbers of storm-petrels eaten are not sustainable and may result in substantial long-term population declines. Under current conditions, maintenance of large populations of both Leach’s storm-petrels and great skuas at St Kilda appears to be mutually exclusive.     

Diet of Antarctic fur seals Arctocephalus gazella at the Danco Coast,Antarctic Peninsula

The diet of non-breeding male Antarctic fur seals, Arctocephalus gazella, was investigated at the Danco Coast, Antarctic Peninsula, by the analysis of 31 and 149 scats collected from January to March 1998 and 2000, respectively. Overall, fish and krill, followed by penguins and squids, were the most frequent prey and constituted the bulk of the diet. The importance of the remaining taxa represented in the samples (octopods, gastropods, bivalves, isopods, polychaetes and poriferans) was negligible. Among fish, channichthyids constituted the bulk of the diet, with Chionodraco rastrospinosus and Chaenodraco wilsoni, followed by the nototheniid, Pleuragramma antarcticum, being the main prey. The myctophid, Electrona antarctica, was the most frequent and numerous fish prey. The results are discussed and compared with those reported for the South Shetland Islands, the closest area for which similar information is available.     

Summer diet of king penguins (Aptenodytes patagonicus) at the Falkland Islands, southern Atlantic Ocean

The diet of king penguins (Aptenodytes patagonicus) brooding chicks was investigated during February 2001 at the Falkland Islands, where a small but increasing population is located at the limit of the breeding range of this species. Fish was the most important food source by number (98.0%) and reconstituted mass (97.8%), squids accounting for the remainder. Myctophid fishes represented the main part of the diet (97.7% by number and 96.6% by reconstituted mass), Protomyctophum choriodon being by far the main prey item (84.2% and 88.1%, respectively). Four other myctophids and one squid species each contributed to more than 1% of the diet by number: Krefftichthys anderssoni (4.8%), Electrona carlsbergi (4.6%), Gymnoscopelus nicholsi (2.2%) and Protomyctophum tenisoni (1.8%), together with small juveniles of Gonatus antarcticus (1.8%). Twelve squid species were identified from accumulated lower beaks, including the ommastrephid Martialia hyadesi (48.3% by number), the onychoteuthids Moroteuthis ingens (15.6%), Kondakovia longimana (10.5%) and Moroteuthis knipovitchi (7.3%), and Gonatus antarcticus (9.2%). The stable-carbon and stable-nitrogen isotopic composition of chick food and adult blood differed in a way that suggests that, during the same trip, adult birds fed for themselves in distant foraging grounds, and fed for their chicks on their way back to the colony. The study emphasizes that king penguins are specialist myctophid eaters throughout their breeding range in summer, and highlights the importance of Protomyctophum choriodon as a link between zooplankton and top predators in the pelagic ecosystem of the southwestern Atlantic Ocean.     

Inter-annual variability in Chinstrap penguin diet at South Shetland and South Orkneys Islands

Inter-annual variability in the diet of Chinstrap penguins (Pygoscelis antarctica) at Laurie Island (South Orkney Islands) and 25 de Mayo/King George and Nelson Islands (South Shetland) was examined based on stomach contents of adults during the 2002/2003–2006/2007 and 2002/2003–2004/2005 breeding seasons, respectively. Krill (Euphausia superba) dominated the diet as frequency of occurrence (in 100% of samples), number (>99%), and percentage contribution in weight (>94.8%). Other prey items were minor and varied between years. The weight of stomach contents was significantly different. The percentage in weight of whole krill was used to compare the feeding conditions across seasons. It differed significantly at the three sites studied. Distribution of krill size varied among years and localities, showing different krill availability for penguins.     

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.     

Regional and temporal variation in diet and provisioning rates suggest weather limits prey availability for an endangered raptor

Understanding variation in food requirements of wild animals is of central importance in population ecology and conservation, as it helps to identify where and when food may be limiting. Studies on diet variation or prey provisioning rates may give useful insights when direct information on prey availability is lacking. We assess spatial and temporal variation in the diet of an endangered predator, the Black Harrier Circus maurus. This raptor is endemic to southern Africa and specializes on small mammals but also feeds on birds and reptiles as alternative prey. Using data on 1679 prey identified in 953 pellets collected in inland and coastal regions from 2006 to 2015, we show that diet composition changed little throughout the breeding season in the coastal region, whereas there was a marked seasonal decline in the occurrence of small mammal prey in the inland region, with a concomitant increase in alternative prey. The proportion of small mammals in the diet declined with increasing maximum temperature, the latter being highest at the inland region late in the breeding season. Using camera recordings at nests in 2014, we further analysed daily patterns of prey provisioning to nestlings. A marked reduction in small mammal provisioning rates occurred during the middle of the day in the hotter inland region but not in the cooler coastal region. Reduced availability of the primary prey, small mammals, in hotter conditions, through a reduction in activity or overall abundance, could explain these patterns. Finally, we show a positive relationship between winter rainfall and interannual differences in the proportion of small mammals in the diet of Black Harriers breeding in the coastal region, suggesting relationships between diet and prey abundance that are mediated through rainfall. We discuss the need to consider spatial variation in food availability in conservation strategies.     

The food and feeding ecology of Blue petrel (Halobaena caerulea) and Dove prion (Pachyptila desolata)

The diet of two small burrow dwelling petrels, Dove prion, Pachyptila desolata and Blue petrel Halobaena caerulea , was studied at the subantarctic island of South Georgia by quantitative analysis of 246 food samples regurgitated by adults about to feed chicks. Crustaceans, and particularly krill Euphausia superba , predominate in both species' diet but fish is considerably more important to the Blue petrel. Other dietary differences between the species chiefly involve the relative proportions of krill, copepods (nearly exclusive to Dove prion) and seven amphipod species taken.
There is no overlap in the species' chick feeding periods but food differences are likely to reflect prey selection rather than resource availability; differences in prey size are ascribed to use of different feeding techniques particularly relating to bill structure and function. Data on frequency of chick feeding, proportion of oil and presence of pumice in food samples are used to indicate that the Blue petrel feeds further away from the breeding colony than the Dove prion. The significance of the various adaptive differences is discussed.     

Dietary shifts,niche relationships and reproductive output of coexisting Kestrels and Long-eared Owls

Summary Food samples of breeding Kestrels (Falco tinnunculus) and Long-eared Owls (Asio otus) were collected in the peak and low phase of their preferred prey (Microtus voles) in western Finland. Diets of pairs that bred as neighbours (1 km) with interspecifics were compared with those of non-neighbours. In both species, neighbouring pairs fed less on Microtus voles and more on alternative prey than did non-neighbours. Competition theory predicts that diet overlap should be lower during prey shortage and that diet similarity should be especially reduced in neighbouring pairs. Observations were consistent with expectations: diet similarity was lower in the low vole years and neighbouring pairs showed less diet overlap that non-neighbours. Differences in habitat composition and prey availability at the sample sites should not confuse the results. In addition to the high diet similarity, hunting habitats and nest sites of the species overlapped almost completely; they only showed clear temporal segregation in hunting. Probably because of food competition, the neighbouring pairs of both species produced significantly fewer young than the non-neighbours. These results contrast with the view that the diet composition and dietary shift of rodent-feeding predatory birds can be interpreted in terms of simple opportunistic foraging. In the breeding season, interspecific competition for food seems to be an important factor that affects the niches of these species, especially in northern areas, where the seasonal low phase of voles in spring and the number of alternative prey are lower than in more southern areas.     

The diet of the Weddell seal <Emphasis Type="Italic">Leptonychotes weddellii</Emphasis> at Laurie Island,South Orkney Islands

The diet of the Weddell seal Leptonychotes weddellii at the South Orkney Islands was investigated by the analysis of 44 and 26 faecal samples collected from the beaches of Laurie Island from May 1999 to January 2000 and from October to December 2001, respectively. The diet was diverse and both pelagic and benthic-demersal organisms were represented in the samples. Fish were the most frequent prey of the samples in both seasons, and fish and krill were the most numerous prey items. Octopods predominated by mass in 1999, whereas krill constituted the bulk of the diet in 2001, a fact that was not observed previously. Among fish, Gobionotothen gibberifrons was largely the species that contributed mostly to the diet, whereas pelagic fish could not be identified in the samples. The results are compared with information from other study areas and discussed in relation to the consumption of krill.     

Biological characteristics of euphausiids preyed upon by Adélie penguins in relation to sea-ice conditions in Lützow-Holm Bay,Antarctica

We examined the biological characteristics of euphausiids found in the stomachs of Adélie penguins in relation to sea-ice conditions in Lützow-Holm Bay over three seasons. Euphausiids, especially Euphausia superba, proved to be a staple food for Adélie penguins irrespective of the ice condition. Body length and maturity-stage compositions of euphausiids were different among seasons, probably reflecting sea-ice condition in summer. The mean body length decreased and maturity regressed during each season in E. superba, which was partly attributable to the selective feeding on large, mature female krill by Adélie penguins. The 1995/1996 year class of E. superba, which was spawned when the sea ice was most developed, was strong and conspicuous in the 1996/1997 and 1997/1998 seasons. This vigor indicates that sea ice provided females with good spawning conditions and larvae with good growth and survival rates.     

Diet and prey consumption of Antarctic petrels Thalassoica antarctica at Svarthamaren, Dronning Maud Land, and at sea outside the colony

The diet of the Antarctic petrel Thalassoica antarctica was studied during two seasons at Svarthamaren, an inland colony in Dronning Maud Land, Antarctica, and in the pack ice off the coast of Svarthamaren. The most important food (wet mass) at Svarthamaren was crustaceans (67%), fish (29%) and squid (5%); however, individuals collected in the pack ice took mostly fish (87%). The prey composition and lengths of prey are comparable to what has been documented in other studies on this species. Estimates of food consumption by birds breeding at Svarthamaren (ca. 250,000 pairs) suggest that approximately 6500 tonnes of crustaceans, 2800 tonnes of fish and 435 tonnes of squid are consumed during the breeding season. The annual consumptions of these birds are estimated to be 34,100 tonnes of crustaceans, 14,700 tonnes of fish, and 2300 tonnes of squid. Satellite telemetry data indicate that Antarctic petrels from Svarthamaren may fly more than 3000 km during one foraging trip, and thus may cover a huge ocean area to obtain their prey. Received: 1 September 1997 / Accepted: 3 February 1998     

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.     

Provisioning and growth in chicks of Wilson's storm-petrels (Oceanites oceanicus) on King George Island, South Shetland Islands

We present data on chick growth and chick feeding in Wilson's storm-petrel (Oceanites oceanicus) in a colony on King George Island, South Shetland Islands. Chicks were repeatedly weighed and the weight differences over 24 h were corrected for metabolic loss in order to obtain an estimation of meal sizes. Chicks were fed on 93% of the nights (n=688 nights). The average meal size for a single feeding was 8.5 g. Chicks received on average 1.2 feedings per night. These results are compared with data for this species from other locations. There was a trend for increased meal sizes from northern to southern populations, parallel to an increase in the adult mass, indicating that Wilson's storm-petrels carry optimal meal sizes according to their body size and may take advantage of increased food abundance by increasing feeding frequencies. We describe chick growth and discuss the influence of egg size, hatching date and feeding frequency on chick growth. The egg size had a positive influence on tarsus growth and body mass of chicks. Later-hatched chicks started wing growth and finished mass growth at a younger age and reached lower peak masses, indicating that late chicks may adapt to the restricted breeding season in their Antarctic breeding grounds by a more rapid development, but will fledge with a lower degree of development and less resources. Accepted: 22 May 2000