Interannual study of Chinstrap penguin’s diet and reproductive success at Laurie Island, South Orkney Islands, Antarctica

Inter-annual variability in the diet of Chinstrap penguins (Pygoscelis antarctica) at Laurie Island, South Orkney Islands was examined based on stomach contents of adults coming ashore during the 1997/1998–2001/2002 breeding seasons. Krill (Euphausia superba) dominated the diet either as frequency of occurrence (in 100% of samples), number (>99%), and percentage contribution in weight (>99.5%). Other prey items were minor but did vary between years. The contribution in weight of fish, amphipods and cephalopods ranged between 0.01–0.5, <0.01–0.14 and 0–0.03, respectively. Although minor components varied slightly along the years, the weight of stomach contents was significantly different. It was found that for a given weight the maximum observed percentage of undigested krill resulted negatively correlated with the stomach contents total weight. The proportion of whole krill observed in individual penguins, expressed as a fraction of the maximum corresponding to its weight was used to characterize the prevailing conditions in each season.     

Identifying patterns in the diet of mackerel icefish (<Emphasis Type="Italic">Champsocephalus gunnari)</Emphasis> at South Georgia using bootstrapped confidence intervals of a dietary index

Ontogenetic, inter-annual and regional variations in diet were investigated for mackerel icefish, Champsocephalus gunnari, in three successive summer seasons around South Georgia. Stomach contents from 2239 C. gunnari (130–560 mm total length) were examined. A bootstrapping technique was used to calculate confidence intervals for an index of relative importance of prey categories (% IRI_DC). Diet varied significantly between years and age classes but there was little regional difference in diet. In general, diet was dominated by krill, Euphausia superba and by the amphipod Themisto gaudichaudii. Smaller (younger) fish tended to prey on a higher proportion of T. gaudichaudii and small euphausiids such as Thysanoessa sp. and took smaller quantities of E. superba. In a season of poor krill availability (summer of 2003–2004) the proportion of krill in the diet, stomach fullness and fish condition (indicated by length–weight relationships) were significantly lower than in the other summer seasons. A large reduction (>80%) in the estimated annual (2005) biomass of the C. gunnari stock directly followed the season of poor krill availability. This decline was largely because of mortality of 2+ and 3+ fish, which were more krill dependent than 1+ fish. Younger fish appear to have survived, leading to an increase in the estimated population biomass in 2006.     

Competition between the krill fishery and penguins in the South Shetland Islands

Among the South Shetland Islands, the potential competition for krill (Euphausia superba) between the Japanese fishery and krill-eating breeding penguins was assessed. A low level of competition was apparent due to several factors. Spatial overlap between the main fishing and foraging areas was insignificant. Firstly, the large colonies of the dominant penguin (chinstrap penguins,Pygoscelis antarctica) occurred where sea ice disappears early in spring, and not necessarily where krill are abundant in summer, i.e. the area of krill fishery. Secondly, overlap between trawling depth and foraging dive depth of penguins was marginal, with the latter depth being shallower. Moreover, overlap in the size-frequency of krill caught by trawlers and those captured by penguins was not complete; the penguins took larger krill on average. Finally, the present small fishery is unlikely to impact upon local krill biomass in the region. Krill biomass was estimated to be 250–1500 × 10³ tonnes within the preferred fishing areas during summer. The present catch rate by the fishery (≤13 × 10³ tonnes/half-month period) is smaller by an order of magnitude, and the fishing area does not include the main foraging areas of breeding penguins.     

Penguins, fur seals, and fishing: prey requirements and potential competition in the South Shetland Islands, Antarctica

Antarctic and sub-Antarctic seabirds, marine mammals, and human fisheries concentrate their foraging efforts on a single species, Antarctic krill (Euphausiasuperba). Because these predators may have a significant effect on krill abundance, we estimated the energy and prey requirements of Adelie (Pygoscelisadeliae), chinstrap (Pygoscelisantarctica), and gentoo (Pygoscelispapua) penguins and female Antarctic fur seals (Arctocephalusgazella) breeding on the South Shetland Islands, Antarctica and compared these estimates with catch statistics from the Antarctic krill fishery. Published data on field metabolic rate, population size, diet, prey energy content, and metabolic efficiency were used to estimate prey requirements of these breeding, adult, land-based predators and their dependent offspring. Due to their large population size, chinstrap penguins were the most significant krill predators during the period examined, consuming an estimated 7.8 × 10⁸ kg krill, followed by Adelie penguins (3.1 × 10⁷ kg), gentoo penguins (1.2 × 10⁷ kg), and Antarctic fur seals (3.6 × 10⁶ kg). Total consumption of all land-based predators on the South Shetland Islands was estimated at 8.3 × 10⁸ kg krill. The commercial krill fishery harvest in the South Shetland Island region (1.0 × 10⁸ kg) was approximately 12% of this. Commercial harvest coincides seasonally and spatially with peak penguin and fur seal prey demands, and may affect prey availability to penguins and fur seals. This differs from the conclusions of Ichii et al. who asserted that the potential for competition between South Shetland predators and the commercial krill fishery is low. Received: 26 August 1997 / Accepted: 16 December 1997     

The contrasting diet of Harpagifer antarcticus (Notothenioidei, Harpagiferidae) at two localities of the South Shetland Islands, Antarctica

The diet of Harpagifer antarcticus was studied at two localities of the South Shetland Islands, Antarctica. The analysis of the stomach contents of specimens collected in tide pools at Potter Cove, King George Island, indicated that gammarid amphipods (mainly Gondogeneia sp.) were the main prey of this fish, followed by polychaetes, gastropods and isopods. By contrast, the specimens from Harmony Point, Nelson Island, which were recovered from stomach contents of Antarctic shags Phalacrocorax bransfieldensis feeding at depths of 46–110 m, preyed almost exclusively on the Antarctic krill Euphausia superba. These results are discussed and compared with previous studies. Received: 27 August 1997 / Accepted: 17 November 1997     

Aspects of the foraging behaviour of the Antarctic Tern Sterna vittata gaini at Harmony Point, South Shetland Islands

During January and February of 2002 and 2003, we studied the diet of the Antarctic Tern Sterna vittata gaini at two colonies in Nelson Island, South Shetland Islands, by identifying the prey fed to chicks by breeders. The fish Notothenia coriiceps was the main prey in both seasons, followed by the myctophid Electrona antarctica, Antarctic krill Euphausia superba and gammarid amphipods. The contribution of fish to the diet increased as chicks grew older. Fish and amphipods were brought to chicks during the day, whereas adults brought Antarctic krill at sunrise and sunset. Both the duration of the feeding trips and the number of trips per foraging bout varied according to the type of prey caught. Preliminary information suggests that, among other causes, the foraging strategy is strongly influenced by the predation pressure of skuas on chicks. Results are compared with the only two previous study on the diet of the Antarctic Tern at the South Shetland Islands.     

Cycles of <Emphasis Type="Italic">Euphausia superba</Emphasis> recruitment evident in the diet of Pygoscelid penguins and net trawls in the South Shetland Islands,Antarctica

Size and sex of Antarctic krill taken from chinstrap and gentoo penguin diet were compared to those from scientific net surveys in the South Shetland Islands from 1998 to 2006 in order to evaluate penguin diet as a sampling mechanism and to look at trends in krill populations. Both penguin diet and net samples revealed a 4–5 year cycle in krill recruitment with one or two strong cohorts sustaining the population during each cycle. Penguin diet samples contained adult krill of similar lengths to those caught in nets; however, penguins rarely took juvenile krill. Penguin diet samples contained proportionately more females when the krill population was dominated by large adults at the end of the cycles; net samples showed greater proportions of males in these years. These patterns are comparable to those reported elsewhere in the region and are likely driven by the availability of different sizes and sexes of krill in relation to the colony.     

Diet and reproductive success of Adélie and chinstrap penguins: linking response of predators to prey population dynamics

The diet and reproductive performance of two sympatric penguin species were studied at Signy Island, South Orkney Islands between 1997 and 2001. Each year, Adélie (Pygoscelis adeliae) and chinstrap (P. antarctica) penguins fed almost exclusively (>99% by mass) on Antarctic krill; however, there was considerable inter-annual variation in reproductive output. In 1998, chinstrap penguins were adversely affected by extensive sea-ice in the vicinity of the colony, whereas Adélie penguins were unaffected by this. However, in 2000, both species suffered reduced reproductive output. Detailed analysis of the population-size structure of krill in the diet indicated a lack of recruitment of small krill into the population since 1996. A simple model of krill growth and mortality indicated that the biomass represented by the last recruiting cohort would decline dramatically between 1999 and 2000. Thus, despite the lack of a change in the proportion of krill in the diet, the population demographics of the krill population suggested that the abundance of krill may have fallen below the level required to support normal breeding success of penguins sometime before or during the 2000 breeding season. The role of marine predators as indicator species is greatly enhanced when studies provide data reflecting not only the consequences of changes in the ecosystem but also those data that elucidate the causes of such changes.     

Fish in the diet of the Antarctic shag Phalacrocorax bransfieldensis at Laurie Island, South Orkney Islands

The analysis of 29 stomach contents collected from a colony close to Pirie Peninsula, South Orkney Islands, from 2 January to 18 February 1995, showed that fish were by far the most important prey of the Antarctic shag Phalacrocorax bransfieldensis, followed by octopods, decapods and gammarids. The fish Notothenia coriiceps constituted the bulk of the diet; however, its importance decreased by the end of the study when Lepidonotothen nudifrons was the most important prey. The composition of the stomach contents varied throughout the breeding season according to changes in the chicks' energetic demands; as chicks grew older the adults increased the mass of the loads carried to the nests, preying on larger fish, a phenomenon also observed at the South Shetland Islands. Received: 3 July 1996 / Accepted: 9 March 1997     

The diet of the Antarctic fur seal <Emphasis Type="Italic">Arctocephalus gazella</Emphasis> at the South Orkney Islands in ten consecutive years

To improve the knowledge on the Antarctic fur seal foraging behavior, evaluate whether changes in its diet are associated with changes in prey availability, and evaluate whether fisheries had negative impacts on Antarctic fur seal populations, a total of 1359 scats of non-breeding males were collected in ten consecutive sampling periods between 1994 and 2003 at Laurie Island, South Orkney Islands. Antarctic krill was the most numerous prey throughout the sampling period followed in importance by fish prey. Antarctic krill also predominated by reconstructed mass, except during 1994 and 1998 when penguins were the most important prey, and during 1995 when fish dominated. Among fish, demersal-benthic species (mainly Gobionotothen gibberifrons, Chaenocephalus aceratus and Chionodraco rastrospinosus) dominated the diet except in 1997, 1998, 1999 and 2002 when myctophids (mainly Gymnoscopelus nicholsi and Electrona antarctica) were the most important fish prey. In seasons when fish dominated the diet, the diet was primarily comprised of demersal species. Although our results are in overall agreement with previous studies, the contribution to the diet of the main preys changed throughout the sampling period, probably in parallel with changes in prey abundance. According to historical results on fish consumption, the fisheries carried out in the past have had negative impacts on seal populations from the South Orkney Islands.     

Analysis of the sources of variance in the mean size of krill consumed by Chinstrap and Adélie penguins at South Orkney Islands

The CCAMLR Ecosystem Monitoring Program aims at detecting possible impacts of the krill fishery on predator populations. Population parameters of Chinstrap and Adélie penguins are monitored at Point Martin, Laurie Island, South Orkneys Islands as part of the Program. Diet information is collected to aid in the interpretation of population data. In the present study, the diet of both species is compared in order to detect differences between them as well as to ascertain whether the krill consumed by both species might be pooled in single estimates of prey parameters. A nested ANOVA model was fitted to the dataset obtained in the seasons from 1997/1998 to 2005/2006. The overall variance was decomposed into variance components: within penguins (krill in stomachs), between penguins (means within each date), between dates (date means in each season), and between years (annual means). The within penguins component was similar in both species (18?% difference), while the between penguins was 64?% larger in Adelies. The within and between penguins variance components of both species showed consistent interannual changes. The annual means of both species differed significantly in three seasons, but not consistently. The sources of variance are tentatively attributed to biological characteristics: the within penguins component would reflect searching and feeding at the individual level; the between penguins might be associated with their foraging ranges, an assumption to be tested tracking individuals. Given their differences, the information from both species should be viewed as complementary and not be pooled.     

Diving behaviour of Adélie penguins (Pygoscelis adeliae) at Dumont D'Urville, Antarctica: nocturnal patterns of diving and rapid adaptations to changes in sea-ice condition

The diving behaviour of Adélie penguins (Pygoscelis adeliae) was studied with time-depth recorders at Dumont D'Urville, Antarctica, during the breeding seasons in 1995/1996 and 1996/1997. We studied penguins foraging at all breeding stages, in various sea-ice conditions. For the first time in this species we observed nocturnal patterns of diving as the penguins dived more frequently and spent more time underwater around midnight than around noon. This behaviour may be related to the abundance of neritic krill, Euphausia crystallorophias, in the diet. Dive depth and duration varied extensively over the cycle, and appeared related to sea ice conditions rather than representative of the locality (22 m/78 s and 40 m/102 s with and without sea-ice, respectively). Comparisons with other studies showed that different diving behaviour previously observed in different localities can also occur at the same locality, and in some cases over a single foraging trip of a single penguin when short-term variation of external conditions occurred. Accepted: 27 September 1999     

The food and feeding ecology of Adélie penguins (Pygoscelis adeliae) and Chinstrap penguins (P. antarctica) at Signy Island, South Orkney Islands

Adélie Pygoscelis adeliae and Chinstrap P. antarctica penguins are important consumers of Southern Ocean marine resources. The stomach contents of adult penguins at Signy Island, South Orkney Islands, were analysed quantitatively throughout the chick-rearing period. They consisted almost exclusively of Antarctic krill Euphausia superba , Adélies eating 35–63% by number and 23–28% by weight of juvenile krill and Chinstraps 72–87% by number and 90–95% by weight of mature krill, as well as small amounts offish and amphipods. Interspecific dietary differences may partly be attributable to Adélies starting breeding one month before Chinstraps but, as they persist when both are simultaneously rearing chicks, the two species may also forage in somewhat different areas.
Krill data from net hauls indicate a substantial overlap in the size of krill taken by scientific, and probably also commercial, operations and by Adélie and Chinstrap penguins.
Chicks were fed c. 300 g of food 0–5-0-8 times per day, Chinstrap chicks without a sibling being fed most frequently. Chicks of both species were most often fed in the late afternoon, and from estimates of swimming speed and feeding frequency adults may feed quite extensively at night.     

The diet of Antarctic fur sealsArctocephalus gazella during the breeding season at South Georgia

 The diet of lactating female Antarctic fur seals Arctocephalus gazella at South Georgia was investigated during the pup-rearing period (January/March) of 1991–1994. Antarctic krill Euphausia superba was the main prey item, occurring in 88% of all scats (n=497), whereas fish occurred in 47% and squid in 5% of all scats. There was considerable intra- and inter-annual variation in the characteristics of krill taken by fur seals. The distribution of krill sizes taken suggests that fur seals are not actively selecting particular sizes of krill and, therefore, that the krill in the diet reflects the krill available around South Georgia. The absence of group 3 krill (44–48 mm in length) in the South Georgia area, as indicated by their absence in the diet of seals, is suggested as a possible reason for low availability of krill and the subsequent reproductive failure among krill predators. The frequency of occurrence of fish was much higher than in previous studies; the pattern of fish consumption showed a consistent seasonal pattern in 3 of the 4 years studied. Of the total number of the myctophid Protomyctophum choriodon, the most numerous fish taxon, 98% were taken between early February and the middle of March. Champsocephalus gunnari and Lepidonotothen larseni agg., which both feed on krill, dominated the fish component of the diet outside this period and together constituted 94% of the total estimated biomass of fish consumed. The intra- and inter-annual variability in the diet of Antarctic fur seals emphasise the need for diet studies to be conducted during the entire pup-rearing periods of several years. Received: 15 March 1995/Accepted: 13 May 1995     

Using carapace measurements to determine the sex of Antarctic krill, Euphausia superba

Krill (Euphausia superba) carapace measurements (length and width; mm) collected from plankton tows in the South Shetland Islands (SSI), Antarctica are used to test the generality of a common discriminant function developed to reconstruct krill length frequencies in Antarctic fur seal diets for the area surrounding South Georgia (SG). Total length and sex ratio of krill in the SSI were overestimated by 5.6 and 154%, respectively, when the SG allometric equations were applied to 3 years (2003–2005) of data. These errors arise and increase as a result of krill population dynamics, specifically recruitment that contributes large proportions of immature krill, misclassified as males by the SG discriminant function. We develop sex-specific regression models based on separate discriminant functions that provide significantly better discriminatory power. However, our analysis indicates that reconstructions of krill sex ratio and length composition in the ocean environment are less reliable in years when the ratio of immature to mature krill is high. For the SSI area, five out of 14 years (35.7%) surveyed (1992–2005) had proportions of immature to mature adult krill ≥ 0.50.     

Daily activity and minimum food requirements during winter for gentoo penguins (Pygoscelis papua) in the South Shetland Islands, Antarctica

Estimates of daily activity and consequent demand for food during winter are scarce for many polar seabirds, yet essential for assessing constraints on foraging effort, demand for food, and potential competition with local fisheries. We affixed archival temperature tags to gentoo penguins (Pygoscelis papua) from two colonies in the South Shetland Islands to measure the frequency, timing, and duration of foraging trips and to estimate minimum food requirements during winter. Foraging trip frequencies ranged from 0.85 to 1.0 trips day⁻¹ and were positively correlated with day length. Early winter foraging trips more closely matched day length than late winter foraging trips. The data suggest that individuals maximize foraging time during the early winter period, likely to recover body mass following the breeding season and molt. The more attenuated response of foraging trip durations to increasing day length in late winter may be related to differences in local resource availability or individual behaviors prior to the upcoming breeding season. Minimum food requirements also exhibited a seasonal cycle with a mid-winter minimum. On average, minimum food requirements were estimated at 0.70 ± 0.12 kg day⁻¹. Extrapolated to the regional population of gentoo penguins, winter food requirements by gentoo penguins were equivalent to roughly 33% of annual krill catches by commercial fisheries in the South Shetland Island region over the past decade. Current expansion of the gentoo population and the krill fishery in the southern Scotia Sea warrants continued monitoring of gentoo penguins during winter.     

Diet of two icefish species from the South Shetland Islands and Elephant Island,<Emphasis Type="Italic"> Champsocephalus gunnari</Emphasis> and<Emphasis Type="Italic"> Chaenocephalus aceratus</Emphasis>

The summer diet of two species of icefishes (Channichthyidae) from the South Shetland Islands and Elephant Island, Champsocephalus gunnari and Chaenocephalus aceratus, was investigated from 2001 to 2003. Champsocephalus gunnari fed almost exclusively on krill (Euphausia superba) in all years. The importance of other taxa (Themisto gaudichaudii, mysids, myctophids) in the diet was negligible. The average feeding rate of Champsocephalus gunnari inferred from an exponential gastric evacuation model was between 1.0 and 1.5% body weight per day. Most of the stomachs of Chaenocephalus aceratus were empty. Stomachs with food contained mainly krill, mysids and fish. Among the fish taken, locally abundant species formed the bulk of the diet: Gobionotothen gibberifrons in 2001, Lepidonotothen larseni and Champsocephalus gunnari in 2002 and L. larseni in 2003. An ontogenetic shift in feeding preference of Chaenocephalus aceratus was observed: fish smaller than 30 cm fed on krill and mysids, while larger animals relied primarily on fish.     

Geographical variation in the diet of the Antarctic fur seal<Emphasis Type="Italic"> Arctocephalus gazella</Emphasis>

The diet of non-breeding male Antarctic fur seals Arctocephalus gazella was investigated at different localities of the Antarctic Peninsula (Cierva Point and Hope Bay), South Shetland Islands (Deception Island and Potter Peninsula) and the South Orkney Islands (Laurie Island), by the analysis of 438 scats collected from January to March 2000. The composition of the diet was diverse, with both pelagic and benthic-demersal prey represented in the samples. Antarctic krill Euphausia superba was the most frequent and numerous prey at all the study sites except at Cierva Point, followed by fish, penguins and cephalopods. Antarctic krill also predominated by mass, followed by either fish or penguins. Fish were the second most important prey by mass at the Antarctic Peninsula whereas penguins were the second most important prey by mass at the South Shetland and South Orkney Islands. Among fish, Pleuragramma antarcticum was the most important species in the diet of the Antarctic fur seals at the Antarctic Peninsula whereas Gymnoscopelus nicholsi predominated at the South Shetland and South Orkney Islands. The results are compared with previous studies, and the possibility of implementing monitoring studies on the distribution/abundance of myctophids and P. antarcticum based on the analysis of the diet of the Antarctic fur seal is considered.     

Summer diet of leopard seals (<Emphasis Type="Italic">Hydrurga leptonyx</Emphasis>) in Prydz Bay,Eastern Antarctica

The diet of male and female leopard seals (Hydrurga leptonyx) was investigated in Prydz Bay, Eastern Antarctica. A total of 70 scats, 1 regurgitate and 3 stomach contents were collected, during the austral summer, between November 1999 and March 2002. Eight prey species were identified, including birds, mammals, fish and invertebrates. Adelie penguins (Pygoscelis adeliae) were the main prey item and crabeater seals (Lobodon carcinophagus), benthic and pelagic fish, amphipods and krill were found to supplement the diet. Cephalopods did not occur in the diet. Crabeater seals were still being captured well after weaning, and were found in the diet of both male and female leopard seals.     

Sex differences in Adélie penguin foraging strategies

Consistent sex differences in foraging trip duration, feeding locality and diet of breeding Adélie penguins (Pygoscelis adeliae) were demonstrated at two widely separated locations over several breeding seasons. Differences in foraging behaviour were most pronounced during the guard stage of chick rearing. Female penguins made on average longer foraging trips than males, ranged greater distances more frequently and consumed larger quantities of krill. In contrast, males made shorter journeys to closer foraging grounds during the guard period and fed more extensively on fish throughout chick rearing. Mean guard stage foraging trip durations over four seasons at Béchervaise Island, Eastern Antarctica and over two seasons at Edmonson Point, Ross Sea ranged between 31 and 73 h for females and 25 and 36 h for males. Ninety percent of males tracked from Béchervaise Island by satellite during the first 3 weeks post-hatch foraged within 20 km of the colony, while the majority (60%) of females travelled to the edge of the continental shelf (80–120 km from the colony) to feed during this period. Received: 10 December 1997 / Accepted: 10 April 1998