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.     

Satellite tracking of Adélie penguins

Summary Female Adélie penguins (Pygoscelis adeliae) that take too long on their first post-laying foraging trip are a major cause of breeding failure, but in the ice-filled waters of Antarctica, determining where they go and why they are away so long has proved difficult. Here we describe the first successful attempt to track penguins at sea using satellite telemetry. Four females foraged in different locations, dispelling the notion of a common feeding ground. They moved up to 272 km from the rookery and covered from 551 to 1,121 km on their trips, swimming at minimum average speeds around 1.2 m/s. The birds were most likely to be in the water between 0630 and 1430 when light intensity, important for a visual predator, was greatest. Carrying the transmitters reduced rates of fat deposition (weight gain), increasing the duration of foraging trips of females, and suggested that they may forage until their fat depots reach a minimum threshold level. This has two implications: (i) durations of these postlaying foraging trips could potentially be used as an indicator of krill abundance (Euphausia sp), the almost exclusive food of Adélie penguins during this period, and (ii) any reduction in krill stocks caused by harvesting could increase foraging trip durations with a concomitant increase mi breeding failures.     

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.     

Krill-feeding behaviour of gentoo penguins as shown by animal-borne camera loggers

Animal-borne camera loggers were used to examine the patterns of prey encounter and feeding behaviour of gentoo penguins at King George Island, Antarctica. The still images from the camera loggers showed that the penguins encountered the swarms of krill for 25.5% (range: 8–38%) of their dives (>5 m) on average, during their foraging trips (mean duration of 5.4 h, n = 7 trips). They encountered krill swarms during the dives to 10–70 m depth, in pelagic as well as benthic habitats. In the benthic habitat, the penguins swam just above the sea floor and headed downward over a krill swarm, probably using the sea floor to assist them to feed on mobile swarms. The shallow coastal waters would be the important foraging habitat of gentoo penguins breeding in King George Island.     

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.     

The effects of prey demography on humpback whale (Megaptera novaeangliae) abundance around Anvers Island, Antarctica

Baleen whales and Adelie penguins in the near-shore waters around the Antarctic Peninsula forage principally on Antarctic krill. Given the spatial overlap in the distribution of these krill predators (particularly humpback whales) and their dependence on krill, the goals of this paper are to determine if the inter-annual community structure and relative abundance of baleen whales around Anvers Island is related to krill demography and abundance, and if the potential exists for inter-specific interactions between Adelie penguins and baleen. We use whale sightings and prey data from both net tows and Adelie penguin stomach samples to correlate the abundance of humpback whales with krill demography and abundance from 1993 to 2001. We find significant relationships between whale abundance and the size–frequency distribution of krill targeted by Adelie penguins, as well as the foraging success of Adelie penguins. These findings suggest both krill predators share common prey preferences in the upper portions of the water column around Anvers Island. These findings highlight the need for better knowledge of baleen whale foraging ecology and inter-specific interactions with penguins, as sea ice and krill populations around the Antarctic Peninsula are affected by rapid changes in climate.     

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.     

Can macaroni penguins keep up with climate- and fishing-induced changes in krill?

Macaroni penguins have evolved to cope with the highly variable conditions of the Southern Ocean. However, changes in prey supply and patchiness potentially associated with changes in climate and krill fishing activity may be occurring too rapidly for the penguins to adapt. We use a stochastic dynamic programming model to examine how changes in both the mean and patchiness of krill supply may affect the foraging decisions, and therefore breeding success, of female macaroni penguins at South Georgia. We predict that rapid changes in the mean supply of prey will have more of an effect on the condition of the female and chick than changes in prey patchiness, and that changes in foraging behavior compensate for changes in prey up to a threshold point, beyond which breeding success is likely impacted. In particular, we predict that the location of the threshold is affected by whether or not the penguins are adapted to the prey environment in which they are foraging, with the female and chick receiving on average 20% less of their daily energetic requirement if the female is not foraging optimally.     

Euphausia superba dominates in the diet of Adélie penguins feeding under fast sea-ice in the shelf areas of Enderby Land in summer

Adélie penguins Pygoscelis adeliae in Enderby Land, Antarctica feed mainly on Euphausia superba during the chick rearing season in shelf areas where fast sea-ice remains: indicating that E. superba is abundant under the fast sea-ice in these areas. The shelf areas in Enderby Land, therefore, are unique since the previous studies of Adélie penguin diet in Ross Sea, Adélie Land and Prydz Bay show that E. crystallorophias is the most abundant krill species in shelf areas in general.     

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.     

Outbreaks of avian cholera in Hope Bay, Antarctica

During austral summers 1999-2000 and 2000-01, two outbreaks of avian cholera occurred in the Hope Bay area (63 degrees 24'S, 56 degrees 59'W), located on the tip of the Antarctic Peninsula. Eighty-six dead birds were found: five kelp gulls (Larus dominicanus), 36 skuas (Stercorarius sp.), and 45 Adelie penguins (Pygoscelis adeliae). The carcasses were studied using clinical, pathological, and microbiological criteria. Water samples from ponds where birds were settled and samples from 90 healthy birds also were analyzed during the second outbreak. Pasteurella multocida isolates were identified by biochemical tests, capsular type, somatic serotype, and susceptibility to nine antibiotics. Molecular subtyping was performed by ApaI and SmaI pulsed-field gel electrophoresis (PFGE) and enterobacterial repetitive intergenic consensus (ERIC-PCR). In February 2000, mortality in skuas was 16% and 2% in kelp gulls. In the 2000-01 breeding season, mortality in south polar skuas was 47%, 24% in brown skuas, 1.4% in kelp gulls, and 0.01% in Adelie penguins. All birds had lesions of avian cholera. In kelp gulls the presentation was chronic, whereas skuas and penguins suffered subacute and acute disease, respectively. Fifty-five isolates recovered from dead birds and one from water were identified as P. multocida gallicida, type A:1. The strains presented a unique molecular pattern by PFGE and ERIC-PCR. A possible hypothesis to explain the origin of the outbreaks was that nonbreeder kelp gulls carried P. multocida gallicida to Hope Bay, and avian cholera was transmitted through water to skuas and penguins. This study reports avian cholera in new bird species, their potential role in the transmission of the disease, and the different responses of these species to the disease.     

Diving pattern and performance in the macaroni penguin Eudptes chrysolophus

The pattern and characteristics of diving in two female macaroni penguins Eudyptes chrysolophus was studied, during the brooding period, using continuous-recording time-depth recorders, for a total of I8 days (15 consecutive days) during which the depth, duration and timing of 4876 dives were recorded. Diving in the first 11 days was exclusively diurnal, averaging 244 dives on trips lasting 12 hours. Near the end of the brooding period trips were longer and included diving at night. About half of all trips (except those involving continuous night-time diving) was spent in diving and dive rate averaged 14–25 dives per hour (42 per hour at night). The duration of day time dives varied between trips, and averaged 1.4–1.7 min, with a subsequent surface interval of 0.5–0.9 min. Dive duration was significantly directly related to depth, the latter accounting for 53% of the variation. The average depths of daytime dives were 20–35 m (maximum depth 11 5 m). Dives at night were shorter (average duration 0.9 min) and much shallower (maximum 11 m); depth accounted for only 6% of the variation in duration. Estimates of potential prey capture rates (3–5 krill per dive; one krill every 17–20 s) are made. Daily weight changes in chicks were directly related to number of dives, but not to foraging trip duration nor time spent diving. Of the other species at the same site which live by diving to catch krill, gentoo penguins forage exclusively diurnally, making longer. deeper dives; Antarctic fur seals, which dive to similar depths as macaroni penguins, do so mainly at night.     

Warming effects in the western Antarctic Peninsula ecosystem: the role of population dynamic models for explaining and predicting penguin trends

The western Antarctica Peninsula and Scotia Sea ecosystems appear to be driven by complex links between climatic variables, primary productivity, krill and Avian predators. There are several studies reporting statistical relationships between climate, krill and Penguin population size. The Adélie (Pygoscelis adeliae), Chinstrap (P. antarctica) and Gentoo (P. papua) penguins appear to be influenced by interannual variability in sea-ice extent and krill biomass. In this paper we developed simple conceptual models to decipher the role of climate and krill fluctuations on the population dynamics of these three Pygoscelis penguin species inhabiting the Antarctic Peninsula region. Our results suggest that the relevant processes underlying the population dynamics of these penguin species at King George Island (South Shetland Islands) are intra-specific competition and the combined effects of krill abundance and sea-ice cover. Our results using population theoretical models appear to support that climate change, specifically regional warming on the western Antarctic Peninsula, represents a major driver. At our study site, penguins showed species-specific responses to climate change. While Chinstrap penguins were only influenced by krill abundance, the contrasting population trends of Adélie and Gentoo penguins appear to be better explained by the “sea-ice hypothesis”. We think that proper population dynamic modeling and theory are essential for deciphering and proposing the ecological mechanisms underlying dynamics of these penguin populations.     

Super-aggregations of krill and humpback whales in Wilhelmina Bay, Antarctic Peninsula

Ecological relationships of krill and whales have not been explored in the Western Antarctic Peninsula (WAP), and have only rarely been studied elsewhere in the Southern Ocean. In the austral autumn we observed an extremely high density (5.1 whales per km(2)) of humpback whales (Megaptera novaeangliae) feeding on a super-aggregation of Antarctic krill (Euphausia superba) in Wilhelmina Bay. The krill biomass was approximately 2 million tons, distributed over an area of 100 km(2) at densities of up to 2000 individuals m(-3); reports of such 'super-aggregations' of krill have been absent in the scientific literature for >20 years. Retentive circulation patterns in the Bay entrained phytoplankton and meso-zooplankton that were grazed by the krill. Tagged whales rested during daylight hours and fed intensively throughout the night as krill migrated toward the surface. We infer that the previously unstudied WAP embayments are important foraging areas for whales during autumn and, furthermore, that meso-scale variation in the distribution of whales and their prey are important features of this system. Recent decreases in the abundance of Antarctic krill around the WAP have been linked to reductions in sea ice, mediated by rapid climate change in this area. At the same time, baleen whale populations in the Southern Ocean, which feed primarily on krill, are recovering from past exploitation. Consideration of these features and the effects of climate change on krill dynamics are critical to managing both krill harvests and the recovery of baleen whales in the Southern Ocean.     

An Interpretation of the Growth of the Adelie Penguin Rookery at Cape Royds, 1955-1990

The population dynamics of the Cape Royds rookery were modelled by computer, in order to determine the probable causes of the dramatic increase since 1980 in the numbers of Adelie penguins, Pygoscelis adeliae, breeding in the Ross Sea region, Antarctica. Variations in the extent of sea-ice around the rookery during incubation and chick rearing cannot feasibly explain the population increase and another factor or event must be introduced, which increases chick production per breeding pair and decreases adult mortality. The timing of the event is critical and rules out the cessation of human impacts or the depletion of competing baleen whales as causal factors. The event is seen as most probably the result of a recent warming of the Ross Sea climate.     

Health evaluation of penguins (Sphenisciformes) following mortality in the Falklands (South Atlantic)

In the Falklands, heavy mortality of rock-hopper penguins Eudyptes chrysocome occurred during the 1985-86 breeding season. Starvation was diagnosed as the primary cause of death, possibly caused by a shortage of euphausiid crustaceans (krill) due to unusual meterological conditions. 'Puffinosis' may possibly have been a contributory factor; otherwise no conclusive evidence of infectious disease or toxicosis was found and also no evidence of radioactive contamination. In the 1986-87 breeding season no unusual mortality occurred, but 99 apparently healthy penguins were examined, i.e., rockhoppers Eudyptes chrysocome syn E. crestatus, gentoos Pygoscelis papua and Magellanics Spheniscus magellanicus. Full necropsies were carried out on 54. Tissue examinations were made for cadmium, copper, iron, manganese, mercury, lead and zinc. High tissue cadmium concentrations found in healthy birds in 1987 were similar to those in penguins which died in 1986, and therefore not considered to be of pathological significance. Although there has been no repetition of the unusually hot 1985-86 breeding season in the Falklands, penguins and other seabirds have had fluctuating breeding successes since then. The precise cause, including the roles of meteorological conditions and overexploitation of some forms of prey species, is unclear.     

Stable isotopes reveal Holocene changes in the diet of Adélie penguins in Northern Victoria Land (Ross Sea, Antarctica)

Adélie penguin (Pygoscelis adeliae) modern and fossil eggshells and guano samples collected from ornithogenic soils in Terra Nova Bay (Victoria Land, Ross Sea) were processed for carbon and nitrogen isotopic ratios with the aim of detecting past penguin dietary changes. A detailed and greatly expanded Adélie penguin dietary record dated back to 7,200 years BP has been reconstructed for the investigated area. Our data indicate a significant dietary shift between fish and krill, with a gradual decrease from past to present time in the proportion of fish compared to krill in Adélie penguin diet. From 7,200 to 2,000 years BP, δ¹³C and δ¹⁵N values indicate fish as the most eaten prey. The dietary contribution of lower-trophic prey in penguin diet started becoming evident not earlier than 2,000 years BP, when the δ¹³C values reveal a change in the penguin feeding behavior. Modern eggshell and guano samples reveal a major dietary contribution of krill but not a krill-dominated diet, since δ¹³C values remain much too high if krill prevail in the diet. According to the Holocene environmental background attested for Victoria Land, Adélie penguin dietary shifts between fish and krill seem to reflect penguin paleoecological responses to different paleoenvironmental settings with different conditions of sea-ice extension and persistence. Furthermore, Adélie penguin diet appears to be particularly affected by environmental changes in a very specific period within the breeding season, namely the egg-laying period when penguin dietary and feeding habit shifts are clearly documented by the δ¹³C of eggshell carbonate.     

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     

Divergent responses of Pygoscelis penguins reveal a common environmental driver

The responses of predators to environmental variability in the Antarctic Peninsula region have exhibited divergent patterns owing to variation in the geographic settings of colonies and predator life-history strategies. Five breeding colonies of Pygoscelis penguins from King George Island and Livingston Island, South Shetland Islands, Antarctica, were examined to (1) compare the responses of sympatric congeners to recent changes in their Antarctic ecosystem and (2) assess underlying causes for such responses. We used linear regression and correlation analyses to compare indices of abundance, recruitment, and summer breeding performance of the Adélie (P. adeliae), gentoo (P. papua), and chinstrap penguins (P. antarctica). Breeding colonies of Adélie and chinstrap penguins have declined by roughly 50% since the mid-1970s, and recruitment indices of Adélie penguins have declined by roughly 80%, but no such patterns are evident for gentoo penguins. Fledging success, however, has remained stable at all breeding colonies. The different trends in abundance and recruitment indices for each species, despite generally similar indices of summer performance, suggest that winter conditions contribute to the divergent responses among the penguins. In particular, strong correlations between indices of penguin and krill recruitment suggest that penguins in the South Shetland Islands may live under an increasingly krill-limited system that has disproportionate effects on the survival of juvenile birds.     

Occurrence of cyclopoid copepods and faecal material in the Halley Bay region,Antarctica, during January–February 1991

Faecal material and cyclopoid copepods were collected during the expedition ANT IX/3, in the Halley Bay area (Weddell Sea, Antarctica), between January and February 1991. Faecal material comprised pellets produced by krill, copepods, ostracods and appendicularians. Cyclopoid copepods were represented by two genera, Oithona and Oncaea. In the Halley Bay area, higher concentrations of krill faecal material (420.9 mm³ m^–2) and chl.-a (39.3 mg m^–2) were found within the upper 200 m of the water column of the polynya than in ice-covered open-ocean areas (58.2 mm³m^–2 and 25.5 mg m^–2, respectively). At an ice-drift station, high concentrations of krill faecal strings under fest-ice were found. In addition, similarities between diatom assemblages in the pack-ice algae and krill faecal strings contents suggest an active utilization of ice-algae by krill populations. Sedimented material collected at 50 m depth by a sediment trap was dominated by krill faecal strings. Contents of small oval pellets (of probable cyclopoid copepod origin) resemble those of krill faecal pellets suggesting that coprophagy was involved. This suggestion is supported by: (1) The small quantity of food particles (other than krill faecal matter) available in the water column (< 0.3 g chl.-al^–1). (2) The negative in situ correlation between krill faecal strings and cyclopoid copepods. (3) The structure of cyclopoid copepod buccal appendages, which are more adapted for raptorial feeding.