Pygoscelid penguins in a swim canal

Summary In Antarctica, we investigated the energy consumption of Adélie (Pygoscelis adeliae), Gentoo (P. papua) and Chinstrap (P. antarctica) penguins while resting in the water (8.4 W-kg^–1) and swimming underwater at various speeds, using a 21m long canal filled with sea-water at 4°C in conjunction with respirometry. The birds swam at will and consumed 15.7, 16.1 and 10 W·kg^–1 at the speed where cost of transport was minimal (2.1, 2.3 and 2.5 m·s^–1 in Adélie, Gentoo and Chinstrap penguins, respectively). Thermal conductance in pygoscelid penguins was 3.3 W·°C^–1. m^–2 and energy expenditure (P_i, W·kg^–1) while resting in the water is given by P_j = -0.3 t_a+9.6, where t_a is water temperature in °C. During the breeding season, pygoscelid penguins spend 25–40% of their daily energy expenditure while foraging at sea. The importance of accurate estimates of at-sea activity and energy consumption is discussed.     

The diets of five summer breeding seabirds in Adélie Land,Antarctica

Summary The diets of five breeding seabird species were investigated on Adélie Land in January–February 1982. Stomach contents of Adélie penguins, Pygoscelis adeliae, were sampled by a water off-loading method and of Procellariiformes by spontaneous regurgitation. Diet compositions by mass were: Adélie penguin (79% euphausiid, 18% fish, 3% squid); Cape pigeon, Daption capense, (64% euphausiid, 29% fish, 7% carrion); Antarctic fulmar, Fulmarus glacialoides, (64% euphausiid, 20% carrion, 16% fish); snow petrel, Pagodroma nivea, (95% fish, 2% euphausiid, 1% carrion) and Wilson's stormpetrel, Oceanites oceanicus, (39% fish, 37% euphausiid, 13% carrion, 12% various crustaceans). The present Adélie penguin diet is consistent with those reported in other studies, given our knowledge of geographical variation in food availability. Differences in the diets of fulmarine petrels appear to relate to differences in foraging areas. The snow petrel is a fish-eating bird associated with pack-ice. Cape pigeon and Antarctic fulmar are mainly krill-eaters and we infer segregation along a neritic/oceanic gradient because of the importance of the neritic Euphausia crystallorophias in the former and the oceanic E. superba in the latter.     

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.     

People in Antarctica — how much do Adélie Penguins Pygoscelis adeliae care?

Summary In the course of physiological field studies, we opportunistically examined the effects of humans and aircraft on breeding Adélie Penguins Pygoscelis adeliae. Proximity to both aircraft and humans caused substantial increases in penguin heart rate even when no external stress was manifest. A solitary human at 20 m distance from commuting penguins on a well-used pathway caused the birds to deviate by 70 m. Birds at nests exposed to a single human fled much more readily when the brood consisted of large chicks (critical distance 6.1 m) rather than small chicks (critical distance 1.3 m) or eggs (critical distance 0.3 m). Aircraft caused birds to panic at distances greater than 1,000 m and 3 days exposure to a helicopter inhibited birds that had been foraging from returning to their nests, caused bird numbers in the colonies to decrease by 15% and produced an active nest mortality of 8%. Based on this data, we make recommendations to minimize stress on Adélie Penguin colonies exposed to man.     

Field metabolic rates of instrumented Adélie penguins using double-labelled water

Summary Adélie penguins (Pygoscelis adeliae) carrying dummy instruments were used to determine field metabolic rates using double-labelled water. All penguins injected with double-labelled water showed a marked loss of body mass (-4.5%) during the period of the experiments (20–131 h), irrespective of the time of the breeding season. Total body water averaged 57.3% and water flux estimates of field metabolic rates correlated with double-labelled water estimates of field metabolic rate (r ²=0.68), indicating that Adélie penguins do not ingest significant amounts of sea water. Brooding Adélie penguins had a mean field metabolic rate of 10.1 W·kg^-1 and at sea a field metabolic rate of 13.3 W·kg^-1, both of which compare well with previously published estimates based on time/activity budgets and respirometry. Mean field metabolic rate in penguins with crèching chicks was 14.1 W·kg^-1, and the birds spent 65 h absent from the nest as opposed to previous estimates of 7.1 W·kg^-1 and 21 h. The effects of weather, disturbance and manipulation on the behaviour and field metabolic rate of penguins late in the breeding season are discussed. Adélie penguins (crèching chicks) equipped with externally attached instruments spent more time absent from the nest than noninstrumented controls (76 vs 54 h), but had a lower field metabolic rate.Abbreviations ANOVA analysis of variance - DLW double-labelled water - FMR field metabolic rate - MR metabolic rate - RMR resting metabolic rate - TBW total body water - VSMOW Vienna standard mean ocean water - WF water flux     

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.     

Delayed laying and prolonged fasting in Adélie Penguins Pygoscelis adeliae

Observations of nesting Adélie Penguins (Pygoscelis adeliae) were made at Ardley Island during spring 1990 when snow cover was unusually thick at some subcolony sites. Adélie Penguins at these sites had to delay egg laying until the snow melted. Maximum length of fasting periods comprising pre-breeding and incubation was 50 days. Long fasting seemed to have no detrimental effect on breeding. Furthermore, there was no relationship between penguin arrival mass and duration of fast. Even birds with small mass had sufficient reserves to undergo long fasting periods.In spring 1990, when we started with a monitoring study for CEMP (CCAMLR 1990) at Ardley Island, there were still high quantities of snow at the subcolony sites. Adélie Penguins at Ardley Island inhabit both small rocky outcrops and flat, stony hillocks (storm bars). The latter had a distinctly thicker snow cover at this time so that the pebbles necessary for nest building were unattainable. Consequently, we observed the behaviour of the penguins in this situation, recorded the laying dates and lengths of fasting periods.     

Diet of Adélie penguins Pygoscelis adeliae during the post-hatching period at Esperanza Bay,Antarctica, 1987/88

The diet of the Adélie penguin Pygoscelis adeliae was studied at Esperanza Bay, Antarctic Peninsula, during the post-hatching period by quantitative analysis of adult stomach contents. Euphausiids constituted on average 96% by mass, while fish contributed 4% by mass. Amphipods were present in small amounts. Antarctic krill Euphausia superba was the predominant component throughout the sampling period. In contrast, Euphausia crystallorophias occurred rarely. Differences in the size of krill taken by Adélie penguins appear to reflect either local changes in the availability of certain age classes at various times in the breed ing season, or differences in foraging areas, or are due to year-to-year differences in prey availability and abundance, possibly caused by variations in seasonal ice cover. Pleuragramma antarcticum constituted the bulk of the fish portion, particularly during the guard period.     

Acoustic systems are adapted to breeding ecologies: individual recognition in nesting penguins

In all species of penguins studied to date, the display call, or parental call, has been demonstrated experimentally to facilitate identification between mates and between chicks and parents. We investigated parent-chick recognition in two nesting species, the Adélie penguin, Pygoscelis adeliae, and the gentoo penguin, P. papua. Through playback experiments, we tested the capacity of chicks to recognize the parental call at varying levels of background noise. By using modified calls, we found that chicks of neither species used temporal characteristics of the parental call (variations in frequency or amplitude with time) for individual recognition, but that both species used a simpler parameter, the pitch of the call. This finding contrasts with the more sophisticated use of acoustic cues by chicks of two non-nesting species, identified in earlier work. These differences in auditory processing of parental calls may have evolved because of different ecological constraints, particularly whether recognition of a nest site supports mutual identification of parent and offspring. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Responses of <Emphasis Type="Italic">Pygoscelis adeliae</Emphasis> and <Emphasis Type="Italic">P. papua</Emphasis> populations to environmental changes at Isla 25 de Mayo (King George Island)

As part of a monitoring study of Adélie and Gentoo penguin colonies, birds occupying nests with eggs and chicks in crèches were counted annually from the 1995/1996 to the 2006/2007 seasons at Stranger Point, Isla 25 de Mayo (King George Island), Antarctica. During the study period the Adélie penguin population showed a decrease of 62%. The number of chicks in crèches followed a similar trend, the smallest number occurring in 2002, when it was 63% lower than in 1995/1996. In contrast, the Gentoo breeding population size increased by 68%, while chicks produced increased by 63%. Despite the opposing trends in population size between species, there was a positive relation in their interannual variation, although the extent, and for some years the direction, of the change observed always favoured Gentoo penguins. Breeding success (chicks in crèches/nests with eggs) fluctuated between 0.65 and 1.26 for Adélies and between 0.76 and 1.27 for Gentoo penguins, and did not differ significantly between species. The similar breeding success of these species suggests that the contrasting population trends observed were driven by factors operating over winter. We suggest that current changes in environmental conditions may affect adult birds of both species during the previous winter with different intensity but in a roughly similar way, but that juvenile survival of both species and thus the recruitment of new breeders might be affected differentially, with a much lower survival rate of juvenile Adélie penguins.     

Effects of antihelminthic treatment on cell-mediated immunity in Gentoo penguin chicks

Intestinal parasites suppose a cost to hosts as they compete directly for nutritional resources. Therefore, hosts must defend themselves against intestinal parasites by mounting an immune response. Many penguin species acquire parasites through their diet and transfer these parasites to their chicks when feeding them. High parasite loads in penguin chicks could have effects on their growth and body condition, and ultimately on their survival. Here, we evaluated the effect of parasites on the cell-mediated immune system in Gentoo penguin (Pygoscelis papua) chicks at Stranger Point (25 de Mayo/King George Island, South Shetland Islands). To this end, 12 chicks were experimentally deparasitized with a mixture of anthelminthic drugs (albendazole and praziquantel), whereas 10 others were kept as control. We measured cutaneous cell-mediated immunity in response to immunization with phytohemagglutinin (PHA). We also analyzed the leukocyte profile in both treated and control groups before and after the treatment. After the treatment, deparasitized birds showed larger foot-web swelling in response to PHA injection than control birds. Deparasitized penguins also showed lower eosinophil and monocyte counts than controls, whereas heterophils, lymphocytes, and total white blood cell counts did not differ between groups. Our results suggest that Gentoo penguin chicks parasitized with intestinal parasites suffer a cost in terms of reduced cell-mediated immune responses that could ultimately affect their survival.     

Late Holocene occupation of Gentoo Penguins (Pygoscelis papua) on Byers Peninsula, Livingston Island, Antarctica

We report excavations of an abandoned penguin colony on Byers Peninsula, Livingston Island, Antarctica, in austral summer 2008/2009. Nine abandoned pebble mounds were located on Devils Point, near an active Gentoo Penguin (Pygoscelis papua) colony of about 3,000 nests, at an elevation of 40–45 m above sea level. Three of the nine mounds were excavated to recover organic remains for radiocarbon and ancient DNA analyses; two additional mounds were probed to obtain tissue samples for additional dating. All radiocarbon dates were corrected for the marine-carbon reservoir effect using a ΔR of 700 ± 50 years. Calibrated 2-sigma ranges in calendar years before present (B.P.) on 23 samples of egg membrane and bone provided an overall range of 40–1,150 B.P., with most of the dates falling between 225 and 465 B.P. Ancient DNA analysis confirmed that the tissues recovered from these excavations represent Gentoo Penguin. One radiocarbon date from the active Gentoo Penguin colony indicated an age of 285–480 B.P. for the initiation of this current occupation and corresponding in age with most of the abandoned mounds. Although geologic evidence indicates that Byers Peninsula has been ice free for at least 3,000 years, these results indicate that penguin occupation lagged behind deglaciation by over 2,000 years. Small numbers of Chinstrap Penguins (P. antarctica) also occupy the same breeding colonies as Gentoo Penguins at Byers Peninsula, but their absence in the ancient sediments suggests that they have only recently colonized this area.     

Adélie Penguin Population Diet Monitoring by Analysis of Food DNA in Scats

The Adélie penguin is the most important animal currently used for ecosystem monitoring in the Southern Ocean. The diet of this species is generally studied by visual analysis of stomach contents; or ratios of isotopes of carbon and nitrogen incorporated into the penguin from its food. There are significant limitations to the information that can be gained from these methods. We evaluated population diet assessment by analysis of food DNA in scats as an alternative method for ecosystem monitoring with Adélie penguins as an indicator species. Scats were collected at four locations, three phases of the breeding cycle, and in four different years. A novel molecular diet assay and bioinformatics pipeline based on nuclear small subunit ribosomal RNA gene (SSU rDNA) sequencing was used to identify prey DNA in 389 scats. Analysis of the twelve population sample sets identified spatial and temporal dietary change in Adélie penguin population diet. Prey diversity was found to be greater than previously thought. Krill, fish, copepods and amphipods were the most important food groups, in general agreement with other Adélie penguin dietary studies based on hard part or stable isotope analysis. However, our DNA analysis estimated that a substantial portion of the diet was gelatinous groups such as jellyfish and comb jellies. A range of other prey not previously identified in the diet of this species were also discovered. The diverse prey identified by this DNA-based scat analysis confirms that the generalist feeding of Adélie penguins makes them a useful indicator species for prey community composition in the coastal zone of the Southern Ocean. Scat collection is a simple and non-invasive field sampling method that allows DNA-based estimation of prey community differences at many temporal and spatial scales and provides significant advantages over alternative diet analysis approaches.     

Human-induced behaviour in Adélie penguins Pygoscelis adeliae

Summary Adélie penguins Pygoscelis adeliae appear to be little perturbed by man. We examined the incidence of nest desertion and duration of foraging trip in Adélie penguins when manipulated and fitted with devices of differing sizes. Birds with ca. 1 cm clipped from their tail feathers stayed at sea 50% longer than unmarked controls. The length of foraging trip and incidence of nest desertion increased with increasing device volume. Penguins fitted with devices did not reduce foraging trip length to that of unpackaged birds for at least 19 days. The susceptibility of Adélie penguins to disturbance should be carefully considered when activity patterns are being studied.     

Energetics of under-water swimming in Adélie penguins (Pygoscelis adeliae)

Summary The energy consumption of Adélie penguins while at rest in water (8.4 W·kg^-1 at 4°C) or swimming below the surface was determined using a 21 m long canal fitted with respiration chambers at each end. Penguins chose to swim 86% of the time at speeds recorded in nature. Cost of transport was lowest (7.9 J·kg^-1·m^-1) at 1.7–2.3 m·s^-1, corresponding to a power input of 15.8 W·kg^-1, and only 50% as high as previously reported. Assuming a muscle efficiency of 0.25, propulsion efficiency is 0.4 and overall efficiency is 0.1. Calculated food requirements vary between 1060 g krill per adult and foraging trip at the beginning of the breeding season and 2500 g at the period of highest demand, prior to crèching of the chicks.Abbreviations BMR basal metabolic rate - COT cost of transport - DEE daily energy expenditure - DF daily food - M mass - P _i power input - P _o power output - PVC polyvinyl chloride - RMR resting metabolic rate - SE standard error - STPD Standard temperature, pressure and density - VO₂ oxygen consumption - t time     

Contrasting population changes in sympatric penguin species in association with climate warming

Climate warming and associated sea ice reductions in Antarctica have modified habitat conditions for some species. These include the congeneric Adélie, chinstrap and gentoo penguins, which now demonstrate remarkable population responses to regional warming. However, inconsistencies in the direction of population changes between species at different study sites complicate the understanding of causal processes. Here, we show that at the South Orkney Islands where the three species breed sympatrically, the less ice‐adapted gentoo penguins increased significantly in numbers over the last 26 years, whereas chinstrap and Adélie penguins both declined. These trends occurred in parallel with regional long‐term warming and significant reduction in sea ice extent. Periodical warm events, with teleconnections to the tropical Pacific, caused cycles in sea ice leading to reduced prey biomass, and simultaneous interannual population decreases in the three penguin species. With the loss of sea ice, Adélie penguins were less buffered against the environment, their numbers fluctuated greatly and their population response was strong and linear. Chinstrap penguins, considered to be better adapted to ice‐free conditions, were affected by discrete events of locally increased ice cover, but showed less variable, nonlinear responses to sea ice loss. Gentoo penguins were temporarily affected by negative anomalies in regional sea ice, but persistent sea ice reductions were likely to increase their available niche, which is likely to be substantially segregated from that of their more abundant congeners. Thus, the regional consequences of global climate perturbations on the sea ice phenology affect the marine ecosystem, with repercussions for penguin food supply and competition for resources. Ultimately, variability in penguin populations with warming reflects the local balance between penguin adaptation to ice conditions and trophic‐mediated changes cascading from global climate forcing.     

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.     

Population variability in heat shock proteins among three Antarctic penguin species

Heat shock proteins (HSPs) are synthesised under stressful conditions such as exposure to elevated temperatures, contamination, free radicals, UV light or pathophysiological states resulting from parasites and/or pathogens. HSPs function to protect cells by means of modulation of protein folding. In Antarctica, these proteins have been studied in such organisms as protozoa and fishes, without attention to geographical variation. We studied the variation of HSP70 and HSP60 levels in Gentoo, Adelie and Chinstrap penguins among different populations along the Antarctic Peninsula from King George Island (62°15′S) to Avian Island (67°46′S). Our results show that the northern population of Gentoo penguin showed higher levels of HSP70 and HSP60 than the southern population. High temperature, human impact and immunity as a proxy for parasites and diseases in northern locations could explain such variation. Adelie penguin only showed significant geographical variation in HSP70, increasing north to south, a pattern perhaps related to increased UV radiation and decreased temperatures from north to south. Chinstrap penguin shows no population differences in the variation in neither HSP70 nor HSP60, although HSP70 showed marginally significant differences. Sexual differences in the level of these proteins are also discussed.     

Factors affecting fledging weight of Adélie penguin (Pygoscelis adeliae) chicks: a modeling study

An individual-based model is developed to examine mechanisms that potentially underlie the observed constancy in fledging weight (2.8-3.2 kg) of Adélie (Pygoscelis adeliae) penguin chicks, in spite of large variability in the abundance of Antarctic krill (Euphausia superba), the primary food source. The model describes the energetic requirements of the chick, with growth resulting from the difference between assimilated energy and respiration. Parameterizations of these metabolic processes are based upon experimental and field observations. Ingestion of Antarctic krill by the chick is dependent on the frequency of food delivery to the chick by the adults, which is based on measured foraging times. The mass, size, and size frequency distribution of Antarctic krill fed to the chick are specified using empirical data. The energy content of the Antarctic krill provided to the chick is taken to be constant or allowed to vary with size. The simulations show that food availability is most critical in the latter portion of chick development, when growth rates and food demands are high. Low food availability during this time must be compensated by either feeding chicks with larger krill of higher caloric value or by increased assimilation efficiency. Periods when small krill with lower caloric value dominate require more frequent feeding of the chicks in order to attain their observed fledging weight. Thus, although the total food energy given to the chick is the main factor determining chick growth, the distribution of food availability relative to chick size (i.e., different net growth rates) and food quality are also factors influencing the fledging weight of penguin chicks. The simulations provide insight into the compensating effects of food delivery, food quality, and metabolic processes that allow Adélie penguin chicks to reach their observed fledging weight in spite of considerable environmental variability in food supply.