The movements of gentoo penguins Pygoscelis papua from Ardley Island, Antarctica

The movements of gentoo penguins (Pygoscelis papua) in Antarctica were studied by equipping a total of 37 birds captured at Ardley Island, South Shetlands between December 1991 and May 1996 with position-determining devices. Information on area usage was derived from 20 of these devices and covered the incubation period (N = 3 birds), the chick-rearing period (N = 14 birds) and the over-wintering period (N = 3 birds). During incubation birds only ventured further than 50 km from the colony 20% of the time and no individual ranged further than 200 km from the colony. In contrast, no individuals attending chicks ranged further than 16 km from the colony. During winter the maximum distance ranged from the colony was 268 km. Mean distances between the birds and the colony were 80, 81 and 127 km. Individual birds tended to associate with one spot, making short (10 day) forays away before returning to nodal areas. The ranging capacity of gentoo penguins appears considerably less than that of sympatric congeners and may reflect the ability of gentoo penguins to dive deeper and thus exploit prey not accessible to congeners. Received: 1 October 1997 / Accepted: 3 February 1998     

Estimated feeding rates and energy requirements of gentoo penguins,Pygoscelis papua,at Macquarie Island

Summary Water and sodium turnovers of 6–7 week old gentoo penguin chicks and breeding adults were measured using isotopically labelled water and sodium. Influx rates for chicks averaged 188 ml·kg^-1·day^-1 and 13.9 mmol·kg^-1·day^-1 for water and sodium, respectively. Chicks consumed an estimated 228 g·kg^-1·day^-1 fresh food or 886 kJ kg^-1 day. These values correspond to 761 g·day^-1 or 2945 kJ·day^-1 for a gentoo chick mid-way through the growth period. Flux rates for adults attending chicks ranged from 199 to 428 ml·kg^-1·day^-1 for water and from 15 to 36 mmol·kg^-1·ay^-1 for sodium.     

Breeding energetics and food requirements of gentoo penguins (Pygoscelis papua) at Heard and Macquarie Islands

The food and energy requirements of breeding gentoo penguins ( Pygoscelis papua ) were studied at Heard and Macquaric Islands by means of isotope turnover techniques.
The food consumption rates of chicks were measured at various stages of growth, providing estimates of the total food provided by adults to rear a chick to fledging.
The energy expenditures of attending adults were also measured at different stages of chick-rearing, allowing the total energy costs associated with breeding to be established for a pair of adults and at the population level on both islands. We estimate the total annual energy budget of a 6·2 kg breeding gentoo penguin to be 1517 MJ which is equivalent to the consumption of 292 kg prey.     

Health evaluation of wild gentoo penguins (Pygoscelis papua) in the Antarctic Peninsula

Historically wildlife conservation was based on habitat protection and exploitation control. Only recently have diseases been considered an important issue. However, pathogens are usually described during or after disease outbreaks, but to determine which pathogens may be emerging, surveys of wildlife health are critical in a given time. This study deals with the health status of gentoo penguins Pygoscelis papua in two localities at the Antarctica Peninsula and one at Ardley Island off the South Shetland Islands. Cloacal swaps, fresh fecal samples, ectoparasites, and blood smears were collected. We examined and dissected 14 penguin corpses found dead. Fecal samples were positive for Campylobacter, Escherichia coli and in the carcasses four endoparasitic species were found: Diphyllobothrium sp. and Parorchites zederi, Corynosoma shackletoni and Stegophorus adeliae. The tick Ixodes uriae occurred in five of the examined penguins, and the louse Austrogoniodes gressitti on six birds. From the colony grounds, we collected 1,184 I. uriae. We recorded antibiotic-resistant bacteria, such as E. coli, in ecosystems where gentoo penguins breed. Cloacal samples (300) were negative for Chlamydia, as well as for Salmonella, Campylobacter, E. coli, Newcastle and Influenza viruses.     

Foraging range of king penguins Aptenodytes patagonicm during summer at Marion Island

Foraging ranges of king penguins Aptenodytes patagonicus were estimated by combining information on the feeding rates to chicks and brood shift lengths of adults (assessed by daily weighings of large chicks and daily checks of marked birds brooding small chicks) with measurements of travelling speeds and activity budgets at sea (assessed using remote recording devices). Adults brooding small chicks were relieved on average every 13 days and large chicks were fed every four days. Adults with large chicks spent 36% of their time, between attachment of the device and recapture, travelling at an average speed of 8.7 km.h^-1. This gives an estimated mean maximum foraging range of about 300 km. Adults attending small chicks spent 19% of their time away swimming, giving an estimated mean maximum foraging range of 225 km. Extreme foraging ranges for all birds were 75 and 902 km for penguins returning between two and 24 days at sea, respectively. Total distance travelled was highly correlated with time away from the colony.     

Breeding performance of gentoo penguins (Pygoscelis papua) at a colony exposed to high levels of human disturbance

Port Lockroy, situated on the Antarctic Peninsula, is one of the most visited tourist sites in Antarctica. The effects of visitor disturbance on the breeding performance of gentoo penguins (Pygoscelis papua) at Goudier Island, Port Lockroy was investigated during the austral summer of 1996/1997 by comparing pairs in treatment areas (visited by 35–55 tourists every 1–2 days) and control colonies (not visited by tourists). There were no differences between the two groups in the proportion of birds that laid, in hatching success or the proportion of single-chick broods. Pairs in treatment colonies laid a higher proportion of single-egg clutches, but this was related to colony location and unusually high snow accumulation. Most treatment colonies were situated on low-lying ground or in the lee of buildings, and probably had more late layers at the initial census. Only 11% of tourist visits had occurred by laying, making human disturbance an unlikely explanation for the higher proportion of single-egg clutches. Nests monitored in a disturbed colony and a control colony showed no differences in chick mass or survival up to 20 days of age. The overall breeding success, based on counts of creched birds, was similar to other southern populations of gentoo penguins, after correcting for mortality between creching and fledging. Historical data from Goudier Island indicate that the colony established itself in 1985 and has rapidly increased in size since then. The neighbouring colony at Alice Creek, which has been regularly visited by tourists for at least a decade, has also shown a population increase, although this expansion has been at a slower rate. We conclude that disturbance from tourist visits is unlikely to have been a major determinant of gentoo population change at Port Lockroy. Accepted: 8 November 1998     

Behaviour and breeding success of gentoo penguins Pygoscelis papua in areas of low and high human activity

A key factor influencing wildlife responses to human activity is the degree to which animals have been previously exposed to human stimuli. On subantarctic Macquarie Island, gentoo penguins Pygoscelis papua breed in areas of high and low human activity (on and off-station, respectively). We investigated the behaviour and breeding success of gentoo penguins on and off-station, by a) comparing the behavioural responses of guarding gentoos before, during and after exposure to standardised pedestrian approaches, and b) employing an observational study to determine how human activity may have contributed to within-season breeding success in light of other environmental and site variables. Behavioural responses to pedestrian visitation by gentoos off-station were significantly stronger than those of birds breeding on-station. However, no relationship was found between pedestrian activity and breeding success off-station. Breeding success was, however, positively related to colony size, and negatively related to the activity of other penguins, the number of nearby southern elephant seal Mirounga leonina harems and the location of colonies within short grassland. On-station, breeding success was amongst the highest recorded for that season. Habituation, predator exclusion and the relevance of these findings for management are discussed.     

Foraging ecology of Gentoo Penguins Pygoscelis papua at Macquarie Island during the period of chick care

The diet, diving behaviour, swimming velocity and foraging range of Gentoo Penguins Pygoscelis papua were studied at Macquarie Island during the breeding season in the 1993–1994 austral summer. Gentoo Penguins are considered to be inshore feeders, and at Macquarie Island the diet and estimated foraging ranges supported this. The diet consisted of 91.6% fish and 8.3% squid, by mass. The dominant prey taxa were the fish Gymnoscopelus sp. and Paranotothenia magellanica. A mixture of pelagic and benthic prey was consumed, with a greater proportion of benthic species occurring later in the season. The penguins exhibited a strong diurnal pattern in their diving behaviour. Deep diving (≥30 m) began near sunrise (03.00 h) and finished close to sunset (21.00 h). Diving at night was less common and very shallow (<10 m). Early in the breeding season, dive profiles indicated that birds were probably following vertically migrating pelagic prey through the water column and were foraging in waters over 100 m deep. Later in the season, more uniform, shallower depths were used, suggesting an increase in benthic foraging activity. These changes in dive pattern and depth were consistent with the habitat preferences of prey species found in the diet. Gentoo Penguins swam at 1.04 m per s and had a maximum potential foraging range of about 26 km for single-day trips. They tended to forage within 14 km of the colony, with a mean range of 5.4 km. This range encompassed the deep ocean habitat to the west and east of the island and a shallow area to the north.     

Influence of sampling protocol on diet determination of gentoo penguins Pygoscelis papua and Antarctic fur seals Arctocephalus gazella

The influence of two sampling protocols on diet determination of two marine predators, the gentoo penguin (Pygoscelis papua) and Antarctic fur seal (Arctocephalus gazella), was investigated. The collection of diet samples on three occasions over a 2-week period was compared with collecting all samples during a single session, as current CCAMLR Ecosystem Monitoring Programme protocols recommend. Some differences in the mass of food recovered from penguins were found but this was attributed to the mass of penguin sampled. There were no differences in diet composition between protocols and although body mass was a significant determinant of the mean length of krill Euphausia superba recovered from penguins, there were no differences between sampling protocols. This study has shown that differences between sampling frequencies are small and a variety of sampling protocols can produce results acceptable for inter-annual monitoring. The mass of sampled individuals can account for significant variation and should be recorded, especially if sampling frequencies and sizes are low. Accepted: 24 March 1999     

Dietary isotopic discrimination in gentoo penguin (Pygoscelis papua) feathers

Feathers are used commonly for stable isotope analysis to assess the foraging ecology and migration patterns of birds. However, these studies often require knowledge of species-specific feather isotopic discrimination factors (the differences in isotopic ratios between a species’ diet and feathers), which can be influenced by a species’ physiological state during molt. In this study, we determined the isotopic discrimination factors (Δ¹³C_{diet−feather} and Δ¹⁵N_{diet−feather}) between adult gentoo penguin (Pygoscelis papua) diet and feathers in a controlled study. In addition, we tested whether molt duration or the magnitude of voluntary dietary reduction during molt influenced isotopic discrimination, as previous studies have found that nutritional stress can exaggerate ¹⁵N enrichment and in some cases lead to ¹³C depletion in feathers. Contrary to this hypothesis, we found no effect of molt duration or dietary reduction on discrimination factors, suggesting that isotopic discrimination is not linearly related to these measures of fasting intensity in penguins. Furthermore, we found that the range of Δ¹⁵N_{diet−feather} found in several species of penguins, which fast while they molt, was similar to discrimination factors in fish-eating birds, which do not fast during molt. It is likely that species-specific metabolic adaptations that limit nutritional stress while fasting and variation in their relative reliance on endogenous vs. dietary pools during feather growth may confound the use of Δ¹⁵N_{diet−feather} as a general measure of nutritional stress when comparing among species.     

Genetic diversity in populations of Gentoo penguins (Pygoscelis papua)

RAPD analysis was used to examine the extent of genetic polymorphism in two populations of Gentoo penguin (Pygoscelis papua) from Antarctic Islands (Petermann and Livingston). The chosen two of three 10 mer oligonucleotide primers accordingly to preliminary results showed different levels of polymorphism in Gentoo penguins at Petermann Island (from 23.53 to 42.86%) and Livingston Island (from 52.94 to 57.14%). Nei's similarity coefficients were in range from 0.5606 (when Gentoo genome profiles were compared with RAPD profiles of two related penguin species: Pygoscelis adeliae (Adelie) and Pygoscelis antarctica (Chinstrep)) to 0.9281 among observed Gentoo penguin populations. Nei's distances values ranged from 0.0746 to 0.5787 among the populations and species. The obtained results will be used for further estimation of genetic diversity of Gentoo penguins and determination of their taxonomic status.     

Chick growth and survival in gentoo penguins (Pygoscelis papua): effect of hatching asynchrony and variation in food supply

Summary In the gentoo penguin, Pygoscelis papua, we examined the effects of intra-clutch egg size differences and hatching asynchrony on differential chick growth and survival (including post-fledging survival), in five years for which indices of food supply were available. An initial size hierarchy within-broods at hatching was due to hatching asynchrony not intra-clutch egg size differences. In 1988 only (a poor food year), the weight advantage gained by the first-hatched (A) chick persisted to the end of brooding (30 days), with more second-hatched (B) chicks dying. There was no difference between A- and B-chick weights at fledging (60 days) or in overall chick survival between synchronous and asynchronous broods in any year. Postfledging survival (measured in one year) was not related to fledging weight or hatching order. These results provide only partial support for the hypothesis that gentoo penguins operate a brood reduction strategy to optimise chick survival in years of low food availability. We suggest that hatching asynchrony in gentoo penguins may result from selection to keep the first egg warm as soon as it is laid, due to extreme low ambient temperatures.     

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.     

What is out there: diversity in feeding of gentoo penguins (Pygoscelis papua) around the Falkland Islands (Southwest Atlantic)

During austral spring 2000, the diet of Gentoo penguins, Pygoscelis papua, was studied and compared to prey availability, assessed by trawl and plankton surveys, in waters adjacent to a number of colonies along the coast of West Falkland. There was good agreement in size and abundance of key prey species in the penguin diet and from the plankton and trawl surveys. The diet of birds breeding adjacent to shallow sheltered waters was dominated by demersal species, in particular crustaceans, Munida spp. whilst birds breeding adjacent to deeper exposed waters generally preyed upon pelagic species of fish, Falkland herring Sprattus fuegensis and squid, Moroteuthis ingens. Gentoo penguins generally foraged at depths of up to 30 m, relatively close to the coast, and illustrated a certain degree of selection of prey species where possible, thus providing evidence that they are not fully opportunistic predators.     

Diving pattern and performance in relation to foraging ecology in the gentoo penguin, Pygoscelis papua

We present data on diving pattern and performance (dive depth, duration, frequency and organization during the foraging trip) in gentoo penguins Pygoscelis papua , obtained using time-depth recorders ( n = 9 birds, 99 foraging trips). These data are used to estimate various parameters of foraging activity, e.g. foraging range, prey capture rates, and are compared in relation to breeding chronology. Foraging trip duration was 6 h and 10 h, and trip frequency 1.0/day and 0.96/day, during the brooding and creche periods, respectively. Birds spent on average 52%of each foraging trip diving. Dive depth and duration were highly bimodal: shallow dives (< 21 m) averaged 4 m and 0.23 min, and deep dives (> 30 m) 80 m and 2.5 min, respectively. Birds spent on average 71%and 25%of total diving time in deep and shallow dives, respectively. For deep dives, dive duration exceeded the subsequent surface interval, but shallow dives were followed by surface intervals 2–3 times dive duration. We suggest that most shallow dives are searching/exploratory dives and most deep dives are feeding dives. Deep dives showed clear diel patterns averaging 40 m at dawn and dusk and 80–90 m at midday. Estimated foraging ranges were 2.3 km and 4.1 km during the brood and creche period, respectively. Foraging trip duration increased by 4 h between the brood and creche periods but total time spent in deep dives (i.e. time spent feeding) was the same (3 h). Of 99 foraging trips, 56%consisted of only one dive bout and 44%of 2–4 bouts delimited by extended surface intervals > 10 min. We suggest that this pattern of diving activity reflects variation in spatial distribution of prey rather than the effect of physiological constraints on diving ability.     

Population trends, breeding success and diet composition of gentoo Pygoscelis papua, magellanic Spheniscus magellanicus and rockhopper Eudyptes chrysocome penguins in the Falkland Islands. A review

Data on population size, breeding success and diet composition of gentoo (Pygoscelis papua), magellanic (Spheniscus magellanicus) and rockhopper (Eudyptes chrysocome) penguins, collected as part of the Falkland Island Seabird Monitoring Programme from 1986/1987 to 1998/1999, were analysed with regard to spatial and temporal variation, as well as potential interaction with local commercial fisheries. No significant population trends were detectable, mainly because of the short time-series and large spatial and inter-annual variation in the number of breeding pairs in the colonies monitored. However, the breeding success of all three penguin species has improved slightly over the last few years, indicating a potential for increasing populations in the near future. During the breeding season, all three penguin species preyed opportunistically on a mixture of fish, squid and crustaceans. Diet composition too showed a high degree of spatial and temporal variation. However, in all three penguin species studied, squid gradually disappeared from the diet over successive years, to be replaced by fish. Coincidentally, the commercial catches of the squid species Loligo gahi in Falkland Islands waters decreased and the by-catch of nototheniid fish increased. All three penguin species compete directly with the commercial fishing fleet for L. gahi; however, there may also be competition for Patagonian toothfish (Dissostichus eleginoides), hake (Merluccius sp.) and southern blue whiting (Micromesistius australis), because juveniles of these species were found regularly in penguin diets.     

Diving patterns of female macaroni penguins breeding on Marion Island, South Africa

Despite the large biomass of macaroni penguins Eudyptes chrysolophus in the Southern Ocean, their feeding ecology is poorly known at some important breeding localities. We investigated the diving behaviour and diet of female macaroni penguins feeding small chicks on Marion Island (46o52′S, 37o5′E), South Africa, one of the species’ most northerly breeding sites, supporting 4% of their global population. We then compared our results with similar studies from other localities. In December 2008, we collected information on 12 foraging trips from 6 individuals using time-depth recorders, as well as diet from 42 individuals. Median trip duration was 22.8 h (5.6–80.8 h). Penguins performed 42.8 ± 15.9 dives per hour at sea, with dive depths averaging 24.6 ± 8.6 m and lasting 40.8 ± 12.1 s, although 74.3% of dives were <10 m. Euphasids dominated their diet (86% by mass), mainly Thysanoessa vicina. A second peak in dive depths at 55–80 m might reflect the 12% of fish in their diet. The substantial proportion of shallow night dives (30% of total dives) suggests some foraging occurs at night. Differences in diving patterns of individual macaroni penguins in this study confirmed the behavioural flexibility of these birds reported from other breeding localities. However, most other studies assumed that dives <3–5 m were commuting dives whereas our study suggests that at least some prey are caught during shallow dives. We highlight how different analytical methods can change the outcome of studies. Despite macaroni penguins’ apparent flexibility in foraging behaviour during the breeding season, their numbers are decreasing globally. Further investigations of their foraging behaviour are needed to assess potential competition with other predators and krill fisheries.