Sea ice affects the population dynamics of Adélie penguins in Terre Adélie

Overall Adélie penguin population size in Pointe Géologie Archipelago increased between 1984 and 2003 at a rate of 1.77% per year, and averaged 33,726±5,867 pairs. As predicted by the optimum model proposed by Smith et al. (Bioscience 49:393–404, 1999). Adélie penguin population size increased when sea ice extent and concentration (SIE and SIC) decreased six years earlier, indicating that the conditions around reproduction or first years at sea, were determinant. The breeding success averaged 85.2±35.45% and was not related to environmental variables. Adult survival probability varied between years from 0.64 to 0.82. Southern oscillation index (SOI) had a strong negative effect on adult annual survival. Adult survival of Adélie penguins increased during warmer events, especially during winter and spring at the beginning of reproduction. Therefore, we speculate that the rapid decreases in 1988–1991 and 1996 of the breeding population size were related to a decrease in adult mortality. However, adult survival varied little, and could not explain the strong increasing population trend. The sea ice conditions during breeding or during the first year at sea appeared determinant and influenced the population dynamics through cohort effects, probably related to the availability of productive feeding habitats.     

Diving and foraging behaviour of Adélie penguins in areas with and without fast sea-ice

The diving and foraging behaviours of Adélie penguins, Pygoscelis adeliae, rearing chiks at Hukuro Cove, Lützow-Holm Bay, where the fast sea-ice remained throughout summer, were compared to those of penguins at Magnetic Island, Prydz Bay, where the fast sea-ice disappeared in early January. Parent penguins at Hukuro Cove made shallower (7.1–11.3 m) but longer (90–111 s) dives than those at Magnetic Island (22.9 m and 62 s). Dive duration correlated with dive depth at both colonies (r ² = 0.001 ∼ 0.90), but the penguins atg Hukuro Cove made longer dives for a given depth. Parents at Hukuro Cove made shorter foraging trips (8.1–14.4 h) with proportionally longer walking/swimming (diving < 1 m) travel time (27–40% of trip duration) and returned with smaller meals (253–293 g) than those at Magnetic Island, which foraged on average for 57.2 h, spent 2% of time walking/swimming ( < 1 m) travel, and with meals averaging 525 g. Trip duration at both colonies correlated to the total time spent diving. Trip duration at Hukuro Cove, but not at Magnetic Island, increased as walking/swimming ( < 1 m) travel time increased. These differences in foraging behaviour between colonies probably reflected differences in sea-ice cover and the availability of foraging sites. Received: 3 November 1995/Accepted: 29 May 1996     

Impacts of human visitors on breeding success and long-term population trends in Adélie Penguins at Casey,Antarctica

Breeding populations of Aélie Penguins Pygoscelis adeliae have been counted at two localities near Casey Station in Wilkes Land, East Antarctica since 1959/60 and 1968/69. At Whitney Point, the breeding population increased from 1122 pairs in 14 colonies in 1959/60 to 4714 pairs in 36 colonies in 1992/93. All new colonies at Whitney Point established on relict colony sites identified in 1959/60. On Shirley Island, the total breeding population has remained at 7770 pairs ±10% between 1968/69 and 1992/93, except in 1990/91 when the population peaked at 8719 pairs. An association between the age of a colony and its rate of increase was observed at Whitney Point, with new colonies (those established since 1971/72) increasing more rapidly than colonies extant in 1959/60. At Shirley Island, where most of the colonies extant in 1968/69 have decreased in population, the establishment and growth of 13 colonies has offset this decrease; these new colonies also exhibited the association between age and rate of increase. Breeding success (chicks fledged per nest) was significantly lower for Shirley Island colonies than for those at Whitney Point. Human visitors to Shirley Island from Casey station are believed to be responsible for the observed changes in the distribution and abundance of breeding pairs and for maintaining the stable population by reducing overall breeding success through the disturbance associated with visits.     

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.     

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.     

Synchronous diving behavior of Adélie penguins

Synchronizing behavior with other conspecifics has been suggested as serving a function of increased foraging efficiency. However, the potential costs associated with synchronization of behavior have rarely been studied. Adélie penguins Pygoscelis adeliae sometimes dive synchronously in small open waters surrounded by fast sea ice. We examined the diving behavior of three couples and one trio, which were observed to dive synchronously among groups of 12–47 birds for 1.7–4.5 h duration, with time-depth recorders. Timing of diving and surfacing differed slightly between individuals, and one bird tended to initiate diving earlier than the other. Although the duration of the dives differed only slightly between these birds, the maximum depth of the dives differed to a large extent, with one member tending to dive consistently deeper than the other bird in two out of the four cases. Vertical distances between tagged birds in the undulatory phases of the dives (presumed feeding time) were greater than those in the descent and ascent phases, suggesting independent foraging by group members. Duration of the undulatory phase of the dives tended to be shorter in deeper-diving individuals than the others in the synchronously diving group, suggesting a potential cost of reduced feeding time to synchronize diving and surfacing with other birds. A digital video image relating to the article is available at .     

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.     

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

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

The gastrointestinal tract of Adélie penguins – morphology and function

The aim of this study was to provide data on the morphology of the gastrointestinal tract of Adélie penguins (Pygoscelis adeliae). It was found to consist of a long oesophagus, a two-chambered stomach, a small intestine measuring only 5.22body length, two rudimentary caeca and a short colon, typical of carnivorous birds. The stomach comprised a glandular proventriculus and a muscular gizzard that frequently contained grit. An acidic pH was recorded in both chambers. Ultrastructural studies of the small intestinal mucosal membrane revealed epithelial cells with elongated, irregular microvilli and high affinity for toluidine blue, absorptive intestinal epithelial cells and goblet cells. Numerous large lymphocyte-like cells were observed close to the brush border of the epithelium, and empty spaces on the epithelial surface reflected normal cell loss in the small intestine. The rudimentary caeca and colon provide relatively little volume and time for symbiotic bacteria to aid the digestion of crustacean chitin.     

Strategic allocation of ejaculates by male Adélie penguins

Sperm competition theory suggests that males should strategically allocate sperm to those females that will bring them the best possible genetic returns. Although males of a number of species of insects and fishes have been shown to allocate sperm strategically, we provide, to our knowledge, the first evidence that an avian species is also capable of allocating ejaculates. Male Adélie penguins (Pygoscelis adeliae) are more likely to transfer sperm during extra-pair copulations (EPCs) than during pair copulations. We investigated the question of how males allocate ejaculates within the constraints of limited sperm availability and found (i) that males that engaged in EPC attempts ejaculated less often when copulating with their social partner than males that made no EPC attempts, and (ii) that there was no difference between males that were involved in failed EPC attempts and those that were involved in successful EPCs in the proportion of copulations that resulted in sperm transfer. These results indicate that males achieve strategic allocation of sperm within the constraints of limited sperm availability by withholding ejaculates from their social partners.     

Annual and seasonal changes in foraging site and diving behavior in Adélie penguins

Foraging sites, diet, and diving behavior of chick-rearing Adélie penguins, Pygoscelis adeliae, in fast sea-ice areas were investigated during two consecutive seasons with contrasting sea-ice conditions. During 1995/1996, fast sea ice covered the foraging range of penguins during the whole breeding season. In contrast, during 1996/1997, sea ice covered the area in December 1996, but gradually thinned and finally broke up, so that open sea appeared along the coast during February 1997. Foraging sites were concentrated in a small area in 1995/1996 and spread over a wider area in 1996/1997 as more small open-water areas were available. In both seasons, parents traveled to more distant foraging sites as the season progressed and, consequently, the foraging-trip duration increased. In both years, Euphausia superba and Pagothenia borchgrevinki dominated the diet in the early part of the season, while later in the season penguins fed mainly on E. superba in 1995/1996 and Pagothenia borchgrevinki and E. crystallorophias in 1996/1997. In 1995/1996, penguins tended to dive deeper—albeit for a relatively shorter duration—when feeding mainly on krill compared to when feeding on fish. In 1996/1997, there was no difference in the dive depth and duration between krill- and fish-eating trips. Our results suggest that prey distribution changes annually and seasonally, probably according to sea-ice conditions, and that consequently penguins modify their foraging sites, diving patterns, and diet according to these changes.     

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     

Adverse effects of instrumentation in incubating Adélie penguins (Pygoscelis adeliae)

The use of data-loggers has permitted to explore the biology of free-ranging animals. However, this method has also been reported to reduce reproductive success while the reasons of this deleterious effect remain poorly documented. In this study, we aimed to identify critical periods of the breeding cycle of Adélie penguins (Pygoscelis adeliae) when the reproductive success may decrease because of instrumentation. For this purpose, we monitored 40 pairs, where one parent was instrumented before egg laying and 30 pairs without devices (controls). These pairs were followed at least during the incubation period but the majority was monitored during the entire breeding season. Reproductive success was affected in pairs where males were instrumented. This was not due to extra chick mortality during chick rearing but to a significantly lower hatching success. Moreover, the use of artificial eggs recording incubation temperatures and egg rotation indicated that in instrumented incubating males, eggs spent as much time at optimal incubation temperatures as control eggs but were rotated at a higher frequency. In Adélie penguins, males initiate incubation and it has been established that the early stage of incubation is one of the most critical periods for embryonic development. The low hatching rate observed in instrumented males was associated with a higher egg rotation rate, perhaps as a stress response to the presence of the instrument. Even though the causal effects remain unclear, instrumentation severely affected hatching success. For these reasons, we recommend equipping birds after the early incubation.     

Measuring the influence of unmanned aerial vehicles on Adélie penguins

Unmanned aerial vehicles (UAVs) have become a useful tool in polar research. While their performance is already proven, little is known about their impact on wildlife. To assess the disturbance caused on the penguins, flights with a UAV were conducted over an Adélie penguin (Pygoscelis adeliae) colony. Vertical and horizontal flights were performed between 10 and 50 m in altitude. Penguins’ reactions were video-recorded, and the behavioural response was used to indicate the level of disturbance. During any flight mode, disturbance increased immediately after takeoff and remained elevated at all altitudes between 20 and 50 m. When the UAV descended below 20 m, the disturbance increased further with almost all individuals being vigilant. Only at these low altitudes, vertical flights caused an even higher level of disturbance than horizontal ones. Repetitions of horizontal overflights showed no short-term habituation occurring. Since the results are only valid for the specific UAV model used, we recommend a more extensive approach with different UAV specifications. As the highest flight altitudes already caused detectable but not subjectively visible responses, we also recommend to regard subjective impressions of disturbance with caution.     

Using logistic regression models to predict breeding success in male Adélie penguins (Pygoscelis adeliae)

Measures of breeding success are traditionally derived using the proportion of total nests that successfully reach strategic stages across a breeding season, such as pair formation, egg laying or fledging chicks. The use of logistic regression has recently become a popular tool in avian literature for identifying influential factors that predict nest and/or individual breeding success. In this study, we use logistic regression models to assess the importance of a range of factors affecting male Adélie penguin (Pygoscelis adeliae) breeding success during the 2002–2003 austral summer, when the presence of icebergs and extensive sea-ice meant overall breeding success of Adélie penguins was low (16/51 focal males had chicks reach fledging point). Logistic regression models for the early breeding season showed that good/average nest quality and central/middle nest location within the sub-colony were the best predictors of successful pair formation. Later, during incubation, the most successful males were those that not only returned earlier but were also heavier upon arrival and built nests of at least average quality. During the final stage when chicks had begun to fledge, the combined parameters of heavier male weight, early arrival time and good nest quality were the best predictors of breeding success. The logistic regression approach used here showed that the predictive ability of these parameters varied as the season progressed. However, most importantly, our logistic models fit the data well across all breeding stages.     

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.     

Parental mass change and food provisioning in Adélie penguins rearing chicks in colonies with contrasting sea-ice conditions

Adélie penguins (Pygoscelis adeliae) changed the pattern of energy allocation for self-maintenance and food provisioning among three colonies. Parents made longer trips (44-57 h) at Davis, where they foraged in open waters, than in Syowa (17-22 h) where fast sea-ice remained. Those at Dumont d'Urville, where the sea-ice disappeared in mid-summer, made trips of variable duration (22-41 h). During foraging trips, parents at Davis accumulated a greater amount of body tissue (0.40 kg) than parents at other locations (-0.06 to 0.13 kg) to offset the loss of body tissue during previous chick guarding. As a consequence, parents at these colonies had a similar body-mass decrease rate throughout the chick-rearing period (10-19 g/day). Parents with a smaller departure body mass seemed to make longer trips at Davis and Dumont d'Urville, and those making longer trips seemed to accumulate more body tissue at Davis, though these effects were not observed at Syowa. Stomach-content mass was greater at Davis (457 g) and Dumont d'Urville (551 g) than at Syowa (271-363 g), while the frequency of colony visits with meals was smaller at Davis (0.31/day per bird) and Dumont d'Urville (0.39/day per bird) than at Syowa (0.64-0.68/day per bird). Stomach-content mass was independent of trip duration. We propose the hypothesis that parents maintain their own body condition by regulating the accumulation of body tissue during foraging trips but change food provisioning with respect to variable foraging-trip duration.     

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     

Diet and foraging effort of Adélie penguins in relation to pack-ice conditions in the southern Ross Sea

We investigated the diet and aspects of foraging effort among Adélie penguins (Pygoscelis adeliae) breeding at three colonies on Ross Island, in the southwestern Ross Sea – Capes Royds, Bird and Crozier – during the chick-provisioning period of three austral summers, 1994–1995, 1995–1996 and 1996–1997. During the study period, pack-ice cover differed in waters offshore of these colonies, by colony, seasons and year. Diet differed among colonies only slightly. The fish Pleuragramma antarcticum was the most important prey, especially during years or periods within years when little pack ice was present. With respect to krill, which composed the remainder of diet, juvenile Euphausia crystallorophias were consumed predominantly in a year of heavy pack-ice cover; more adult krill were consumed in 2 years when pack ice was sparse. Foraging trip duration differed by colony, season and year and was related directly to distance from the colony to the nearest pack ice. The amount of food brought to chicks increased as trip duration increased, to a point (2 days), but then decreased as duration increased further (up to 4 days). On the basis of data on mass of parents and of meal sizes to chicks, it appeared that on the longest trips more of the food gathered by parents was used for self maintenance; on the longest trips, parents lost body mass. Successful foraging during chick rearing, the period when adult foraging is most intense, appears to depend on the proximity of pack ice to nesting colonies for this penguin species. Received: 1 October 1997 / Accepted: 25 April 1998     

Foraging behavior of Adélie penguins during incubation period in Lützow-Holm Bay

The diving behavior of Adélie penguins Pygoscelis adeliae was investigated using time–depth recorders during the incubation period in the fast sea-ice area of Lützow-Holm Bay, Antarctica. Dive profiles and activity/time allocation suggested that penguins were obligated to walk on the fast-ice for 90–100 km until a polynya, which they used as an access to the pack-ice zone. Dive depth did not differ between males and females, though males’ dive duration was longer than that of females. Dive depth was slightly shallower and dive duration was shorter during the incubation than during the chick-rearing phase. Birds dove throughout the day, although less frequently around midnight, and there was no clear diel change in dive depth. This daily dive pattern during incubation period was similar to that previously observed during the chick-rearing period in a fast sea-ice area, but differed from that observed in sea-ice-free area. Variations in diving behavior resulted from different environmental conditions, such as foraging area with different sea-ice condition, as well as from different life history strategies.