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 .     

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.     

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.     

Changes in Adélie penguin breeding populations in Lützow-Holm Bay, Antarctica, in relation to sea-ice conditions

The Adélie penguin, Pygoscelis adeliae, an important component of the Antarctic marine ecosystem, is closely associated with sea ice. Using data collected by Japanese Antarctic Research Expeditions since the 1960s, we examined trends in breeding populations of this species around Lützow-Holm Bay. Ten colonies ranging in size from 10 to 2,500 individuals were counted along the Soya Coast. Populations fluctuated synchronously, and overall increased at most colonies, except for two: one located deep inside the bay and another where human disturbance was substantial. Populations tended to increase during, or after, periods of sparse sea ice in summer, a condition that occurred once every decade. An increase in population size also occurred 5 years after a winter of extensive sea ice and after a winter of especially reduced sea ice.     

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.     

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.     

Prolactin and parental behavior in Adélie penguins: effects of absence from nest, incubation length, and nest failure

Adélie penguin (Pygoscelis adeliae) males and females, nesting in Antarctica, alternate attendance at the nest with absences of many days to forage at sea. We investigated the importance of tactile input from egg and chicks on prolactin levels by observing nest attendance patterns and obtaining blood samples (1) during the first nest exchange of the incubation stage, (2) from birds whose incubation period was artificially increased or decreased by about 10 days, and (3) from birds whose nests had failed. Prolactin levels in females after 8 to 11 days of absence from the breeding colony did not differ from those in incubating males and did not change after females resumed incubation. Moving eggs between nests resulted in nests in which chicks hatched after about 26, 36 (normal), or 46 days. Duration of incubation did not affect prolactin levels in the parents measured during incubation, at the pip stage, hatch stage, or early brood stage. Adults first left their chicks unguarded on about the same calendar date, regardless of chick age. However, chicks from long incubation nests averaged 8 days younger when they were left unguarded than chicks from control or short-incubation nests. In females, there was no effect of nest failure on prolactin levels. In males, prolactin levels were slightly lower after nest failure than in males tending nests. Testosterone was significantly higher in males after nest failure than in males still tending nests. Prolactin is elevated in Adélie penguins as part of the program of cyclical hormonal changes that accompany the lengthy reproductive season and is relatively independent of tactile input. Sustained prolactin secretion is probably required for the maintenance of parental behavior in offshore feeding species that must be absent from the nest for many days at a time.     

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.     

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.     

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.     

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.     

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     

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.     

Foraging rhythms in Adélie penguins (Pygoscelis adeliae) at hope bay,Antarctica; determination and control

Summary We studied diel periodicity in activity of Adélie penguins (Pygoscelis adeliae) during the austral summer solstice and one month later by counting, hourly, numbers of birds leaving colonies to forage, numbers arriving back after foraging, numbers in a colony and numbers resting outside the colonies. During the solstice large numbers of birds were arriving at and departing from the colony at all times of the day although there was a tendency for more birds to be at sea when light intensity was highest at mid-day. Generally, birds not brooding chicks did not rest on land. A month later, when visibility was poor at mid-night, the percentage of birds at sea was highly positively correlated with light intensity. Birds returning from foraging in the evening fed chicks immediately and then either rested in the colonies or on snow patches between the colonies and the sea until the following morning.     

Alloparental feeding in Adélie penguins: why is it uncommon?

We investigated alloparental interactions and conditions which could facilitate or prevent the expression of alloparental behaviours in Adélie penguins (Pygoscelis adeliae), a long-lived seabird which nests in high-density colonies around Antarctica. Observation sessions were carried out during the crèche stage on 48 identified pairs and 50 identified chicks in a 217-nest subcolony. As the season progressed, young were fed less often by their own parents because these were increasingly absent from the breeding site and less responsive to their offspring’s solicitations. As a consequence, young and particularly those with a low body mass, coming from a two-chick brood, opted for gradually soliciting more from other adults to obtain food, preferentially those nesting in their direct vicinity. Unsuccessful breeders represented a low and constant part of the adult population and were not specifically solicited by unrelated young. Despite the increasing chick demand, only 4.1% (3 out of 73) of alloparental solicitations resulted in feeding, which is negligible compared to parental feeding. To investigate factors that could trigger the appearance of alloparental care, we carried out comparisons with king (Aptenodytes patagonicus) and emperor penguins (Aptenodytes forsteri) which represent the closest species for which data on alloparental behaviour were available. Our results show different trends to those observed in these species and three factors may explain the low occurrence of alloparental behaviour in Adélie penguins: (1) the low and constant proportion of unsuccessful breeders, (2) the absence of chick selectivity towards unsuccessful breeders, and (3) the late period of chick accessibility for potential alloparents.     

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     

Satellite telemetry of the winter migration of Adélie penguins (Pygoscelis adeliae)

Adélie penguins (Pygoscelis adeliae), after breeding in Antarctica during the austral summer, undergo a winter migration before returning to the breeding grounds 8 months later. It is the major source of adult mortality, with about a quarter of them not returning. Here we describe the first attempt to track the winter migration of Adélie penguins using satellite telemetry. Transmitters were attached to two penguins on 16 February 1991 after their post-breeding moult at Cape Bird, Ross Island, Antarctica. Transmissions were received from one penguin (bird #1) for 4.4 months, during which time it travelled 2792.6 km from the rookery (nearly 1500 km straight-line distance). Transmissions were received from the other penguin (bird #2) for 2.5 months during which time it followed a path remarkably similar to that of bird #1. The penguins travelled northwards up the coast of Victoria Land, keeping within 100 km of the coast, rounding Cape Adare soon after 29 March and were midway between the Balleny Islands and the Antarctic coast on 3 May. Thereafter, the record from bird #1 shows that it travelled further westwards until, when opposite the Mastusevich Glacier Tongue of the Mastusevich Glacier, it turned due north and moved away from the coast. By 29 June, when transmissions ended, its progression had slowed and it was northwest of the Balleny Islands near a zone where pack ice covered 75% of the surface of the sea. Two novel points that arise from this study are: (1) that Adélie penguins from Cape Bird undergo winter migrations of not less than 5000 km, and (2) that they may be travelling to common overwinter feeding grounds.     

Nonlinear effects of winter sea ice on the survival probabilities of Adélie penguins

The population dynamics of Antarctic seabirds are influenced by variations in winter sea ice extent and persistence; however, the type of relationship differs according to the region and the demographic parameter considered. We used annual presence/absence data obtained from 1,138 individually marked birds to study the influence of environmental and individual characteristics on the survival of Adélie penguins Pygoscelis adeliae at Edmonson Point (Ross Sea, Antarctica) between 1994 and 2005. About 25% of 600 birds marked as chicks were reobserved at the natal colony. The capture and survival rates of Adélie penguins at this colony increased with the age of individuals, and five age classes were identified for both parameters. Mean adult survival was 0.85 (SE = 0.01), and no effect of sex on survival was evident. Breeding propensity, as measured by adult capture rates, was close to one, indicating a constant breeding effort through time. Temporal variations in survival were best explained by a quadratic relationship with winter sea ice extent anomalies in the Ross Sea, suggesting that for this region optimal conditions are intermediate between too much and too little winter sea ice. This is likely the result of a balance between suitable wintering habitat and food availability. Survival rates were not correlated with the Southern Oscillation Index. Low adult survival after a season characterized by severe environmental conditions at breeding but favorable conditions during winter suggested an additional mortality mediated by the reproductive effort. Adélie penguins are sensitive indicators of environmental changes in the Antarctic, and the results from this study provide insights into regional responses of this species to variability in winter sea ice habitat.     

A method for estimating colony sizes of Adélie penguins using remote sensing imagery

Adélie penguins (Pygoscelis adeliae) are important predators of krill (Euphausia spp.) and Antarctic silverfish (Pleuragramma antarctica) during summer, are a key indicator of the status of the Southern Ocean ecosystem, and are therefore a focal species for the Committee for the Conservation of Antarctic Marine Living Resources (CCAMLR) Ecosystem Monitoring Program. The ability to monitor the population size of species potentially affected by Southern Ocean fisheries, i.e., the Adélie penguin, is critical for effective management of those resources. However, for several reasons, direct estimates of population size are not possible in many locations around Antarctica. In this study, we combine high-resolution (0.6 m) satellite imagery with spectral analysis in a supervised classification to estimate the sizes of Adélie penguin breeding colonies along Victoria Land in the Ross Sea and on the Antarctic Peninsula. Using satellite images paired with concurrent ground counts, we fit a generalized linear mixed model with Poisson errors to predict the abundance of breeding pairs as a function of the area of current-year guano staining identified in the satellite imagery. Guano-covered area proved to be an effective proxy for the number of penguins residing within. Our model provides a robust, quantitative mechanism for estimating the breeding population size of colonies captured in imagery and identifies terrain slope as a significant component influencing apparent nesting density. While our high-resolution satellite imagery technique was developed for the Adélie penguin, these principles are directly transferrable to other colonially nesting seabirds and other species that aggregate in fixed localities.