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     

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.     

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.     

Holocene Adélie penguin diet in Victoria Land,Antarctica

Ornithogenic soils (N = 97) dated up to 7000 Before Present (bp) were sampled in 16 relict and modern breeding colonies of Adélie penguin along the Victoria Land coast (Ross Sea, Antarctica). Taxonomic identification of fish otoliths (N = 677) recovered in these soils allowed to identify the Antarctic silverfish as the most eaten prey (90.1%) throughout the investigated period. A morphometric analysis of the otoliths revealed that the Adélie penguin primarily selected prey averaging 67.23 ± 23 mm of standard length. Temporal distribution of Pleuragramma antarcticum showed a peak between 2,000 and 4,000 years bp, a period corresponding to the maximum spread of Adélie penguin in the Victoria Land. Possible explanations of the variations of the abundance of the fish prey in the diet are discussed in the context of the paleoclimatic events and as possible consequences of dietary shifts due to the temporal variation of prey availability in the Ross Sea ecosystem.     

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

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

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.     

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 .     

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.     

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.     

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.     

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.     

Breeding success and population trends in Adélie penguins in areas with low and high levels of human disturbance

The breeding performance and population trends of Adélie penguins (Pygoscelis adeliae) was studied at Esperanza/Hope Bay, Antarctic Peninsula, by comparing an area with low levels of human disturbance (LLD) and an area with high levels of human disturbance (HLD), close to an Argentine research station. From 1995/1996 to 2004/2005 (except for 1999/2000 and 2003/2004), the following population parameters were measured in both areas: (1) the number of breeding pairs, (2) the number of chicks in creches and (3) the number of chicks produced by breeding pairs. Counts were made for 26 breeding groups situated in the LLD area and 63 breeding groups located in the HLD area. The number of chicks per breeding pair was obtaobtained by following 100 marked nests in each area. All parameters were measured as described in the CCAMLR Monitoring Program protocols. The magnitude and direction (increasing or decreasing) of the changes in breeding population size and the number of chicks creched were similar in both areas. Overall, the number of breeding pairs decreased from 4,744 to 2,968 (37.4%) in the LLD area, and from 8,744 to 5,378 (38.6%) in the HLD area. The number of chicks fledged increased from 3,808 to 4,065 (6.7%) in the LLD area, and decreased from 6,991 to 6,712 (4%) in the HLD area. Breeding success (chicks fledged per marked nest) did not differ significantly between areas for most of the seasons compared. In 1996/1997, breeding success was significantly higher in the HLD area. Our data suggest that environmental influences currently exert greater effects than human disturbance on the penguin population at Esperanza Bay.     

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.     

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.     

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.     

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.     

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.     

Community structure of pelagic cnidarians off Adélie Land, East Antarctica, during austral summer 2008

Some studies have suggested that pelagic cnidarians are important components of the Southern Ocean ecosystem due to their high abundance and diversity and their high predatory effects, although little information on these animals is available. Thus, we examined the spatial distribution of pelagic cnidarians from the oceanic to neritic zone off Adélie Land, East Antarctica. Discrete depth sampling was conducted from the surface to 2,000 m depth from late January to early February 2008. In total, 3347 individuals representing 45 species/taxa from eight orders were collected. Cluster analysis revealed three major clusters: (1) an epipelagic group in the oceanic zone composed mainly of Pegantha martagon, the abundance and species diversity of which were very low; (2) a meso- and bathy-pelagic group characterised by high abundance and species diversity with dominance of Dimophyes arctica, Vogtia serrata, and Halicreas minimum; and (3) a neritic group represented by a high abundance of Diphyes antarctica. Cnidarian communities in the epipelagic zone were divided by hydrographic structures such as the Southern Boundary of the Antarctic Circumpolar Current and Antarctic Slope Front, whereas those in the meso- and bathy-pelagic layers dominated by circumpolar deep water were relatively stable with higher diversity throughout the oceanic zone.     

Analysis of the sources of variance in the mean size of krill consumed by Chinstrap and Adélie penguins at South Orkney Islands

The CCAMLR Ecosystem Monitoring Program aims at detecting possible impacts of the krill fishery on predator populations. Population parameters of Chinstrap and Adélie penguins are monitored at Point Martin, Laurie Island, South Orkneys Islands as part of the Program. Diet information is collected to aid in the interpretation of population data. In the present study, the diet of both species is compared in order to detect differences between them as well as to ascertain whether the krill consumed by both species might be pooled in single estimates of prey parameters. A nested ANOVA model was fitted to the dataset obtained in the seasons from 1997/1998 to 2005/2006. The overall variance was decomposed into variance components: within penguins (krill in stomachs), between penguins (means within each date), between dates (date means in each season), and between years (annual means). The within penguins component was similar in both species (18?% difference), while the between penguins was 64?% larger in Adelies. The within and between penguins variance components of both species showed consistent interannual changes. The annual means of both species differed significantly in three seasons, but not consistently. The sources of variance are tentatively attributed to biological characteristics: the within penguins component would reflect searching and feeding at the individual level; the between penguins might be associated with their foraging ranges, an assumption to be tested tracking individuals. Given their differences, the information from both species should be viewed as complementary and not be pooled.     

Inter-colony and sex differences in the effects of parental body condition and foraging effort on the brood growth of Adélie penguins

Among colonies with different foraging distances, central-place-foraging seabirds may change their foraging and reproductive efforts. We compared the body condition, meal frequency, and diving behavior of male and female Adélie penguins at two locations: Dumont d'Urville, where there was little sea ice and they foraged in open waters far from the colony; and Syowa, where there was heavy, fast sea ice and they foraged in ice cracks close to the colony. The parental mass decrease rate during the chick-rearing period was similar between colonies and between sexes. A large individual variation in meal frequency positively affected the brood growth rate, but daily underwater time did not. A weak but significant positive effect of body condition on brood growth rate was found only in males at Syowa. It was suggested that males work with better body condition than females. We propose the hypothesis that the regional difference in the distance to the feeding sites and the sex difference in body energy reserve might constrain the capacity to regulate reproductive effort.