Path characteristics and colony arrival-departure statistics of king penguins (Aptenodytes patagonicus) on Possession and Kerguelen Island

Orientation based on visual cues can be extremely difficult in crowded bird colonies due to the presence of many individuals. We studied king penguins (Aptenodytes patagonicus) that live in dense colonies and are constantly faced with such problems. Our aims were to describe adult penguin homing paths on land and to test whether visual cues are important for their orientation in the colony. We also tested the hypothesis that older penguins should be better able to cope with limited visual cues due to their greater experience. We collected and examined GPS paths of homing penguins. In addition, we analyzed 8 months of penguin arrivals to and departures from the colony using data from an automatic identification system. We found that birds rearing chicks did not minimize their traveling time on land and did not proceed to their young (located in creches) along straight paths. Moreover, breeding birds' arrivals and departures were affected by the time of day and luminosity levels. Our data suggest that king penguins prefer to move in and out of the colony when visual cues are available. Still, they are capable of navigating even in complete darkness, and this ability seems to develop over the years, with older breeding birds more likely to move through the colony at nighttime luminosity levels. This study is the first step in unveiling the mysteries of king penguin orientation on land.

At surface behaviour of emperor penguins from Pointe Géologie, Adélie Land, Antarctica, from expedition DDU 2005

The foraging distributions of 20 breeding emperor penguins were investigated at Pointe Géologie, Terre Adélie, Antarctica by using satellite telemetry in 2005 and 2006 during early and late winter, as well as during late spring and summer, corresponding to incubation, early chick-brooding, late chick-rearing and the adult pre-moult period, respectively. Dive depth records of three post-egg-laying females, two post-incubating males and four late chick-rearing adults were examined, as well as the horizontal space use by these birds. Foraging ranges of chick-provisioning penguins extended over the Antarctic shelf and were constricted by winter pack-ice. During spring ice break-up, the foraging ranges rarely exceeded the shelf slope, although seawater access was apparently almost unlimited. Winter females appeared constrained in their access to open water but used fissures in the sea ice and expanded their prey search effort by expanding the horizontal search component underwater. Birds in spring however, showed higher area-restricted-search than did birds in winter. Despite different seasonal foraging strategies, chick-rearing penguins exploited similar areas as indicated by both a high 'Area-Restricted-Search Index' and high 'Catch Per Unit Effort'. During pre-moult trips, emperor penguins ranged much farther offshore than breeding birds, which argues for particularly profitable oceanic feeding areas which can be exploited when the time constraints imposed by having to return to a central place to provision the chick no longer apply.

(Table 1) Diving behaviour of male Adélie penguins (Pygoscelis adeliae) with and without corticosterone implants at Pointe Géologie

The amount of energy that organisms can allocate to self-maintenance and/or reproduction largely depends on their foraging strategies. Because of corticosterone (CORT) involvement in the control of energy metabolism, food intake and locomotor activity, recent studies have sought to demonstrate the role of this hormone in foraging decisions and performance. Moreover, considerable recent advances in animal-attached loggers now allow the study of behaviour in free-living animals. In order to assess the effects of CORT administration on the foraging behaviour of free-living Adelie Penguins Pygoscelis adeliae, we studied a group with CORT implants and a control group without CORT implants, by attaching time-depth recorders to the two groups and monitoring them throughout up to seven consecutive foraging trips during the guard stage (in Adelie Land, Antarctica). We found that foraging trips duration was similar between both groups. Dive durations, time spent at the bottom phase of dives, and the number of undulations per dive of CORT-implanted birds were all significantly higher than those of controls. However, CORT-implanted birds performed fewer dives overall (ca. 4,400) than controls (ca. 6,250) and spent many (13 and 6 times for penguins #3 and #4, respectively) long periods (>3 h) without diving. The low foraging effort and long resting periods support the view that CORT-implanted birds probably gained less energy than did the control birds. CORT treatment appears then to result in redirecting bird behaviour from costly activity (i.e. reproduction) to a behaviour promoting the preservation of energy reserves. Future studies are therefore needed to assess body condition and reproductive success of CORT-manipulated birds in parallel with the recording of their diving performances.

(Table 1) Foraging trip characteristics of male Adélie penguins (Pygoscelis adeliae) during their first incubation trip at Pointe Géologie

Knowledge of habitat use by top marine predators in response to environmental conditions is crucial in the current context of global changes occurring in the Southern Ocean. We examined the at-sea locations of male Adelie penguins (Pygoscelis adeliae) breeding at Dumont d'Urville during their first, long incubation trip. Compared with the chick-rearing period, penguins performed longer trips, going to oceanic waters as far as 320 km from the colony. We observed 3 strategies: (1) five individuals covered large distances to the north, targeting open-ocean areas and following the currents of two persistent eddies; (2) five individuals foraged to the north-west, close to the Antarctic shelf slope at the limit of the pack ice; and (3) three individuals covered much shorter distances (northwards or eastwards). The foraging range also seemed to be limited by the body condition of the penguins before their departure to sea.

(Table 1) Breeding parameters for 60 male Adélie penguins (Pygoscelis adeliae) at station Dumont d'Urville (Antarctica) depending on treatment (control vs. CORT)

Corticosterone, the main stress hormone in birds, mediates resource allocation, allowing animals to adjust their physiology and behaviour to changes in the environment. Incubation is a time and energy-consuming phase of the avian reproductive cycle. It may be terminated prematurely, when the parents' energy stores are depleted or when environmental conditions are severe. In this study, the effects of experimentally elevated baseline corticosterone levels on the parental investment of incubating male Adelie penguins were investigated. Incubation duration and reproductive success of 60 penguins were recorded. The clutches of some birds were replaced by dummy eggs, which recorded egg temperatures and rotation rates, enabling a detailed investigation of incubation behaviour. Corticosterone levels of treated birds were 2.4-fold higher than those of controls 18 days post treatment. Exogenous corticosterone triggered nest desertion in 61% of the treated birds; consequently reducing reproductive success, indicating that corticosterone can reduce or disrupt parental investment. Regarding egg temperatures, hypothermic events became more frequent and more pronounced in treated birds, before these birds eventually abandoned their nest. The treatment also significantly decreased incubation temperatures by 1.3 °C and lengthened the incubation period by 2.1 days. However, the number of chicks at hatching was similar among successful nests, regardless of treatment. Weather conditions appeared to be particularly important in determining the extent to which corticosterone levels affected the behaviour of penguins, as treated penguins were more sensitive to severe weather conditions. This underlines the importance of considering the interactions of organisms with their environment in studies of animal behaviour and ecophysiology.

(Table 1) Egg laying and hatching dates and guarding duration of Adélie penguins (Pygoscelis adeliae) on Petrel Island, Antarctica

1. In Polar Regions, the extent and dynamics of sea-ice are changing. This affects the ocean productivity which consecutively impacts plankton communities and polar top predators like penguins. Yet, the underlying behavioural and physiological mechanisms remain poorly understood. 2. Here we monitored the ecophysiological responses of Adelie penguin (Pygoscelis adeliae) pairs during two seasons of contrasting timing of sea-ice retreat. Beside classical breeding parameters like foraging trip duration, body mass and reproductive success, we also investigated food-related stress (via plasma corticosterone concentration), nutritional state (via metabolite levels) and the use of penguins' habitat (via blood isotopic values). 3. Body mass and reproductive success remained unchanged but foraging trips were shorter when sea-ice retreated earlier. Constant plasma corticosterone concentrations indicated that none of the feeding conditions resulted in a food-related stress. However metabolite levels were lower when sea-ice retreated early, suggesting that the foraging performance and the quality/quantity of food differed. Indeed isotopic ratios indicated that coastal prey like fish contributed more to the penguins' diet when sea-ice retreated prematurely. 4. The early sea-ice retreat was related to higher chlorophyll concentrations, known to favour krill recruitment. Paradoxically, this was not associated to a higher krill contribution in the penguins' diet. We propose that a shift in the phytoplankton quality (rather than quantity), affecting krill recruitment, forced penguins to switch to more available prey like coastal fish. 5. In some Antarctic regions, sea-ice is retreating earlier and earlier. In the present study, even though the timing of sea-ice retreat and the consecutive ocean productivity differed drastically between the 2 years, Adelie penguins were not severely affected because they were able to adjust their at-sea behaviour and thus maintained their body condition and reproductive success unchanged. 6. This suggests that the timing of sea-ice retreat does not represent an important threat to populations of Adelie penguins at least as long as alternative resources are still available and other environmental parameters like winter sea-ice extent are not dramatically altered.

(Tables 1,2) Site characteristics and phenotypic attributes of King Penguins (Aptenodytes patagonicus) on Possession Island

In colonial species, it is often assumed that locations in the center of the colony are of highest quality and provide highest breeding success. We tested this prediction, known as the "central-periphery model," in a King Penguin colony in the subantarctic Crozet Archipelago. Breeding activity and survival of 150 penguins, fitted with transponder tags, were monitored over an entire breeding season. Among these 150 birds, 50 bred on the slope at the upper periphery of the colony, where the rates of predation and parasitism by ticks were high. Fifty birds bred in the center of the colony, where rates of predation and tick parasitism were low, and 50 bred at the lower end of the colony, where the rate of tick parasitism was low but predation and flooding were important risks. We predicted that the center of the colony should provide the safest breeding place and consequently be characterized by the highest breeding success and be used by the highest-quality individuals. Yet we found that penguins breeding in the center of the colony had the same breeding success as those at both peripheral locations. In addition, penguins breeding on the upper slope had a higher survival rate than penguins breeding at the center or bottom of the slope and were likely of higher quality. Our study does not support the central-periphery model and emphasizes the complexity behind the relationships among breeding site, breeding success, and individual quality.

(Table 1) Comparison of dive duration and speed of Adélie penguins (Pygoscelis adeliae) equipped with internal and external devices

Bio-logging studies suffer from the lack of real controls. However, it is still possible to compare indirect parameters between control and equipped animals to assess the level of global disturbance due to instrumentation. In addition, it is also possible to compare the behaviour of free-ranging animals between individuals equipped with different techniques or instruments to determine the less deleterious approach. We instrumented Adelie Penguins (Pygoscelis adeliae) with internal or external time-depth recorders and monitored them in parallel with a control group during the first foraging trip following instrumentation. Foraging trip duration was significantly longer in the internally-equipped group. This difference was due to a larger number of dives, reflecting a lower foraging ability or a higher food demand, and longer periods of recovery at the surface. These longer recovery periods were likely to be due to a reduced efficiency to ventilate at the surface, probably because the implanted devices pressurised adjacent organs such as air sacs. Moreover, descent and ascent rates were slightly lower in externally-equipped penguins, presumably because external instrumentation increased the bird drag. Looking at our results, implantation appears more disadvantageous - at least for short-term deployment - than external equipment in Adelie Penguins, while this method has been described to induce no negative effects in long-term studies. This underlines the need to control for potential effects due to methodological aspects in any study using data loggers in free-ranging animals, to minimise disturbance and collect reliable data.

(Table 1) Characteristics of breeding male Adélie penguins (Pygoscelis adeliae) treated with varying degrees of corticosterone, Adélie land, Antarctica

Breeding individuals enter an emergency life-history stage when their body reserves reach a minimum threshold. Consequently, they redirect current activity toward survival, leading to egg abandonment in birds. Corticosterone (CORT) is known to promote this stage. How and to what extent CORT triggers egg abandonment when breeding is associated with prolonged fasting, however, requires further investigation. We manipulated free-living male Adelie penguins with CORT-pellets before their laying period. We then examined their behavioral response with respect to nest abandonment in parallel with their prolactin levels (regulating parental care), and the subsequent effects of treatment on breeding success in relieved birds. Exogenous CORT triggered nest abandonment in 60% of the treated penguins -14 days after treatment and induced a concomitant decline in prolactin levels. Interestingly, prolactin levels in treated penguins that did not abandon their nest were higher at the point of implantation and also after being relieved by females, when compared with abandoning penguins. Among successful birds, the treatment did not affect the number of chicks, nor the brood mass. Our results show the involvement of CORT in the decision-making process regarding egg abandonment in Adelie penguins when incubation is associated with a natural long fast. However, we suggest that CORT alone is not sufficient to trigger nest abandonment but that 1) prolactin levels need to reach a low threshold value, and 2) a rise in proteolysis (i.e. utilization of protein as main energy substrate) seems also to be required.

Antarctic penguins; a study of their social habits,

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Sexual behavior in penguins

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