At-sea associations in foraging little penguins

Prey distribution, patch size, and the presence of conspecifics are important factors influencing a predator's feeding tactics, including the decision to feed individually or socially. Little is known about group behaviour in seabirds as they spend most of their lives in the marine environment where it is difficult to observe their foraging activities. In this study, we report on at-sea foraging associations of little penguins (Eudyptula minor) during the breeding season. Individuals could be categorised as (1) not associating; (2) associating when departing from and/or returning to the colony; or (3) at sea when travelling, diving or performing synchronised dives. Out of 84 separate foraging tracks, 58 (69.0%) involved associations with conspecifics. Furthermore, in a total of 39 (46.4%), individuals were found to dive during association and in 32 (38.1%), individuals were found to exhibit synchronous diving. These behaviours suggest little penguins forage in groups, could synchronise their underwater movements and potentially cooperate to concentrate their small schooling prey.

Influence of environmental conditions on foraging behaviour and its consequences on reproductive performance in little penguins

During the breeding season, seabirds are central place foragers and have to adapt their foraging behaviour in response to environmental variation to maximize efficiency and reproductive output. Due to its small size and swimming mode of transport, the little penguin (Eudyptula minor) is expected to be greatly susceptible to such fluctuations. The links between local-, meso- and macro-scale environmental conditions and inter-annual variation in foraging behaviour and reproductive performance of little penguins were investigated during three consecutive breeding seasons at two colonies in south-eastern Australia marked by contrasting oceanographic conditions. At a local scale, foraging effort was correlated positively with wind direction and negatively with wave height. At a regional scale, foraging effort of individuals from both colonies was negatively correlated with higher sea surface temperature (SST) off the Bonney Coast in the previous Austral summer, suggesting a weaker Bonney Upwelling event and a cascade of effects throughout the Bass Strait region. At a larger scale, the El Niño Southern Oscillation was also found to correlate with foraging behaviour, with lower foraging effort being observed during La Niña event. Although individuals increased their foraging effort during years with poorer conditions, they were not able to maintain high breeding success. In addition, peak egg-laying was found to coincide with a decrease in local SST and a peak of sea surface chlorophyll-a concentration. In conclusion, these results highlight how different environmental conditions could influence foraging behaviour and ultimately reproductive success of little penguins. It also showed that under certain circumstances, these individual strategies were not sufficient to cope with environmental variability.

(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.

(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.

The influence of animal mobility on the assumption of uniform distances in aerial line-transect surveys

Line-transect distance sampling is a widely used method for estimating animal density from aerial surveys. Analysis of line-transect distance data usually relies on a requirement that the statistical distribution of distances of animal groups from the transect line is uniform. We show that this requirement is satisfied by the survey design if all other assumptions of distance sampling hold, but it can be violated by consistent survey problems such as responsive movement of the animals towards or away from the observer. We hypothesise that problems with the uniform requirement are unlikely to be encountered for immobile taxa, but might become substantial for species of high mobility. We test evidence for non-uniformity using double-observer distance data from two aerial surveys of five species with a spectrum of mobility capabilities and tendencies. No clear evidence against uniformity was found for crabeater seals or emperor penguins on the pack-ice in East Antarctica, while minor non-uniformity consistent with responsive movement up to 30 m was found for Adelie penguins. Strong evidence of either non-uniformity or a failure of the capture-recapture validating method was found for eastern grey kangaroos and red kangaroos in Queensland.

Carbon fluxes and pelagic ecosystem dynamics near two western Antarctic Peninsula Adélie penguin colonies: an inverse model approach

An inverse food-web model for the western Antarctic Peninsula (WAP) pelagic food web was constrained with data from Palmer Long Term Ecological Research (PAL-LTER) project annual austral summer sampling cruises. Model solutions were generated for 2 regions with Adelie penguin Pygoscelis adeliae colonies presenting different population trends (a northern and a southern colony) for a 12 yr period (1995-2006). Counter to the standard paradigm, comparisons of carbon flow through bacteria, microzooplankton, and krill showed that the diatom-krill-top predator food chain is not the dominant pathway for organic carbon exchanges. The food web is more complex, including significant contributions by microzooplankton and the microbial loop. Using both inverse model results and network indices, it appears that in the northern WAP the food web is dominated by the microbial food web, with a temporal trend toward its increasing importance. The dominant pathway for the southern WAP food web varies from year to year, with no detectable temporal trend toward dominance of microzooplankton versus krill. In addition, sensitivity analyses indicated that the northern colony of Adelie penguins, whose population size has been declining over the past 35 yr, appears to have sufficient krill during summer to sustain its basic metabolic needs and rear chicks, suggesting the importance of other processes in regulating the Adelie population decline.

Influence of sea-ice structure on minke whale and other krill-predator distribution

This thesis investigated the extent to which sea-ice structure and complexity could be measured to identify suitable feeding habitat for krill-eating predators such as the Antarctic minke whale (Balaenoptera bonaerensis). The results of this study suggest that the distribution and movement of minke whales is limited by the structure of the sea ice and that the sea ice influences minke whales' habitat use more than their overall distribution. Similar relationships were found for other krill predators such as Adelie penguins and crabeater seals.

Tab.1: Tabulation of pinguin population within the Antarctic Peninsula region

Six species of penguins breed on the Antarctic continent, the Antarctic Peninsula, the South Shetland and South Orkney Islands. Their breeding populations within the Antarctic Peninsula, and the South Orkney and South Shetland Is., and estimates of global populations are given. Typical breeding seasons are also presented, but it must be noted that these will vary inter-annually and intra-annually under the influence of factors such as sea-ice extent and ENSO (interannual) and the location of each breeding colony (southerly localities will be later than northerly localities, as their breeding season is "compressed" within the shorter summer). Their foraging strategies (categorized as near-shore or offshore) and typical durations of foraging trips are also tabulated. As with breeding season events, foraging behaviour will vary intra-seasonally and inter-seasonally (in terms of dive duration, dive depth, foraging location, etc). The distribution of known penguin breeding colonies is circum-continental, with Emperor and Adelie penguins predominant on approximately 75 % of the coast, with two major concentrations in the Ross Sea and in Prydz Bay. The third concentration is in the Antarctic Peninsula region, where some of the largest penguin colonies are present. All six species breed within the area (predominantly Chinstrap Penguins), and the Peninsula region has a greater diversity than the remainder ofthe Antarctic with respect to penguins. The distribution at sea of nonbreeding penguins is less cIear. Non-breeding individuals of all six species move throughout the Southern Ocean, and in many cases, to areas well north of the winter pack-ice zone. However, it is not possible to estimate densities of penguins at sea as there are no estimates of non-breeding penguin populations the extent of their travels.

(Table 1) Concentrations of organic pollutants in soil samples from Adélie penguin (Pygoscelis adeliae) colonies and reference sites, Hop Island, Antarctica

Although long-range atmospheric transport has been described as the predominant mechanism for exposing polar regions to persistent organic pollutants (POPs), recent studies have suggested that bird activity can also contribute substantially to contaminant levels in some environments. However, because the species so far reported have all been migratory, it has not been demonstrated conclusively whether locally elevated contamination represents transport from lower latitudes by the migrating birds or, alternatively, redistribution and concentration of contaminants that were already present in the high-latitude environments. The present study demonstrates, for the first time, that several POPs are present in elevated concentrations in an environment frequented by a non-migratory species (Adelie penguins) that spends its entire life in the Antarctic. Levels of POPs, such as p,p'-DDE, hexachlorobenzene (HCB), chlordanes (CHLs) and polychlorinated biphenyls (PCBs), were 10 to 100-fold higher in soil samples from penguin colonies than from reference areas. This significant difference is likely related to local penguin activity, such as a higher abundance of guano and the presence of bird carcasses. This hypothesis is also supported by a higher percentage of persistent congeners (PCB 99, 118, 138 and 153) in the soil from the colonies compared to the reference areas. This profile of PCB congeners closely matched profiles seen in penguin eggs or penguin blood.

As aves antárticas estão livres de hemoparasitos? Um estudo de caso de pinguins (Pygoscelis spp.) e de skuas (Catharacta spp.) antárticos da Baía do Almirantado, Ilha Rei George, Antártica.

O parasitismo é uma importante força seletiva em populações, assim como a competição e a predação. Os parasitos sanguíneos podem afetar a coloração da plumagem, a seleção sexual e o sucesso reprodutivo em aves. As aves da região Antártica têm sido mencionadas na literatura como livres de hemoparasitos. A Baía do Almirantado, na Ilha Rei George, Península Antártica, é a maior Baía da região, abrigando diferentes espécies de aves durante o período reprodutivo. Dentre elas, estão duas espécies de skuas, as mais frequentes da Antártica, skua-sub-antártica (Catharacta lonnbergi) e skua-polar-do-sul (C. maccormicki) e três espécies de pinguins, pinguim-antártico (Pygoscelis antarctica), pinguim-papua (P. papua) e pinguim-de-adélia (P. adeliae). Skuas e pinguins são aves que se dispersam durante o inverno austral, podendo ser potenciais reservatórios e transmissores de parasitos, embora resultados negativos de hemoparasitos tenham sido encontrados para diversas outras aves marinhas e também para a região Antártica. O objetivo do presente trabalho foi investigar a presença de hemoparasitos em pinguins e skuas antárticos na Baía do Almirantado. Amostras de lâminas de esfregaço sanguíneo e de sangue para análises moleculares de pesquisa de Plasmodium/Haemoproteus foram coletadas em dois períodos reprodutivos, de dezembro de 2010 a março de 2011 e de dezembro de 2011 a fevereiro de 2012. Um total de 185 amostras de aves foram coletadas, incluindo 120 pinguins e 65 skuas. Skuas foram tiveram resultados negativos para hemoparasitos. As três espécies de pinguins foram positivas para Plasmodium/Haemoproteus , via técnica molecular, incluindo dois P. papua,dois P. antarctica etrês P. adeliae. Apenas um indivíduo confirmado positivo pela técnica molecular, pertencente a P. papua, foi positivo utilizando a técnica de esfregaço sanguíneo, com diagnóstico de Plasmodium sp. Não houve diferença significativa entre indivíduos machos e fêmeas das espécies parasitadas, assim como entre adultos e filhotes. As aves parasitadas (n=7), foram categorizadas abaixo do peso (n=5) e acima do peso (n=2). O presente estudo é o primeiro a relatar hemoparasitos na região Antártica e também é o primeiro registro de presença de hemoprotozoários para as três espécies de pinguins analisadas. A ausência de hemoparasitos em aves antárticas tem sido justificada pela ausência de potenciais vetores na região. Portanto, é possível que os pinguins parasitados tenham adquirido a infecção durante a dispersão por ocasião do inverno austral. No entanto, skuas antárticas também são aves migratórias, que podem atingir regiões com potenciais vetores reconhecidos, mas nunca foram diagnosticadas com hemoparasitos, o que foi reforçado pelos resultados negativos do presente estudo. Nesse caso, acredita-se que skuas, podem ter um sistema imune competente ou que a ausência de hemoparasitos nessas aves seja justificada por confinamentos filogenéticos entre parasito-hospedeiro. Entretanto, pouco se sabe sobre a existência de vetores na Antártica, rotas migratórias das aves da região e especificidade parasito-hospedeiro. Os resultados inéditos encontrados no presente estudo devem, portanto, servir como ponto de partida para o entendimento das interações parasito-hospedeiro, de forma a contribuir para a preservação do ambiente antártico.