Although penguins are key marine predators from the Southern Ocean, their migratory behaviour during the inter-nesting period remains widely unknown. Here, we report for the first time, to our knowledge, the winter foraging movements and feeding habits of a penguin species by using geolocation sensors fitted on penguins with a new attachment method. We focused on the macaroni penguin Eudyptes chrysolophus at Kerguelen, the single largest consumer of marine prey among all seabirds. Overall, macaroni penguins performed very long winter trips, remaining at sea during approximately six months within the limits of the Southern Ocean. They departed from Kerguelen in an eastward direction and distributed widely, over more than 3.10 6 km 2 . The penguins spent most of their time in a previously unrecognized foraging area, i.e. a narrow latitudinal band (47–49° S) within the central Indian Ocean (70–110° E), corresponding oceanographically to the Polar Frontal Zone. There, their blood isotopic niche indicated that macaroni penguins preyed mainly upon crustaceans, but not on Antarctic krill Euphausia superba , which does not occur at these northern latitudes. Such winter information is a crucial step for a better integrative approach for the conservation of this species whose world population is known to be declining.

Ecological niche theory predicts segregation mechanisms that mitigate potential competition between closely related organisms. However, little is known outside the breeding season, when central-place foraging animals may move on larger scales. This study tested for segregation mechanisms within the same 2007 inter-breeding period on three neighboring populations of avian predators from the southern Indian Ocean: Eastern Rockhopper Penguins Eudyptes filholi from Crozet and Kerguelen and Northern Rockhopper Penguins E. moseleyi from Amsterdam. Using state-of-the-art geolocation tracking and stable isotope analysis techniques, we quantified and compared the ecological niches in time, space, and diet. The three populations showed large-scale movements over deep oceanic waters near the Subantarctic Front, with generally little individual variation. The two neighboring populations of Eastern Rockhopper Penguins showed strikingly distinct distribution in space, while foraging in similar habitats and at the same trophic level (crustacean-eaters). In contrast, Northern Rockhoppers showed marked spatial overlap with birds of the sibling Eastern species, but their temporal delay of two months enabled them to effectively avoid significant overlap. Our results highlight parsimonious mechanisms of resource partitioning operating at the population level that may explain how animals from neighboring localities can coexist during the nonbreeding period.

How top avian predators are distributed at sea during the inter-breeding period remains poorly known. Here we focused on the pelagic seabird that is the single greatest avian consumer of marine resources: the macaroni penguin. Our aims were (1) to investigate where these penguins are distributed whilst at sea during winter, and (2) to determine how their distribution and trophic ecology varies in 1 colony between successive years and between 2 colonies from neighbouring localities at the same time. We surveyed a total of 30 penguins from Crozet and Kerguelen Islands (southern Indian Ocean). Penguins from Kerguelen were tracked in 2006 and 2007, and those from Crozet in 2007. Habitat use was investigated using miniaturised light-based geolocators, and trophic ecology by the use of stable isotope analyses. Results showed large-scale patterns of distribution over deep oceanic waters of the Polar Frontal Zone characterised by a temperature of 3.0 to 3.5°C during July. Birds from Kerguelen had a consistent inter-annual winter distribution at the population level. Birds tracked from Crozet were distributed in distinct areas, separate from the birds from Kerguelen. Trophic ecology (blood δ 15 N value) was very similar between years and colonies, and indicated that these penguins preyed upon low trophic level prey, most probably swarming crustaceans, at the end of winter. We conclude that inter-annual fidelity in winter habitat provides evidence of highly favourable and predictable foraging areas, while the distinct inter-colony distributions suggest very heterogeneous distribution of profitable feeding areas for this species in the southern Indian Ocean.

Infectious diseases may be particularly critical for the conservation of endangered species. A striking example is the recurrent outbreaks that have been occurring in seabirds on Amsterdam Island for the past 30 years, threatening populations of three Endangered seabird species and of the endemic, Critically Endangered Amsterdam albatross Diomedea amsterdamensis. The bacteria Pasteurella multocida (avian cholera causative agent), and to a lesser extent Erysipelothrix rhusiopathiae (erysipelas causative agent), were both suspected to be responsible for these epidemics. Despite this critical situation, demographic trends were not available for these threatened populations, and the occurrence and characterization of potential causative agents of epizootics remain poorly known. The aims of the current study were to (i) provide an update of population trends for four threatened seabird species monitored on Amsterdam Island, (ii) assess the occurrence of P. multocida, and E. rhusiopathiae in live birds from five species, (iii) search for other infectious agents in these samples and, (iv) isolate and genotype the causative agent(s) of epizooties from dead birds. Our study shows that the demographic situation has worsened substantially in three seabird species during the past decade, with extremely low reproductive success and declining populations for Indian yellow-nosed albatrosses Thalassarche carteri, sooty albatrosses Phoebetria fusca, and northern rockhopper penguins Eudyptes moseleyi. Pasteurella multocida or E. rhusiopathiae were detected by PCR in live birds of all five investigated species, while results were negative for eight additional infectious agents. A single strain of P. multocida was repeatedly cultured from dead birds, while no E. rhusiopathiae could be isolated. These results highlight the significance of P. multocida in this particular eco-epidemiological system as the main agent responsible for epizootics. The study stresses the urgent need to implement mitigation measures to alter the course of avian cholera outbreaks threatening the persistence of seabird populations on Amsterdam Island.

Abstract:The juvenile phase is poorly known in Antarctic seabirds, despite being a critical period for individual survival. To better understand the ecology of young Antarctic seabirds, we surveyed for the first time the three-dimensional habitat use of six juvenile emperor penguins during their post-natal dispersal from Terre Adélie, using bio-telemetric tags. The tags transmitted location and activity data for nearly 100 days on average. One individual was followed during eight months and covered 7000 km, which represents the longest continuous individual survey for the species. Studied individuals first dispersed away from Antarctica, up to 54.78S and 1250 km north of the pack-ice edge, in the Polar Frontal Zone. This highlighted a much looser association with sea ice and a greater at-sea range compared to previous knowledge on breeding adults. Juvenile penguins then moved southwards close to the extending pack-ice during autumn and winter. Over the survey duration, juveniles showed a contrasting use of marine habitats, with less mobility, less time underwater, and shallower dives (generally not over 50-100 m) in the pack ice, versus greater distances travelled, more time spent underwater, especially deeper than 100 m (up to 250-300 m) in open water. We discuss hypotheses which could explain the northward exodus of juvenile emperor penguins across contrasting habitats.

For land-breeding marine organisms such as seabirds, knowledge about their habitat use has mainly been gained through studies of breeding individuals that are constrained to return frequently to their breeding grounds. In this study we set out to measure whether: a) habitat selection in the non-breeding period predicts habitat selection in the breeding period, and b) whether breeding individuals concentrated their activity on the closest suitable habitats. Macaroni Eudyptes chrysolophus and gentoo Pygoscelis papua penguins, two marine predators with contrasting foraging strategies, were tracked from the Iles Kerguelen and their habitat selection investigated through Mahalanobis distances factorial analysis. This study presents the first data about gentoo penguins' juvenile dispersal. For both species, results showed 6.9 times larger maximum ranges and up to 12.2 times greater distances travelled during the non-breeding period. Habitat suitability maps suggested both species made similar environmental selections whatever the period. Macaroni penguins targeted pelagic areas beyond the shelf break while gentoo penguins always remained over the shelf. We consider the ecological significance of larger scale movements made outside the breeding period and suggest that this non-breeding period is of particular interest when attempting to understand an animal's habitat selection.

The development of biologging techniques has been instrumental in studying the behaviour of wild animals and interpreting it with respect to the bio-physical features of their habitat. Light-based geolocation currently appears to be the only technique suitable for the study of farranging small species, particularly marine species, over long periods, but it provides locations with low precision. In this study, we sought to improve the exploitation of these data. Specifically, the goals were to (1) correct rather than reject estimates, especially during equinox periods, (2) perform reproducible time-saving routine analyses, and (3) determine the spatial likelihood of the estimations to adapt inferences on habitat use by the population. We therefore applied an existing data-processing method based on spatial template fitting, using Markov Chain Monte Carlo and state-space modelling (Kalman filter) improved by a facultative sea surface temperature-matching procedure and a land mask. The main functions used for geolocation are grouped under the R package 'TripEstimation', freely available online. We focused on a typical example of animal movement that at present can only be ethically obtained from light-based geolocation. The method made it possible to estimate realistic positions during equinox periods and to routinely process the 12 available datasets. We thus obtained the most probable location for each sunrise/sunset as well as the posterior distribution around each estimated location, allowing an intuitive habitat use investigation at a scale of 100 km. This paper describes the method used and provides the complete and comprehensively annotated commands required for its use.

Global change is contributing to unprecedented expansions of infectious diseases in wildlife. Recurrent avian cholera outbreaks are causing dramatic chick mortality and population decline in endangered albatross colonies on Amsterdam Island, a critical seabird breeding ground in the Southern Indian Ocean. We manufactured a killed vaccine using a Pasteurella multocida strain isolated from a dead albatross in the field.We used this same bacterial strain to establish a serological assay allowing the monitoring of antibody levels following bird vaccination. Using this vaccine on chicks 2 weeks posthatching caused 100% seroconversion and reduced the death risk by a factor exceeding 2.5, raising fledging probability from 14% to 46%. These results suggest that using a specifically tailored vaccine could be a key tool to effectively protect endangered seabirds from disease outbreaks threatening them with extinction.