Colonial breeding is widespread among animals. Some, such as eusocial insects, may use agonistic behavior to partition available foraging habitat into mutually exclusive territories; others, such as breeding seabirds, do not. We found that northern gannets, satellite-tracked from twelve neighboring colonies, nonetheless forage in largely mutually exclusive areas and that these colony-specific home ranges are determined by densitydependent competition. This segregation may be enhanced by individual-level public information transfer, leading to cultural evolution and divergence among colonies.Main Text: Colonial animals are constrained by their colony locations, which are ultimately limited by resource availability (1). However, within species, potential colony home ranges often overlap, implying competition among colonies may also be limiting (2). In eusocial central-place foragers the spatial effects of direct competition among colonies are well understood (2). In contrast, the spatial influences of indirect competition and information transfer on non-territorial species (e.g. seals, swallows and seabirds), where levels of relatedness are much lower, remain conjectural. For example, the hinterland model (3) predicts that breeding seabirds segregate along colonial lines, because of inequalities in travel costs from each colony. Predicted home ranges therefore comprise Voronoi polygons (Fig. 1A), as seen in some territorial animals (2). Food availability is assumed to be proportional to polygon area, limiting colony size. An alternative model proposes that density-dependent competition among colony members is limiting (4). As colonies grow, local prey depletion or disturbance requires birds to travel further to provision their young. However, this model ('Ashmole's halo') does not consider interactions among colonies and tacitly assumes that adjacent colonies' home ranges overlap (5).Indirect evidence exists to support both models (3,6,7) and recent tracking studies suggest that seabirds and pinnipeds segregate along colonial lines (8-12). However, these studies proved inconclusive on the causes and ubiquity of segregation, largely because few colonies were sampled or tracking resolution was low. Here we use high resolution satellite-tracks of the foraging movements of 184 chick-rearing northern gannets Morus bassanus (hereafter gannets) from 12 of the 26 colonies fringing the British Isles (median 17 birds/colony), representing ~80% of the area's breeding population (Fig. 1A, Table S1), to test whether among-colony segregation occurs in a model colonial non-territorial central-place forager. We then use population-and individual-level models to explore potential mechanisms underlying spatial segregation.Gannets are wide-ranging (max. foraging range ~700 km) pelagic seabirds that forage in patches of enhanced production, primarily on shoaling, mesotrophic fish and to a lesser extent fisheries discards (13)(14)(15). In almost all cases we tracked birds from adjacent colonies simultaneously (16). Individua...

Abstract. Extreme climatic conditions and their ecological impacts are currently emerging as critical features of climate change. We studied extreme sea ice condition (ESIC) and found it impacts both life-history traits and population dynamics of an Antarctic seabird well beyond ordinary variability.The Southern Fulmar (Fulmarus glacialoides) is an ice-dependent seabird, and individuals forage near the ice edge. During an extreme unfavorable year (when sea ice area is reduced and distance between ice edge and colony is high), observed foraging trips were greater in distance and duration. As a result, adults brought less food to their chicks, which fledged in the poorest body condition. During such unfavorable years, breeding success was extremely low and population growth rate (k) was greatly reduced. The opposite pattern occurred during extreme favorable years.Previous breeding status had a strong influence on life-history traits and population dynamics, and their responses to extreme conditions. Successful breeders had a higher chance of breeding and raising their chick successfully during the following breeding season as compared to other breeding stages, regardless of environmental conditions. Consequently, they coped better with unfavorable ESIC. The effect of change in successful breeder vital rates on k was greater than for other stages' vital rates, except for pre-breeder recruitment probabilities, which most affected k.For environments characterized by ordinary sea ice conditions, interindividual differences were more likely to persist over the life of individuals and randomness in individual pathways was low, suggesting individual heterogeneity in vital rates arising from innate or acquired phenotypic traits. Additionally, unfavorable ESIC tended to exacerbate individual differences in intrinsic quality, expressed through differences in reproductive status.We discuss the strong effects of ESIC on Southern Fulmar life-history traits in an evolutionary context. ESICs strongly affect fitness components and act as potentially important agents of natural selection of life histories related to intrinsic quality and intermittent breeding. In addition, recruitment is a highly plastic trait that, if heritable, could have a critical role in evolution of life histories. Finally, we find that changes in the frequency of extreme events may strongly impact persistence of Southern Fulmar populations.

Seasonal long-distance migration is likely to be experienced in a contrasted manner by juvenile, immature and adult birds, leading to variations in migratory routes, timing and behaviour. We provide the first analysis of late summer movements and autumn migration in these three life stages, which were tracked concurrently using satellite tags, geolocators or GPS recorders in a long-ranging migratory seabird, the Scopoli’s shearwater (formerly named Cory’s shearwater, Calonectris diomedea ) breeding on two French Mediterranean islands. During the late breeding season, immatures foraged around their colony like breeding adults, but they were the only group showing potential prospecting movements around non-natal colonies. Global migration routes were broadly comparable between the two populations and the three life stages, with all individuals heading towards the Atlantic Ocean through the strait of Gibraltar and travelling along the West African coast, up to 8000 km from their colony. However, detailed comparison of timing, trajectory and oceanographic conditions experienced by the birds revealed remarkable age-related differences. Compared to adults and immatures, juveniles made a longer stop-over in the Balearic Sea (10 days vs 4 days in average), showed lower synchrony in crossing the Gibraltar strait, had more sinuous pathways and covered longer daily distances (240 km.d-1 vs 170 km.d-1). Analysis of oceanographic habitats along migratory routes revealed funnelling selection of habitat towards coastal and more productive waters with increasing age. Younger birds may have reduced navigational ability and learn progressively fine-scale migration routes towards the more profitable travelling and wintering areas. Our study demonstrates the importance of tracking long-lived species through the stages, to better understand migratory behavior and assess differential exposure to at-sea threats. Shared distribution between life stages and populations make Scopoli’s shearwaters particularly vulnerable to extreme mortality events in autumn and winter. Such knowledge is key for the conservation of critical marine habitats.