Demographic profiles and environmental drivers of variation relate to individual breeding state in a long-lived trans-oceanic migratory seabird, the Manx shearwater.

Understanding the points in a species breeding cycle when they are most vulnerable to environmental fluctuations is key to understanding interannual demography and guiding effective conservation and management. Seabirds represent one of the most threatened groups of birds in the world, and climate change and severe weather is a prominent and increasing threat to this group. We used a multi-state capture-recapture model to examine how the demographic rates of a long-lived trans-oceanic migrant seabird, the Manx shearwater Puffinus puffinus, are influenced by environmental conditions experienced at different stages of the annual breeding cycle and whether these relationships vary with an individual's breeding state in the previous year (i.e., successful breeder, failed breeder and non-breeder). Our results imply that populations of Manx shearwaters are comprised of individuals with different demographic profiles, whereby more successful reproduction is associated with higher rates of survival and breeding propensity. However, we found that all birds experienced the same negative relationship between rates of survival and wind force during the breeding season, indicating a cost of reproduction (or central place constraint for non-breeders) during years with severe weather conditions. We also found that environmental effects differentially influence the breeding propensity of individuals in different breeding states. This suggests individual spatio-temporal variation in habitat use during the annual cycle, such that climate change could alter the frequency that individuals with different demographic profiles breed thereby driving a complex and less predictable population response. More broadly, our study highlights the importance of considering individual-level factors when examining population demography and predicting how species may respond to climate change.

Drivers and fitness consequences of dispersive migration in a pelagic seabird.

Animals can be flexible in their migration strategies, using several wintering sites or a variety of routes. The mechanisms promoting the development of these migratory patterns and their potential fitness consequences are poorly understood. Here, we address these questions by tracking the dispersive migration of a pelagic seabird, the Atlantic puffin Fratercula arctica, using over 100 complete migration tracks collected over 7 years, including repeated tracks of individuals for up to 6 consecutive years. Because puffins have high flight costs, dispersion may generate important variation in costs of migration. We investigate differences in activity budgets and energy expenditure between different strategies. We find that puffins visit a range of overwintering destinations, resulting in a diversity of migratory routes differing in energy expenditures; however, they show interindividual similarity in the timings and location of major movements. We consider 3 hypothetical mechanisms that could generate this pattern: 1) random dispersion; 2) sex segregation; and 3) intraspecific competition or differences in individual quality. First, we dismiss random dispersion because individuals show strong route fidelity between years. Second, we find that sex differences contribute to, but do not account fully for, the migratory variation observed. Third, we find significant differences in breeding success between overwintering destinations, which, together with differences in foraging levels between routes, suggest that birds of different quality may visit different destinations. Taken together, our results show that dispersive migration is a complex phenomenon that can be driven by multiple factors simultaneously and can shape a population's fitness landscape.

Foraging flexibility and search patterns are unlinked during breeding in a free-ranging seabird.

In order to maximize foraging efficiency in a varying environment, predators are expected to optimize their search strategy. Environmental conditions are one important factor affecting these movement patterns, but variations in breeding constraints (self-feeding vs. feeding young and self-feeding) during different breeding stages (incubation vs. chick-rearing) are often overlooked, so that the mechanisms responsible for such behavioral shifts are still unknown. Here, to test how search patterns are affected at different breeding stages and to explore the proximate causes of these variations, we deployed data loggers to record both position (global positioning system) and dive activity (time-depth recorders) of a colonial breeding seabird, the razorbill Alca torda. Over a period of 3 years, our recordings of 56 foraging trips from 18 breeders show that while there is no evidence for individual route fidelity, razorbills exhibit higher foraging flexibility during incubation than during chick rearing, when foraging becomes more focused on an area of high primary productivity. We further show that this behavioral shift is not due to a shift in search patterns, as reorientations during foraging are independent of breeding stage. Our results suggest that foraging flexibility and search patterns are unlinked, perhaps because birds can read cues from their environment, including conspecifics, to optimize their foraging efficiency.

A dispersive migration in the Atlantic Puffin and its implications for migratory navigation.

Navigational control of avian migration is understood, largely from the study of terrestrial birds, to depend on either genetically or culturally inherited information. By tracking the individual migrations of Atlantic Puffins, Fratercula arctica, in successive years using geolocators, we describe migratory behaviour in a pelagic seabird that is apparently incompatible with this view. Puffins do not migrate to a single overwintering area, but follow a dispersive pattern of movements changing through the non-breeding period, showing great variability in travel distances and directions. Despite this within-population variability, individuals show remarkable consistency in their own migratory routes among years. This combination of complex population dispersion and individual route fidelity cannot easily be accounted for in terms of genetic inheritance of compass instructions, or cultural inheritance of traditional routes. We suggest that a mechanism of individual exploration and acquired navigational memory may provide the dominant control over Puffin migration, and potentially some other pelagic seabirds, despite the apparently featureless nature of the ocean.