Understanding how geographic range limits are shaped is a central and challenging question in ecology that has become particularly critical in the context of global environmental changes. While such limits are often studied for resident or breeding species, range limits are equally important for migratory species in winter when population regulation may occur due to limited resources in the non-breeding season. A central hypothesis in several theories for range limitations is that the density, fitness and performance of individuals decrease towards the edge of the range as organisms become maladapted when approaching the limit of their environmental tolerance (‘Abundant-centre Hypothesis’). Energy is a critical resource, especially in winter when environmental conditions deteriorate, and this hypothesis predicts that high energy expenditure (low performance) at the range limit would lead to rapidly dwindling body mass and reduced fitness. We investigated this hypothesis in an Arctic-breeding seabird wintering in the North-Atlantic, the black-legged kittiwake (Rissa tridactyla). From 2008 to 2019, we tracked 117 adult kittiwakes (n=176 tracks) with geolocation devices and saltwater immersion sensors to estimate the migratory strategies, time-activity budget and energy expenditure of individuals during winter, and estimated their reproductive success after their return to the colony during summer. Population density was indeed higher towards the center of the range. However, contrary to the predictions, the energy expenditure of individuals was higher at the centre of the range and decreased towards the edge. In contrast, there were no spatial differences in the reproductive success of individuals wintering at the centre versus at the edge of their range. We conclude that performance and fitness did not increase towards the centre of the wintering range, implying that although resource acquisition was likely higher at the abundant centre, energy expenditure was also higher, so that individual fitness was constant across the winter range.

Understanding how geographic range limits are shaped is a central and challenging question in ecology that has become particularly critical in the context of global environmental changes. A central hypothesis in several theories for range limitations is that the density, fitness and performance of individuals decrease towards the edge of the range as organisms become maladapted when approaching the limit of their environmental tolerance (‘Abundant-centre Hypothesis’). Energy is a critical resource, especially in winter when environmental conditions deteriorate, and this hypothesis predicts that high energy expenditure (low performance) at the range limit would lead to rapidly dwindling body mass and reduced fitness. We investigated this hypothesis in an Arctic breeding seabird wintering in the North-Atlantic, the black-legged kittiwake (Rissa tridactyla). From 2008 to 2019, we tracked 118 adult kittiwakes (n= 178 tracks) with geolocation devices and saltwater immersion sensors to estimate the time-activity budget and energy expenditure of individuals during winter, and estimated their reproductive success after their return to the colony during summer. Density was indeed higher towards the center of the range. However, contrary to the predictions, the energy expenditure of individuals was higher at the centre of the range and decreased towards the edge. In contrast, there were no spatial differences in the reproductive success of individuals wintering at the centre versus at the edge of their range. We conclude that performance and fitness did not increase towards the centre of the range, implying that although resource acquisition was likely higher at the abundant centre, energy expenditure was also higher, so that individual fitness was constant across the range.

Animal-borne telemetry devices provide essential insights into the life-history strategies of far-ranging species and allow us to understand how they interact with their environment. Many species in the seabird family Alcidae undergo a synchronous moult of all primary flight feathers during the non-breeding season, making them flightless and more susceptible to environmental stressors, including severe storms and prey shortages. However, the timing and location of moult remains largely unknown, with most information coming from studies on birds killed by storms or shot at sea. Using light-level geolocators with saltwater immersion loggers, we develop a method for determining flightless periods in the context of the annual cycle. Four Atlantic puffins (Fratercula arctica) were equipped with geolocator/immersion loggers on each leg to attempt to overcome issues of leg-tucking in plumage while sitting on the water, which confounds the interpretation of logger data. Light level and saltwater immersion time-series data were combined to correct for this issue. This approach was adapted and applied to 40 puffins equipped with the standard practice deployments of geolocators on one leg only. Flightless periods consistent with moult were identified in the dual-equipped birds, whereas moult identification in single-equipped birds was less definitive and should be treated with caution. Within the dual-equipped sample, we present evidence for two flightless moult periods per non-breeding season in two puffins that undertook more extensive migrations (> 2000km), and were flightless for up to 76 days in a single non-breeding season. A biannual flight feather moult is highly unusual among non-passerine birds, and may be unique to birds that undergo catastrophic moult, i.e. become flightless when moulting. Though our conclusions are based on a small sample, we have established a freely available methodological framework for future investigation of the moult patterns of this and other seabird species.