Studying fine-scale movements of seabirds during migration is logistically challenging, but GPS technologies allow accurate tracking of individuals on their migratory journeys. Such data provide essential information in the context of offshore wind farm (OWF) developments, notably to anticipate spatial OWF overlap with migratory corridors and main foraging areas used along the routes of vulnerable species. Using high-precision GPS-GSM tags, we investigated the end of summer, southbound migration of two emblematic seabirds of French waters: Juvenile and adult Northern gannets Morus bassanus, and adult Balearic shearwaters Puffinus mauretanicus. Both species travel along the Southwestern European coast, between the Bay of Biscay and Western Africa, or the Mediterranean Sea. Adult gannets thereby migrate through the EEZ of up to 10 countries, six for juvenile gannet, and four for shearwaters. Combining behavioural segmentation based on hidden Markov models and utilization distribution modelling, we found that between two and 6 % of migration routes overlapped with proposed OWFs, with similar impacts on transit and foraging/resting areas. Studied seabirds were most at risk within Portuguese compared to Spanish waters, as they flew closest to OWFs (<10 km on average). While massive OWF developments are being planned within Western European coastal areas, our study suggests that offshore developments should be set >22 km away from the coast, to preserve transnational seabird migratory corridors.

High pathogenicity avian influenza virus (HPAIV) caused the worst seabird mass-mortalities on record in Europe across 2021-2022. The northern gannet ( Morus bassanus ) was one of the most affected species, with tens of thousands of casualties in the northeast Atlantic between April-September 2022. Disease outbreaks can drastically modify the movement ecology of animals and diminish spatial consistency, thereby increasing the potential for disease transmission. To detect potential changes in movement behaviour, we GPS-tracked breeding adults following the initial HPAIV outbreak, at three of the largest gannet breeding colonies where major mortality of adults and chicks occurred (Bass Rock, Scotland, UK; Grassholm, Wales, UK; Rouzic island, Brittany, France). Crucially, GPS-tracked birds remained faithful to their breeding sites and did not prospect other breeding colonies. They performed regular foraging trips at sea, similar to their behaviour before the outbreak. Gannet foraging effort was nonetheless lower than in 2019, thus surviving birds may have benefited from reduced intra- and interspecific food competition. Breeding colony fidelity of adult northern gannets following HPAIV mass-mortalities suggests limited long-term capacity to virus spread, which may contrast with the behaviour of adults during the disease outbreak, or with that of younger individuals.

Body-mounted accelerometers provide a new prospect for estimating power use in flying birds, as the signal varies with the two major kinematic determinants of aerodynamic power: wingbeat frequency and amplitude. Yet wingbeat frequency is sometimes used as a proxy for power output in isolation. There is, therefore, a need to understand which kinematic parameter birds vary and whether this is predicted by flight mode (e.g. accelerating, ascending/descending flight), speed or morphology. We investigate this using high-frequency acceleration data from (i) 14 species flying in the wild, (ii) two species flying in controlled conditions in a wind tunnel and (iii) a review of experimental and field studies. While wingbeat frequency and amplitude were positively correlated, R 2 values were generally low, supporting the idea that parameters can vary independently. Indeed, birds were more likely to modulate wingbeat amplitude for more energy-demanding flight modes, including climbing and take-off. Nonetheless, the striking variability, even within species and flight types, highlights the complexity of describing the kinematic relationships, which appear sensitive to both the biological and physical context. Notwithstanding this, acceleration metrics that incorporate both kinematic parameters should be more robust proxies for power than wingbeat frequency alone.

Early life is a critical phase of the life cycle of animals and is attracting increased attention because little information is available on the behaviour of young individuals during this period. Behaviour during early life is probably influenced by the environmental conditions encountered by young animals, but data on intraspecific variation between breeding sites during this crucial period of life are limited. Here we study variability in the foraging behaviour of juveniles and adults in three colonies of a pantropical seabird, the Red‐footed Booby Sula sula. Both adults and juveniles were measured and fitted with GPS loggers in three remote islands: Genovesa (Galapagos, Eastern Pacific Ocean), Europa (Western Indian Ocean) and Surprise (New Caledonia, Western Pacific Ocean). Foraging behaviour was compared between age‐classes, sex and colonies by examining trip characteristics, different behaviours at sea, potential associations between individuals and morphological characteristics. Compared with adults, juveniles conducted shorter trips that were restricted to around the colony, especially on Genovesa (max. range: 203.4 ± 125.1 km and 3.6 ± 3.1 km, respectively). Juveniles appeared more constrained by poor flight skills and experience rather than by their morphology. Adults travelled 45% of the time during at‐sea trips, whereas juveniles spent a a lower proportion of time travelling but foraged more often using an ‘area‐restricted search’ behaviour, potentially training to catch prey. Associations between juveniles were commonly detected in the three colonies and occurred mostly during foraging, suggesting that social learning is an important strategy. Variability of morphometric measurements in both adults and juveniles was high between sites, with larger birds found on Genovesa. These results suggest that adaptations to local environmental conditions are already visible in their early life. Future studies should continue to investigate the behavioural flexibility of juvenile birds to better understand the effect of local environmental conditions during this critical stage of life.

A widespread hypothesis for the ontogeny of behavior and decision-making is the early-exploration-later-canalization hypothesis. It postulates that juveniles are more exploratory and adults more consistent in their behavior. In addition, it is often assumed that naive juveniles could overcome the costs of individual experience building by copying more the decisions of others than adults (early-conformism-later-self-defining hypothesis). Here, we compare the central place foraging movements of adults and postfledging juveniles in their first flights around the colony before dispersal and migration in two sympatric species of tropical seabirds: red-footed boobies and great frigatebirds. Using GPS records of individual movements, we analyzed the foraging directions of seabirds from the colony across successive trips. Juveniles of both species showed significant within-individual consistency in foraging direction but at lower levels than adults. Juveniles leaving the colony within the same time window showed significant but low between-individual resemblance in foraging direction at levels similar to adults. In both species, homing efficiency was lower in juveniles than in adults. Juvenile foraging directions were initially influenced by wind conditions, particularly in low wing loading frigate-birds. Wind conditions progressively lost influence on juvenile foraging directions during their first weeks of flights. In contrast, within-individual consistency, between-individual resemblance, and homing efficiency did not show signs of progression in juveniles. Our results support the early-exploration-later-canalization hypothesis but not the early-conformism-later-self-defining hypothesis. Relaxed constraints on self-feeding efficiency could favor high variability in postfledging tropical seabirds. Our simple approach could be applied to further test these hypotheses by comparing strategies across a wide range of central place foragers.

Lunar phase and illumination are known to affect nocturnal behavior of many organisms, particularly through predator-prey interactions. Visual predators can benefit from higher light levels to increase their activity, while prey may decrease their activity to avoid predation. The lower number of nocturnal seabirds observed on colonies during full moon nights has been mostly interpreted as a predation avoidance strategy. However, it is also possible that shearwaters take advantage of the moon's illumination to feed also at night, and stay at sea to forage during full moon nights. We used miniaturized GPS-loggers to obtain 179 tracks from 99 wedge-tailed shearwaters breeding in New Caledonia, to investigate moonlight effects on individual behavior. Lunar phase significantly predicted self-provisioning trip duration, with individuals performing longer trips around the full moon. However, this relationship was not significant during chick-provisioning trips when adults have to frequently return to the colony. Adults mostly returned to the colony during moonlit periods, refuting the predation avoidance theory. Tracked individuals showed an unexpectedly high amount of nocturnal foraging activity (28% of total activity), positively influenced by the presence of the moon. δ15N stable isotope values were significantly related to the percentage of nocturnal foraging, but with a weak relationship, impeding our ability to confirm that wedge-tailed shearwaters fed on different prey when foraging at night. This study suggests that reduced colony attendance around the full moon may be linked to greater at-sea foraging opportunities in distant oceanic areas than to increased predation risk on land.

The wedge-tailed shearwater (WTS) population of New Caledonia is one of the largest in the world, yet its biology and foraging ecology are poorly known. We studied WTS from4 colonies in New Caledonia. We examined foraging behaviour and habitats using GPS receivers and light sensors during and outside the breeding season, respectively, and compared our findings with those from other WTS populations worldwide. During breeding, New Caledonian WTS alternated short foraging trips close to the colony over the lagoon, or off the reef edge, with longer trips over distant, deep waters. Whereas neighboring colonies overlapped at sea, especially during short trips, there was a clear separation of foraging zones between the pairs of colonies located in the southern versus northwestern parts ofNewCaledonia. AlthoughWTSactively foraged and commuted to foraging zones during the day, they mainly returned to the colony or rested at night, indicating that they feed mainly during the day. Active foraging did not take place in more productive areas, suggesting that it may instead be related to the presence of sub-surface predators. Outside the breeding season, birds from 3 colonies had similar trans-equatorial migratory behaviour. All left New Caledonia at the same time of the year with a fast, northeasterly movement and wintered over deep waters in the same sector of the northwestern tropical Pacific Ocean. At overwintering sites, they spent most of their non-foraging time presumably sitting on the water, especially at night, making a slow westward movement before returning to New Caledonia. WTS from New Caledonia forage over warm, oligotrophic deep waters throughout their life cycle, and the species appears to have a flexible foraging strategy adapted to the various environmental conditions encountered across its wide tropical range.

Compensating for wind drift can improve goalward flight efficiency in animal taxa, especially among those that rely on thermal soaring to travel large distances. Little is known, however, about how animals acquire this ability. The great frigatebird ( Fregata minor ) exemplifies the challenges of wind drift compensation because it lives a highly pelagic lifestyle, travelling very long distances over the open ocean but without the ability to land on water. Using GPS tracks from fledgling frigatebirds, we followed young frigatebirds from the moment of fledging to investigate whether wind drift compensation was learnt and, if so, what sensory inputs underpinned it. We found that the effect of wind drift reduced significantly with both experience and access to visual landmark cues. Further, we found that the effect of experience on wind drift compensation was more pronounced when birds were out of sight of land. Our results suggest that improvement in wind drift compensation is not solely the product of either physical maturation or general improvements in flight control. Instead, we believe it is likely that they reflect how frigatebirds learn to process sensory information so as to reduce wind drift and maintain a constant course during goalward movement.

In the context of climate change, how extreme climatic events, such as cyclones, will affect the foraging abilities of marine vertebrates is still poorly known. During the course of a study on the foraging behaviour of two tropical seabirds, red-footed boobies and great frigatebirds, several cyclones have affected their breeding grounds and foraging zones, allowing us to study their response to extreme wind conditions. We examined whether adults and young naïve birds were able to predict the arrival of a cyclone and behave accordingly to reduce mortality risks and optimise foraging. We show that when a cyclone approached, juveniles and adults of the two species differed in their decisions to leave the colony for the sea. When the winds reached gale force, the juveniles of both species and adult frigatebirds remained at the colony, whereas adult boobies continued their foraging routine. The mortality of the individuals remaining on land remained limited. When encountering at-sea gale conditions, adult birds were able to avoid the centre of the low pressure systems and moved westward to bypass the route of the cyclones and circumvent the moving cyclone. Frigatebirds climb to high altitudes when close to the eye of the cyclone to bypass it at high speeds. These movements likely reduce the mortality risk at sea but can temporarily cause birds to move outside their normal range at sea or over land masses. We discuss the potential consequences of an increase in cyclonic conditions on seabird populations.

The early life of animals is a period of high mortality, when foraging capacities are assumed to be improved progressively. In birds, this critical period involves the improvement of the flight. How do young birds gain these capacities has rarely been studied in natural conditions especially in seabirds that spend most of their life at sea. We used detailed GPS and body acceleration data on 37 great frigatebirds (Fregata minor), to test the hypothesis that juveniles starting their first flights have lower flying capacities than adults, but that these capacities will improve during a long learning period, before independence from parents, specific to this seabird that can spend months on the wing at sea. We found that most flight components improved over time to tend towards those of adults, especially the travel speed, range, duration and maximum altitude of trips. However, unexpectedly, juveniles had higher ascent rates, soaring and gliding capacities above the sea than adults. Moreover, energy expenditure of juveniles was similar to adults during low cost travelling movements and during active foraging, but juveniles spent more time foraging actively than adults. Our results suggest that flight tactics based on long-distance effortless movements specific to this family are acquired during a long period, but soaring and gliding capacities are already inherited by juveniles and possibly favoured by morphological adaptations specific to juveniles. These adaptations might explain the extreme dispersive capacities of juveniles.

Drones and unmanned aerial vehicles are increasingly used in research on wildlife. Their wide applications can also give interesting insights into habitat use and population distribution. However, the disturbance they might be responsible for, on species and especially in protected areas has yet to be investigated. We assessed and compared the behavioural response of 11 southern seabird species at the Crozet Islands, Southern Indian Ocean, to drone approaches at specific altitudes. We first show that the behavioural response differed between species depending on the altitude of the drone approach. At 50 m of altitude, only one of the studied species showed a detectable reaction, whereas at 10 m, most species showed strong behavioural postures of stress. Adult penguins breeding in large colonies, and some albatross species showed little behavioural response even when the drone was as close as 3 m, whereas other species such as giant petrels or cormorants appeared highly sensitive to drone approaches. Among King Penguins, although incubating adults showed little signs of behavioural stress, non-breeding adults and fledglings in crèches exhibited strong behavioural responses to the drone approach. Monitoring heart rate allowed us to investigate the link between behavioural and physiological response to that specific potential stressor in king penguins. Whereas we confirmed the expected link between physiological and behavioural response in chicks, breeding adults showed no behavioural sign of stress but had a significant increase in heart rate, the relative increase being higher than in chicks. All together these results have important implications for the conservation of species and should be helpful for future legislations on the use of drones.

Migratory behavior varies extensively between bird taxa, from long distance migration to purely sedentary behavior. Variability in migratory behavior also occurs within taxa, where individuals within some species, or even populations, show mixed strategies. The same variability occurs in seabird species. We examined the migratory behavior of distinct populations of great frigatebirds Fregata minor in three distant oceanographic basins. Great frigatebird populations showed extensive variation in post-breeding migratory behavior. Birds from Europa Island (Mozambique Channel) made long-distance migration to numerous distinct roosting sites in the Indian Ocean, New Caledonia birds made shorter distance migrations to roosting sites in the southwestern Pacific Ocean, and Galapagos birds were resident within the archipelago year round. Juvenile birds from Europa Is. and New Caledonia dispersed widely whereas Galapagos juveniles were resident year round. The migratory behavior of Europa Is. and New Caledonia resulted in complete separation of foraging grounds between breeding adults, non-breeding adults, and juveniles, whereas in the Galapagos the overlap was complete. We suggest that population variability in migratory behavior may have arisen because of different environmental conditions at sea, and also depends on the availability of suitable roosting sites on oceanic islands. The results also highlight the capacity of frigatebirds to remain airborne most of the time even outside the breeding season when they have to molt.

The early life stages represent a crucial period that can strongly influence population dynamics. We studied the development of foraging behaviour in the red-footed booby, a tropical seabird with an extensive post-fledging care period (3 to 6 months). Adults and juveniles were observed from shore and tracked at sea using GPS loggers over 3 consecutive 12-day periods. Juveniles initially made a majority of flights inland, likely to practice flying, and formed groups of up to 10 juveniles before making short trips at sea. They left the island later and returned earlier than the adults, allowing them to be fed on the nest. Over time, juveniles left the colony alone more frequently and increased the range of their trips while remaining significantly closer to the colony than the adults. They spent more time intensively foraging (slow and sinuous trajectory) than adults, which could reflect attempts to capture prey. Juveniles foraged independently of their parents but associated frequently with congeners, particularly during area-restricted search (ARS) behaviour. The extensive post-fledging care period observed may be explained by the need to develop proper foraging skills adapted to tropical waters, where resources are particularly scarce and unpredictable.

While interspecific differences in foraging behaviour have attracted much attention, less is known about how foraging behaviour differs between populations of the same species. Here we compared the foraging strategy of a pantropical seabird, the red-footed booby Sula sula, in 5 populations breeding in contrasted environmental conditions. The foraging strategy strongly differed between sites, from strictly diurnal short trips in Europa Island (Mozambique channel) to long trips including up to 5 nights at sea in Genovesa Island (Galapagos archipelago). The Expectation Maximisation binary Clustering (EMbC) algorithm was used to determine the different behaviours of individuals during their foraging trips (travelling, intensive foraging, resting and relocating). During the day, the activity budget was similar for all the breeding colonies. During the night, birds were primarily on the water, drifting with currents. At all sites, birds similarly performed intensive foraging in zones of area-restricted search (ARS), although the size and duration of ARS zones differed markedly. Red-footed boobies foraged over deep oceanic waters, with chlorophyll a concentrations varying between sites. Birds did not appear to target areas with higher productivity. We suggest that range differences between populations may be linked to other factors such as intraand interspecific competition.

Understanding how animals respond to atmospheric conditions across space is critical for understanding the evolution of flight strategies and long-distance migrations. We studied the three-dimensional movements and energetics of great frigate birds (Fregata minor) and showed that they can stay aloft for months during transoceanic flights. To do this, birds track the edge of the doldrums to take advantage of favorable winds and strong convection. Locally, they use a roller-coaster flight, relying on thermals and wind to soar within a 50- to 600-meter altitude band under cumulus clouds and then glide over kilometers at low energy costs. To deal with the local scarcity of clouds and gain longer gliding distances, birds regularly soar inside cumulus clouds to use their strong updraft, and they can reach altitudes of 4000 meters, where freezing conditions occur.

Understanding how animals respond to atmospheric conditions across space is critical for understanding the evolution of flight strategies and long-distance migrations. We studied the three-dimensional movements and energetics of great frigate birds (Fregata minor) and showed that they can stay aloft for months during transoceanic flights. To do this, birds track the edge of the doldrums to take advantage of favorable winds and strong convection. Locally, they use a roller-coaster flight, relying on thermals and wind to soar within a 50- to 600-meter altitude band under cumulus clouds and then glide over kilometers at low energy costs. To deal with the local scarcity of clouds and gain longer gliding distances, birds regularly soar inside cumulus clouds to use their strong updraft, and they can reach altitudes of 4000 meters, where freezing conditions occur.

Seabirds are considered to be good indicators of the marine environment. However, little is known about the effects of environmental variability on the foraging behaviour of tropical seabirds. Red-footed boobies (RFB) nesting on Europa Island (Mozambique Channel) were fitted with GPS devices over four years and different breeding stages. We first show that the durations of foraging trips vary extensively according to the stage of the breeding, being short during brooding, intermediate during incubation and long during fledging. This result highlights the importance of considering breeding stage when conducting comparisons of foraging between sites or years. In addition, we show that RFB adjusted their foraging behaviour between years (2003, 2011, 2012 and 2013) according to the prevailing environmental conditions. During 2011, RFB made longer foraging trips with larger area-restricted search (ARS) zones over a larger total surface area, suggesting that the foraging conditions were probably poor. This year was characterized by a decrease of the major environmental drivers of the Mozambique Channel system, i.e. particularly low chlorophyll concentrations in the northern part of the Mozambique Channel, as well as a weak eddy activity. This observation suggests that environmental conditions may have altered the southward transport and concentration processes structuring the trophic chain, leading to adverse conditions for a central-place forager like the RFB. Our results emphasize that environmental and breeding stage variation should be taken into account to better understand the distribution of these predators in marine tropical ecosystems.

The annual moult creates the highest physiological stress during a penguin's breeding‐cycle and is preceded by a period of hyperphagia at sea. Although crucial to individual survival, foraging strategies before moult have been little investigated in keystone marine consumers in the Southern Ocean. The Macaroni Penguin Eudyptes chrysolophus demonstrates how individuals may adjust their foraging strategies during this period in line with constraints such as potential intraspecific competition between localities, foraging ability between dimorphic sexes and timing at sea between breeding and non‐breeding population components. We recorded pre‐moult behaviour at sea for 22 Macaroni Penguins from Crozet and Kerguelen Islands (southern Indian Ocean) during 2009 and 2011, using light‐based geolocation and stable isotope analysis. Penguins were distributed in population‐specific oceanic areas with similar surface temperatures (3.5 °C) south of the archipelagos, where they foraged at comparable trophic levels based on stable isotopes of their blood. Bayesian ‘broken stick’ modelling with concurrent analysis of seawater temperature records from the animal‐borne devices showed that within each population, females remained 6 days longer than males in the colder waters before heading back towards their colonies. Finally, 17 other non‐breeding individuals that moulted earlier had a higher mean blood δ 15 N value than did post‐breeding birds, meaning that early moulters probably fed more on fish than did late moulters. Our findings of such adjustments in foraging strategies developed across locality, sex and breeding status help understanding of the species' contrasted pre‐moult biology across its range and its ecology in the non‐breeding period.

Marine Protected Areas (MPAs) are being established across all marine regions but their validity for the conservation of highly mobile marine vertebrates has been questioned. We tested the hypothesis that French coastal MPAs primarily designed for coastal and benthic biota are also beneficial for the conservation of a pelagic seabird, the Vulnerable yelkouan shearwater (Puffinus yelkouan), an endemic species to the Mediterranean Sea. We used a vast spectrum of electronic devices (GPS, temperature-depth-recorders, satellite transmitters and geolocators) and stable isotopic analyses to study the year-round movements and the trophic status of yelkouan shearwaters from the Hyères archipelago (France). In addition we conducted large-scale ship and aircrafts observation surveys to investigate spatio-temporal density patterns of shearwaters (genus Puffinus) in the western Mediterranean Sea. This extensive investigation permitted the first comprehensive study of the at-sea ecology of yelkouan shearwaters showing strikingly coastal habits, partial migration, unsuspected diving capabilities (max dive depth of 30m), and a broad diet ranging from zooplankton to small pelagic fish. Importantly, 31% of yelkouan shearwaters GPS positions associated with foraging, 38% of diving positions, and 27% of resting positions were within the three French MPAs during the breeding season. These high scores confirmed by year-round distribution derived from GLS, PTTs, at-sea and aerial observations, validated our hypothesis of the major importance of coastal MPAs for the conservation of yelkouan shearwater. Our case-study is therefore a major contribution to research efforts aiming at linking the spatial ecology of highly mobile marine vertebrates with effective conservation of marine biodiversity.