Résumé: Coordinated movements of seabirds exploiting a prey patch are known to increase prey encounter and capture rates of individuals. These behaviours, based on effective cooperation between seabirds, have only been reported at small scale, i.e. the scale of the prey patch. However, the efficient prey exploitation by seabirds in vast oceans require larger scale processes such as information transfers between individuals. Indeed, information transfers between foraging seabirds (e.g. changes in behaviour) reduce their search cost while increasing their prey encounter rate. Whether or not these information transfer processes imply active cooperation is unknown. Using images from fishing boat radars in the eastern tropical Atlantic, we show the existence of frequent medium-scale patterns of coordinated flights of seabird groups, consisting in seabird fronts ('rake' patterns) of 0.3-4.4 km width, displacing cohesively over 1.2-10.6 km and lasting between 2 and 19 min. For these rakes to be maintained, seabird groups have to adjust their flight speeds and directions, while they are on average distant of 500 m from each other, what cannot occur by chance. These findings suggest the existence of collective and coordinated movements in seabirds during prey searching at several kilometres' scale. This potential cooperation between foraging seabird groups brings new insight in the evolutionary trajectories of seabirds life-style.

Résumé: In marine ecosystems top predator populations are shaped by environmental factors affecting their prey abundance. Coupling top predators’ population studies with independent records of prey abundance suggests that prey fluctuations affect fecundity parameters and abundance of their predators. However, prey may be abundant but inaccessible to their predators and a major challenge is to determine the relative importance of prey accessibility in shaping seabird populations. In addition, disentangling the effects of prey abundance and accessibility from the effects of prey removal by fisheries, while accounting for density dependence, remains challenging for marine top predators. Here, we investigate how climate, population density, and the accessibility and removal of prey (the Peruvian anchovy Engraulis ringens) by fisheries influence the population dynamics of the largest sedentary seabird community (≈ 4 million individuals belonging to guanay cormorant Phalacrocorax bougainvillii, Peruvian booby Sula variegata and Peruvian pelican Pelecanus thagus) of the northern Humboldt Current System over the past half-century. Using Gompertz state–space models we found strong evidence for density dependence in abundance for the three seabird species. After accounting for density dependence, sea surface temperature, prey accessibility (defined by the depth of the upper limit of the subsurface oxygen minimum zone) and prey removal by fisheries were retained as the best predictors of annual population size across species. These factors affected seabird abundance the current year and with year lags, suggesting effects on several demographic parameters including breeding propensity and adult survival. These findings highlight the effects of prey accessibility and fishery removals on seabird populations in marine ecosystems. This will help refine management objectives of marine ecosystems in order to ensure sufficient biomass of forage fish to avoid constraining seabird population dynamics, while taking into account of the effects of environmental variability.

Résumé: Seabirds are known to concentrate on prey patches or at predators aggregations standing for potential feeding opportunities. They may search for prey using olfaction or by detecting visually feeding con-specifics and sub-surface predators, or even boats. Thus, they might form a foraging network. We hypothesized that conditionally to the existence of a foraging network, the visual detection ability of seabirds should have a bearing on their medium-scale distribution at sea. Using a fishing-boat radar to catch the instantaneous distribution of seabirds groups within 30 km around the vessel, we conducted a spatial clustering of the seabird-echoes. We found 7,657 clusters (i.e. aggregations of echoes), lasting less than 15 minutes and measuring 9.2 km in maximum length (median). Distances between seabirds groups within clusters showed little variation (median: 2.1 km; CV: 0.5), while area varied largely (median: 21.9 km2; CV: 0.8). Given existing data on seabirds’ reaction distances to boats or other marine predators, we suggest that these structures may represent active foraging sequences of seabirds spreading themselves in space such as to possibly cue on each others. These seabird clusters were not previously described and are size compatible with the existence of a foraging network.

Résumé: Most seabirds are very noisy at their breeding colonies, when aggregated in high densities. Calls are used for individual recognition and also emitted during agonistic interactions. When at sea, many seabirds aggregate over patchily distributed resources and may benefit from foraging in groups. Because these aggregations are so common, it raises the question of whether seabirds use acoustic communication when foraging at sea? We deployed video-cameras with built in microphones on 36 Cape gannets (Morus capensis) during the breeding season of 2010-2011 at Bird Island (Algoa Bay, South Africa) to study their foraging behaviour and vocal activity at sea. Group formation was derived from the camera footage. During similar to 42 h, calls were recorded on 72 occasions from 16 birds. Vocalization exclusively took place in the presence of conspecifics, and mostly in feeding aggregations (81% of the vocalizations). From the observation of the behaviours of birds associated with the emission of calls, we suggest that the calls were emitted to avoid collisions between birds. Our observations show that at least some seabirds use acoustic communication when foraging at sea. These findings open up new perspectives for research on seabirds foraging ecology and their interactions at sea.

Résumé: The spatiotemporal distribution of animals is dependent on a suite of factors, including the distribution of resources, interactions within and between species, physiological limitations, and requirements for reproduction, dispersal, or migration. During breeding, reproductive constraints play a major role in the distribution and behavior of central place foragers, such as pelagic seabirds. We examined the foraging behavior and marine habitat selection of Laysan (Phoebastria immutabilis) and black-footed (P. nigripes) albatrosses throughout their eight month breeding cycle at Tern Island, Northwest Hawaiian Islands to evaluate how variable constraints of breeding influenced habitat availability and foraging decisions. We used satellite tracking and light-based geolocation to determine foraging locations of individuals, and applied a biologically realistic null usage model to generate control locations and model habitat preference under a case–control design. Remotely sensed oceanographic data were used to characterize albatross habitats in the North Pacific.

Résumé: Past tracking studies of marine animals have primarily targeted adults, biasing our understanding of at-sea habitat use toward older life stages. Anthropogenic threats persist throughout the at-sea ranges of all life stages and it is therefore of interest to population ecologists and managers alike to understand spatiotemporal distributions and possible niche differentiation between age-classes. In albatrosses, particularly little is known about the juvenile life stage when fledglings depart the colonies and venture to sea with no prior experience or parental guidance. We compared the dispersal of 22 fledgling Black-footed Albatross Phoebastria nigripes between 2006 and 2008 using satellite telemetry and 16 adults between 2008 and 2009 using geolocaters from Midway Atoll National Wildlife Refuge, Northwest Hawaiian Islands. Following tag deployment, all fledglings spent several days within the calm atoll waters, then travelled northward until reaching 750-900km from the colony. At this point, fledgling distributions approached the productive North Pacific Transition Zone (NPTZ). Rather than reaching the high chlorophyll a densities on the leading edge of this zone, however, fledglings remained in areas of low productivity in the subtropical gyre. In contrast, adult albatrosses from the same breeding colony did not utilize the NPTZ at this time of year but rather ranged throughout the highly productive northern periphery of the Pacific Ocean Basin among the shelf regions off Japan and the Aleutian Islands. The dichotomy in habitat use between fledglings and adults from Midway Atoll results in complete spatial segregation between age-classes and suggests ontogenetic niche separation in this species. This research fills a large knowledge gap in at-sea habitat use during a little known yet critical life stage of albatrosses, and contributes to a more comprehensive understanding of differential mortality pressure between age-classes and overall conservation status for the vulnerable Black-footed Albatross.

Résumé: During breeding, animal behaviour is particularly sensitive to environmental and food resource availability. Additionally, factors such as sex, body condition, and offspring developmental stage can influence behaviour. Amongst seabirds, behaviour is generally predictably affected by local foraging conditions and has therefore been suggested as a potentially useful proxy to indicate prey state. However, besides prey availability and distribution, a range of other variables also influence seabird behavior, and these need to be accounted for to increase the signal-to-noise ratio when assessing specific characteristics of the environment based on behavioural attributes. The aim of this study was to use continuous, fine-scale time-activity budget data from a pelagic seabird (Cape gannet, Morus capensis) to determine the influence of intrinsic (sex and body condition) and extrinsic (offspring and time) variables on parent behaviour during breeding. Foraging trip duration and chick provisioning rates were clearly sex-specific and associated with chick developmental stage. Females made fewer, longer foraging trips and spent less time at the nest during chick provisioning. These sex-specific differences became increasingly apparent with chick development. Additionally, parents in better body condition spent longer periods at their nests and those which returned later in the day had longer overall nest attendance bouts. Using recent technological advances, this study provides new insights into the foraging behaviour of breeding seabirds, particularly during the post-guarding phase. The biparental strategy of chick provisioning revealed in this study appears to be an example where the costs of egg development to the female are balanced by paternal-dominated chick provisioning particularly as the chick nears fledging.

Résumé: ABSTRACT Seabirds forage in a highly dynamic environment and prey on fish schools that are patchily distributed. Colonially breeding seabirds regularly commute back and forth from their colony to foraging areas and need to acquire information on the location of food before and/or during each foraging trip. The use of conspecifics as cues to locate prey has long been debated, and although the hypothesis was backed up by modeling studies, observations have been contradictory. We deployed GPS devices coupled with micro video cameras on Cape Gannets to observe the social context of foraging seabirds and the influence of conspecifics on the movement of individuals. The Cape Gannets reached their first patch using a succession of flights interrupted by stops on the water, during which the birds were mainly preening. During flight, the birds reacted to conspecifics by changing direction, either flying in the opposite direction of conspecifics that were flying toward the colony or following conspecifics outward. The time to reach the first patch was significantly reduced (by half) when the birds reacted to conspecifics in these different ways, compared with the birds that did not react. The use of conspecifics flying toward the colony to find food is consistent with the hypothesis that colonies can act as a focal place for information transfer, with foragers updating their flying direction when they detect conspecifics flying toward the colony. The fine-scale reaction of seabirds toward each other at sea, and the associated improved foraging efficiency, as well as the division of trips into a succession of flights, constitute elements that indicate the existence and the use of a structured network among foraging Cape Gannets. , RÉSUMÉ Les oiseaux marins se nourrissent dans un environnement fortement dynamique et sur des proies agrégées en bancs. Les oiseaux coloniaux font régulièrement des aller-retours entre la colonie et les zones d'alimentation, et doivent alors acquérir de l'information sur la localisation de leurs proies avant et pendant chaque voyage en mer. L'utilisation de congénères comme source d'information pour localiser des proies a longtemps été débattue, et bien que cette hypothèse soit soutenue par des modèles théoriques, les observations empiriques restent contradictoires. Nous avons déployé des GPS et micro-caméras sur des individus de Morus capensis afin d'observer le contexte social de ces oiseaux au cours de leur recherche alimentaire et l'influence de congénères sur le déplacement des individus. Ceux-ci ont rejoint leur première zone d'alimentation par une succession de vols, interrompus par des arrêts sur l'eau durant lesquels ils faisaient principalement leur toilette. En vol, les oiseaux ont réagi à leurs congénères en modifiant leur direction, soit pour aller en direction opposée de congénères qui volaient vers la colonie, soit pour suivre des congénères au large. Le temps pour rejoindre la première zone d'alimentation était significativement réduit (de moitié) pour les oiseaux ayant réagi à leurs congénères de ces différentes manières, comparé aux oiseaux n'ayant pas réagi. L'utilisation de congénères volant vers la colonie pour trouver de la nourriture est en accord avec l'hypothèse de l'utilisation de la colonie comme point central pour l'échange d'information, à partir duquel les individus en recherche de nourriture pourraient ajuster leur direction de vol au fur et à mesure qu'ils rencontrent des congénères de retour vers la colonie. Les réactions à fine échelle entre oiseaux en mer, associées à l'amélioration de leur efficacité pour trouver de la nourriture, ainsi que le découpage du trajet en vols successifs, constituent des éléments en faveur de l'existence et de l'utilisation d'un réseau structuré de recherche alimentaire chez M. capensis. Mots-clés: biologging, caméra, centre d'information, GPS, oiseaux marins, prédateur à place centrale, recherche alimentaire en réseau, suivi

Résumé: The study of ecological and behavioral processes has been revolutionized in the last two decades with the rapid development of biologging-science. Recently, using image-capturing devices, some pilot studies demonstrated the potential of understanding marine vertebrate movement patterns in relation to their proximate, as opposed to remote sensed environmental contexts. Here, using miniaturized video cameras and GPS tracking recorders simultaneously, we show for the first time that information on the immediate visual surroundings of a foraging seabird, the Cape gannet, is fundamental in understanding the origins of its movement patterns. We found that movement patterns were related to specific stimuli which were mostly other predators such as gannets, dolphins or fishing boats. Contrary to a widely accepted idea, our data suggest that foraging seabirds are not directly looking for prey. Instead, they search for indicators of the presence of prey, the latter being targeted at the very last moment and at a very small scale. We demonstrate that movement patterns of foraging seabirds can be heavily driven by processes unobservable with conventional methodology. Except perhaps for large scale processes, local-enhancement seems to be the only ruling mechanism; this has profounds implications for ecosystem-based management of marine areas.

Résumé: Although oriented migrations have been identified in many terrestrial bird species, the post-breeding-season movements of seabirds are generally regarded as dispersive. We used geolocator tags to reveal post-breeding movements and winter distribution of northern gannets (Morus bassanus) at a meta-population scale. By focusing on five breeding colonies of European gannets, we show that their breeding and wintering grounds are connected by a major flyway running along the coasts of Western Europe and Africa. Moreover, maximum winter distance to colony was similar across colonies despite their wide latitudinal range. In contrast with the general opinion that large pelagic birds such as gannets have unlimited ranges beyond the breeding season, our findings strongly suggest oriented chain migration in northern gannets (a pattern in which populations move uniformly southward) and highlight the benefit of meta-population approaches for studying seabird movements. We argue that the inclusion of such processes in ocean management plans is essential to improve efforts in marine biodiversity conservation.

Résumé: A goal of seabird ecology is to relate the physiology, population dynamics, distribution and behaviour of seabirds with their biotic and abiotic environments. One of the most challenging aspects is to understand how seabirds interact with their environment when direct observations are not always possible or practical. In the present paper, we reviewed 218 published studies that examined associations between seabird distribution, behaviour and their environment, in order to assess current trends, weaknesses and the future directions of research. Based on the number of publications, it is evident that the field is growing rapidly and that methods for evaluating seabird distribution are becoming increasingly more sophisticated and are changing from Eulerian (grid-like) to Lagrangian (particle-like) data types. This has been accompanied by a reduction in the spatial and temporal scale of observation, where, in most cases, no behavioural information is inferred from Lagrangian data; instead they are often used as if they were Eulerian data. In parallel, environmental remote sensing is becoming more common; however, we did not record significant changes in the statistical approaches used to describe seabird distributions and used to link them with oceanographic variables. In particular, despite the spatially explicit nature of the data, spatial statistics have rarely been used. The vast majority of studies used environmental variables that described water masses (descriptive approach), whereas a few studies determined oceanographic features that enhance prey availability to seabirds (process-based approach). Future studies could enhance their ecological interpretation of seabird–environment interactions by making greater use of ad hoc statistical approaches that facilitate appropriate pattern detection (e.g. area-restricted searching pattern for birds, mesoscale patterns for environment). Furthermore, appropriate hypothesis testing and modelling that accounts for the spatially explicit, multiscale and multivariate nature of the interaction between seabirds and their habitats is recommended. Although quantitative methods currently exist (but are rarely used), further application could greatly improve our understanding of the processes linking seabird distribution to their environment.

Résumé: When two closely related species co-occur, each exhibiting sex-specific differences in size, resource partitioning is expected. We studied sex-specific foraging behaviour of two sympatric seabird species in the Gulf of California to disentangle the respective influence of species and sex, but also mass and size of individuals, on foraging behaviour observed. We used highly accurate data loggers to study movements, diving behaviour and activity of brown and blue-footed boobies rearing young chicks. Interspecific differences were limited; brown boobies had longer foraging trips and spent less time on the water than blue-footed. The major differences observed were sex specific, females of each species tended to have longer foraging trips, foraged farther from the colony, flew greater distances and had larger zones of area restricted search. These sex-specific differences were more prominent in brown than in blue-footed boobies. Analyses of diet and stable isotopes show that during the study period, both species fed mainly on sardines, at similar trophic levels and in similar zones; outside the breeding season the carbon and nitrogen signatures from feathers were also similar on average. In these sympatric species feeding on a super abundant prey, sex-specific differences appear to have a greater role than species-specific differences, and it is suggested that sex-specific differences may be mainly related to breeding involvement, males being more involved in nest attendance and defence and females being greater provisioners. However, we show that several sex specific differences in foraging behaviour observed were partly or totally explained by body size (flight speeds, foraging range, flapping frequency) or by body mass (depths attained during diving, and duration of dives), parameters influenced by biomechanic constraints such as flight and diving.