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Although mechanisms of genetic and social inheritance have been implicated in determining the migratory routes of birds, it is unclear what their relative contributions are in species where outbound and return migration routes differ (‘loop migrants’). Here, we used biologging devices to follow Manx Shearwaters Puffinus puffinus, a long-lived seabird with a trans-Atlantic loop migration, from before their first migration until their 3rd calendar year. We found that Shearwaters undertake first migration without their parents, setting off almost immediately upon fledging and moving along a more direct trajectory than adults, before wintering in the same part of the South Atlantic as adult conspecifics and subsequently iteratively developing their return migration route over the next 3 years, each time returning – unlike adults – via a Western Atlantic route. We propose that the first outbound migration in Manx Shearwaters is broadly consistent with a genetically inherited vector, that both the outbound and the return migration trajectories are unlikely to be learnt from experienced conspecifics, and that return migration in Manx Shearwaters (and perhaps loop migrants more generally) may be informed by genetically inherited information and/or local environmental conditions.

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.

Funding: This work was supported by the University of Oxford Christopher Welch Scholarship (to J.A.K.); ASAB Undergraduate Project Scholarship (to J.A.K.); UKRI BBSRC scholarship grant number BB/M011224/1 (to J.W. and N.G.); The Queen’s College, University of Oxford (to A.L.F.); Junior Research Fellowship at St. John’s College, University of Oxford (to O.P.); Merton College, University of Oxford (to T.G.); Mary Griffiths Award (to T.G.); BBSRC David Phillips Fellowship grant numbers BB/G023913/1 and BB/ G023913/2 (to C.R.); and Jesus College, University of Oxford (to G.K.T.). This project has received funding from the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreement no. 682501) (to G.K.T.) Dynamic soaring harvests energy from a spatiotemporal wind gradient, allowing albatrosses to glide over vast distances. However, its use is challenging to demonstrate empirically and has yet to be confirmed in other seabirds. Here, we investigate how flap-gliding Manx shearwaters optimize their flight for dynamic soaring. We do so by deriving a new metric, the horizontal wind effectiveness, that quantifies how effectively flight harvests energy from a shear layer. We evaluate this metric empirically for fine-scale trajectories reconstructed from bird-borne video data using a simplified flight dynamics model. We find that the birds' undulations are phased with their horizontal turning to optimize energy harvesting. We also assess the opportunity for energy harvesting in long-range, GPS-logged foraging trajectories and find that Manx shearwaters optimize their flight to increase the opportunity for dynamic soaring during favorable wind conditions. Our results show how small-scale dynamic soaring affects large-scale Manx shearwater distribution at sea. Publisher PDF

Dynamic soaring harvests energy from a spatiotemporal wind gradient, allowing albatrosses to glide over vast distances. However, its use is challenging to demonstrate empirically, and has yet to be confirmed in other seabirds. Here we investigate how flap-gliding Manx Shearwaters optimise their flight for dynamic soaring. We do so by deriving a new metric, the horizontal wind effectiveness, that quantifies how effectively flight harvests energy from a shear layer. We evaluate this metric empirically for fine-scale trajectories reconstructed from bird-borne video data using a simplified flight dynamics model. We find that the birds’ undulations are phased with their horizontal turning to optimise energy harvesting. We also assess the opportunity for energy harvesting in long-range, GPS-logged foraging trajectories, and find that Manx Shearwaters optimise their flight to increase the opportunity for dynamic soaring during favourable wind conditions. Our results show how small-scale dynamic soaring impacts large-scale Manx Shearwater distribution at sea.TeaserFlap-gliding shearwaters harvest wind energy by fine-scale trajectory optimization and this impacts their large-scale distribution at sea.

Albatrosses are the iconic aerial wanderers of the oceans, supremely adapted for long-distance dynamic soaring flight. Perhaps because of this they are considered poorly adapted for diving1, in contrast to many smaller shearwater and petrel relatives, despite having amphibious eyes2, and an a priori mass advantage for oxygen-storage tolerance3. Modern biologging studies have largely confirmed this view4,5, casting doubt on earlier observations using capillary tube maximum depth gauges1, which may exaggerate depths, and emphasising albatrosses’ reliance on near-surface feeding. Nevertheless, uncertainty about albatross diving remains an important knowledge gap since bycatch in human fisheries (e.g. birds becoming hooked when diving for longline bait fish) is thought to be driving many population declines in this most threatened group of birds6. Here we show, using miniature electronic depth loggers (TDRs), that black-browed albatross, Thalassarche melanophris, can dive to much greater depths (19 m) and for much longer (52 s) than previously thought — three times the maxima previously recorded for this species (6 m and 15 s), and more than twice the maxima reliably recorded previously for any albatross (from 113.7 bird-days of tracking4,5,7). Further evidence that diving may be a significant behavioural adaptation in some albatrosses comes from co-deployed 3-axis accelerometers showing that these deeper dives, which occur in most individuals we tracked, involve active under-water propulsion without detectable initial assistance from momentum, sometimes with bottom phases typical of active prey pursuit. Furthermore, we find (from co-deployed GPS) that diving occurs primarily in the distal portions of long-distance foraging trips, with deeper dives occurring exclusively during daylight or civil twilight, confirming the importance of visual guidance. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

For studies on magnetic compass orientation and navigation performance in small bird species, controlled experiments with orientation cages inside an electromagnetic coil system are the most prominent methodological paradigm. These are, however, not applicable when studying larger bird species and/or orientation behaviour during free flight. For this, researchers have followed a very different approach, attaching small magnets to birds, with the intention of depriving them of access to meaningful magnetic information. Unfortunately, results from studies using this approach appear rather inconsistent. As these are based on experiments with birds under free-flight conditions, which usually do not allow exclusion of other potential orientation cues, an assessment of the overall efficacy of this approach is difficult to conduct. Here, we directly tested the efficacy of small magnets for temporarily disrupting magnetic compass orientation in small migratory songbirds using orientation cages under controlled experimental conditions. We found that birds which have access to the Earth's magnetic field as their sole orientation cue show a general orientation towards their seasonally appropriate migratory direction. When carrying magnets on their forehead under these conditions, the same birds become disoriented. However, under changed conditions that allow birds access to other (i.e. celestial) orientation cues, any disruptive effect of the magnets they carry appears obscured. Our results provide clear evidence for the efficacy of the magnet approach for temporarily disrupting magnetic compass orientation in birds, but also reveal its limitations for application in experiments under free-flight conditions., Summary: Orientation tests in a small migratory songbird provide clear evidence for the efficacy of the magnet approach for temporarily disrupting magnetic compass orientation, but also reveal its limitations in experiments under free-flight conditions.

There is increasing evidence for impacts of light pollution on the physiology and behaviour of wild animals. Nocturnally active Procellariiform seabirds are often found grounded in areas polluted by light and struggle to take to the air again without human intervention. Hence, understanding their responses to different wavelengths and intensities of light is urgently needed to inform mitigation measures. Here, we demonstrate how different light characteristics can affect the nocturnal flight of Manx shearwaters Puffinus puffinus by experimentally introducing lights at a colony subject to low levels of light pollution due to passing ships and coastal developments. The density of birds in flight above the colony was measured using a thermal imaging camera. We compared number of flying shearwaters under dark conditions and in response to an artificially introduced light, and observed fewer birds in flight during ‘light-on’ periods, suggesting that adult shearwaters were repelled by the light. This effect was stronger with higher light intensity, increasing duration of ‘light-on’ periods and with green and blue compared to red light. Thus, we recommend lower light intensity, red colour, and shorter duration of ‘light-on’ periods as mitigation measures to reduce the effects of light at breeding colonies and in their vicinity.

The conservation of migratory marine species, including pelagic seabirds, is challenging because their movements span vast distances frequently beyond national jurisdictions. Here, we aim to identify important aggregations of seabirds in the North Atlantic to inform ongoing regional conservation efforts. Using tracking, phenology, and population data, we mapped the abundance and diversity of 21 seabird species. This revealed a major hotspot associated with a discrete area of the subpolar frontal zone, used annually by 2.9–5 million seabirds from ≥56 colonies in the Atlantic: the first time this magnitude of seabird concentrations has been documented in the high seas. The hotspot is temporally stable and amenable to site-based conservation and is under consideration as a marine protected area by the OSPAR Commission. Protection could help mitigate current and future threats facing species in the area. Overall, our approach provides an exemplar data-driven pathway for future conservation efforts on the high seas. info:eu-repo/semantics/publishedVersion

Significance Procellariiform seabirds homing from distant foraging locations present a natural situation in which the homing route can become obstructed by islands or peninsulas because birds will not travel long distances over land. By measuring initial orientation from Global Positioning System (GPS) tracks during homing, we found that the Manx shearwater fails to encode such obstacles while homing, implying a navigation system that encodes the direction of home rather than a learned route. Nonetheless, shearwaters timed their journeys home, implying that their navigational system provides them with information about both direction and distance home, providing evidence that for routine, yet long-distance navigation, seabirds probably ascertain homeward direction by comparing their current position and the location of home with 2 or more intersecting field gradients., While displacement experiments have been powerful for determining the sensory basis of homing navigation in birds, they have left unresolved important cognitive aspects of navigation such as what birds know about their location relative to home and the anticipated route. Here, we analyze the free-ranging Global Positioning System (GPS) tracks of a large sample (n = 707) of Manx shearwater, Puffinus puffinus, foraging trips to investigate, from a cognitive perspective, what a wild, pelagic seabird knows as it begins to home naturally. By exploiting a kind of natural experimental contrast (journeys with or without intervening obstacles) we first show that, at the start of homing, sometimes hundreds of kilometers from the colony, shearwaters are well oriented in the homeward direction, but often fail to encode intervening barriers over which they will not fly (islands or peninsulas), constrained to flying farther as a result. Second, shearwaters time their homing journeys, leaving earlier in the day when they have farther to go, and this ability to judge distance home also apparently ignores intervening obstacles. Thus, at the start of homing, shearwaters appear to be making navigational decisions using both geographic direction and distance to the goal. Since we find no decrease in orientation accuracy with trip length, duration, or tortuosity, path integration mechanisms cannot account for these findings. Instead, our results imply that a navigational mechanism used to direct natural large-scale movements in wild pelagic seabirds has map-like properties and is probably based on large-scale gradients.

Seabirds care for their offspring in remote breeding colonies where foraging sites are distant and may be unpredictable, and where chicks are left unaccompanied for extended periods during their parents’ foraging trips, leaving them vulnerable to predation or starvation. One way to mitigate this risk is for individuals to coordinate parenting duties with their partner. Many biparental and cooperatively breeding species are now known to coordinate their care, though the mechanisms underlying this are not well understood. In Manx shearwaters (Puffinus puffinus), both parents alternate shifts of incubation in a coordinated manner. To resolve which processes may underlie this routine, we imposed a wing-loading handicap on parents to reduce their foraging efficiency, forcing them to choose between an extended foraging trip or to return to the nest before their condition has recovered to optimal levels. We found that handicapped parents took significantly longer trips than normal, to which their partner responded by lengthening their incubation shift, suggesting shift durations are not pre-determined. However, the duration of foraging trips and the mass at which foraging birds returned to the nest appeared to be mediated by the condition of the partner. These results suggest that while foraging trip duration is largely driven by the need for the foraging bird to recoup its own condition losses, information-transfer between the parents may facilitate a more cooperative mechanism whereby the decisions made by foraging birds still account for the condition of their partner.

The functions of display between breeding pairs of animals have been given little attention outside of sexual selection. Yet evidence suggests that display between partners is in fact most commonly observed following mate choice, and is often just as elaborate. In many bird species, allopreening, when one member of a pair preens the other, is a major component of display both pre- and post-pair formation. Despite this, there has been little investigation into its functions. Explanations that have been put forward tend to focus on its role in feather hygiene, which has limited phylogenetic support, or its function in the maintenance of the pair bond, though how this might occur or indeed what this actually represents has not been adequately explained. Phylogenetic evidence reveals that allopreening is most commonly observed in those species exhibiting high levels of partner retention and biparental care, and it appears to be functional in maintaining cooperation in parental behaviour in at least one species. In our observational study, we explored the patterns and putative functions of allopreening during the nest-relief displays of breeding pairs of Black-browed Albatrosses Thalassarche melanophris during incubation and chick-provisioning. Allopreening was an important feature of displays, constituting 30% of display time. We found that the bird returning from its foraging trip usually initiated allopreening, and preened more than its partner prior to change-over of nesting duties. We further found a positive relationship between the amount of time the pair spent in display and the duration of the subsequent foraging trip, providing tentative support for a function in maintaining cooperative parental behaviour between the parents. Although we cannot be conclusive as to its exact functions, we add to a limited literature the first exploration of functions for this conspicuous behaviour in albatrosses. info:eu-repo/semantics/publishedVersion

Each winter, the North Atlantic Ocean is the stage for numerous cyclones, the most severe ones leading to seabird mass-mortality events called ‘‘winter wrecks.’’ During these, thousands of emaciated seabird carcasses are washed ashore along European and North American coasts. Winter cyclones can therefore shape seabird population dynamics by affecting survival rates as well as the body condition of surviving individuals and thus their future reproduction. However, most often the geographic origins of impacted seabirds and the causes of their deaths remain unclear. We performed the first ocean-basin scale assessment of cyclone exposure in a seabird community by coupling winter tracking ∼ 1,500 individuals of five key North Atlantic seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia, and Rissa tridactyla) and cyclone locations. We then explored the energetic consequences of different cyclonic conditions using a mechanistic bioenergetics model and tested the hypothesis that cyclones dramatically increase seabird energy requirements. We demonstrated that cyclones of high intensity impacted birds from all studied species and breeding colonies during winter but especially those aggregating in the Labrador Sea, the Davis Strait, the surroundings of Iceland, and the Barents Sea. Our broad-scale analyses suggested that cyclonic conditions do not increase seabird energy requirements, implying that they die because of the unavailability of their prey and/or their inability to feed during cyclones. Our study provides essential information on seabird cyclone exposure in a context of marked cyclone regime changes due to global warming. at-sea distributioncyclonesenergy expenditureGLS trackingseabird migrationseascape ecology North Atlantic winter cyclones starve seabirds

Aim Europe's only globally critically endangered seabird, the Balearic shearwater (Puffinus mauretanicus), is thought to have expanded its postbreeding range northwards into UK waters, though its at sea distribution there is not yet well understood. This study aims to identify environmental factors associated with the species’ presence, map the probability of presence of the species across the western English Channel and southern Celtic Sea, and estimate the number of individuals in this area. Location The western English Channel and southern Celtic Sea. Methods This study analyses strip transect data collected between 2013 and 2017 from vessel‐based surveys in the western English Channel and southern Celtic Sea during the Balearic shearwater's postbreeding period. Using environmental data collected directly and from remote sensors both Generalized Additive Models and the Random Forest machine learning model were used to determine shearwater presence at different locations. Abundance was estimated separately using a density multiplication approach. Results Both models indicated that oceanographic features were better predictors of shearwater presence than fish abundance. Seafloor aspect, sea surface temperature, depth, salinity, and maximum current speed were the most important predictors. The estimated number of Balearic shearwaters in the prediction area ranged from 652 birds in 2017 to 6,904 birds in 2014. Main conclusions Areas with consistently high probabilities of shearwater presence were identified at the Celtic Sea front. Our estimates suggest that the study area in southwest Britain supports between 2% and 23% of the global population of Balearic shearwaters. Based on the timing of the surveys (mainly in October), it is probable that most of the sighted shearwaters were immatures. This study provides the most complete understanding of Balearic shearwater distribution in UK waters available to date, information that will help inform any future conservation actions concerning this endangered species., Areas with consistently high probabilities of shearwater presence were identified at the Celtic Sea front. Our estimates suggest that the study area in southwest Britain supports between 2% and 23% of the global population of Balearic shearwaters. This study provides the most complete understanding of Balearic shearwater distribution in UK waters available to date, information that will help inform any future UK conservation actions concerning this endangered species.

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.

Biologging has emerged as one of the most powerful and widely used technologies in ethology and ecology, providing unprecedented insight into animal behaviour. However, attaching loggers to animals may alter their behaviour, leading to the collection of data that fails to represent natural activity accurately. This is of particular concern in free-ranging animals, where tagged individuals can rarely be monitored directly. One of the most commonly reported measures of impact is breeding success, but this ignores potential short-term alterations to individual behaviour. When collecting ecological or behavioural data, such changes can have important consequences for the inference of results. Here, we take a multifaceted approach to investigate whether tagging leads to short-term behavioural changes, and whether these are later reflected in breeding performance, in a pelagic seabird. We analyse a long-term dataset of tracking data from Manx shearwaters (Puffinus puffinus), comparing the effects of carrying no device, small geolocator (GLS) devices (0.6% body mass), large Global Positioning System (GPS) devices (4.2% body mass) and a combination of the two (4.8% body mass). Despite exhibiting normal breeding success in both the year of tagging and the following year, incubating birds carrying GPS devices altered their foraging behaviour compared to untagged birds. During their foraging trips, GPS-tagged birds doubled their time away from the nest, experienced reduced foraging gains (64% reduction in mass gained per day) and reduced flight time by 14%. These findings demonstrate that the perceived impacts of device deployment depends on the scale over which they are sought: long-term measures, such as breeding success, can obscure finer-scale behavioural change, potentially limiting the validity of using GPS to infer at-sea behaviour when answering behavioural or ecological questions.

In migratory animals for whom post-natal care is limited, it is essential that there are inherited mechanisms whereby an individual can navigate—first, to the terminus of their migration, and second, back to a suitable breeding site. In birds, empirical evidence suggests that orientation on first migration is controlled by an inherited navigational vector, a direction and a distance in which to move (the “clock and compass” model) [1, 2, 3, 4, 5]. The mechanism and information that underlie the return to the natal breeding site are, however, almost entirely unknown. A potential solution to this problem would be for an animal to learn the values for spatially and temporally stable gradient cues that specifically indicate the location of the natal site [6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16]. One potential cue for latitude is magnetic inclination. Here, we use ringing recoveries made over the last 80 years to investigate whether magnetic inclination might be used as a navigational cue to control the latitude of recruitment in a trans-global migrant, the Manx shearwater (Puffinus puffinus). We find that small changes in inclination between when a bird fledges and when it returns from first migration correlate with probabilistic changes in latitude at recruitment, in doing so quantitatively fulfilling a priori predictions as to the magnitude and direction of latitudinal shift. This, we believe, suggests that (1) natal magnetic inclination is learnt prior to fledging and (2) is used to provide latitudinal information when making the first return trip from the wintering grounds.

Este artículo contiene 13 páginas, 5 figuras, 1 tabla., Patterns of behavioural variation and migratory connectivity are important characteristics of populations, particularly at the edges of species distributions, where processes involved in influencing evolutionary trajectories, such as divergence, mutual persistence, and natural hybridization, can occur. Here, we focused on two closely related seabird species that breed in the Mediterranean: Balearic shearwaters (Puffinus mauretanicus) and Yelkouan shearwaters (Puffinus yelkouan). Genetic and phenotypic evidence of hybridization between the two species on Menorca (the eastern and westernmost island in the breeding ranges of the two shearwaters, respectively) has provided important insights into relationships between these recently diverged species. Nevertheless, levels of behavioural and ecological differentiation amongst these populations remain largely unknown. Using geolocation and stable isotopes, we compared the at-sea movement behaviour of birds from the Menorcan ‘hybrid’ population with the nearest neighbouring populations of Balearic and Yelkouan shearwaters. The Menorcan population displayed a suite of behavioural features intermediate to those seen in the two species (including migration strategies, breeding season movements and limited data on phenology). Our findings provide new evidence to support suggestions that the Menorcan population is admixed, and indicate a role of non-breeding behaviours in the evolutionary trajectories of Puffinus shearwaters in the Mediterranean., This study was funded by the Natural Environment Research Council, UK (grant numbers: NE/J500227/1, EK192-02/12).The National Oceanography Centre, the Total Foundation and Merton College – University of Oxford provided additional support. The Yelkouan shearwater project was funded by the French Biodiversity Agency for Marine Protected Areas within the ‘Programme PACOMM, Natura2000 en mer’. MG is supported by the Spanish Ministry of Science (CGL2017-85210).

Knowing the spatial scales at which effective management can be implemented is fundamental for conservation planning. This is especially important for mobile species, which can be exposed to threats across large areas, but the space use requirements of different species can vary to an extent that might render some management approaches inefficient. Here the space use patterns of seabirds were examined to provide guidance on whether conservation management approaches should be tailored for taxonomic groups with different movement characteristics. Seabird tracking data were synthesised from 5419 adult breeding individuals of 52 species in ten families that were collected in the Atlantic Ocean basin between 1998 and 2017. Two key aspects of spatial distribution were quantified, namely how far seabirds ranged from their colony, and to what extent individuals from the same colony used the same areas at sea. There was evidence for substantial differences in patterns of space-use among the ten studied seabird families, indicating that several alternative conservation management approaches are needed. Several species exhibited large foraging ranges and little aggregation at sea, indicating that area-based conservation solutions would have to be extremely large to adequately protect such species. The results highlight that short-ranging and aggregating species such as cormorants, auks, some penguins, and gulls would benefit from conservation approaches at relatively small spatial scales during their breeding season. However, improved regulation of fisheries, bycatch, pollution and other threats over large spatial scales will be needed for wide-ranging and dispersed species such as albatrosses, petrels, storm petrels and frigatebirds. Fil: Oppel, Steffen. Royal Society For The Protection Of Birds; Fil: Bolton, Mark. Royal Society For The Protection Of Birds; Fil: Carneiro, Ana P.B.. Birdlife International; Fil: Dias, Maria P.. Birdlife International; Fil: Green, Jonathan A.. University of Liverpool; Reino Unido Fil: Masello, Juan F.. Justus Liebig University Giessen; Fil: Phillips, Richard A.. British Antarctic Survey; Fil: Owen, Ellie. Royal Society For The Protection Of Birds; Fil: Quillfeldt, Petra. Justus Liebig University Giessen; Fil: Beard, Annalea. Environmental And Natural Resources Directorate; Fil: Bertrand, Sophie. Universidad Federal Rural Pernambuco; Brasil. Centre National de la Recherche Scientifique. Institut de Recherche pour le Développement; Francia Fil: Blackburn, Jez. British Trust For Ornithology; Fil: Boersma, P. Dee. University Of Washington, Seattle; Fil: Borges, Alder. Projecto Vitó; Fil: Broderick, Annette C.. University of Exeter; Reino Unido Fil: Catry, Paulo. Ispa - Instituto Universitário; Portugal Fil: Cleasby, Ian. Royal Society For The Protection Of Birds; Fil: Clingham, Elizabeth. Environmental And Natural Resources Directorate; Fil: Creuwels, Jeroen. Naturalis Biodiversity Center; Fil: Crofts, Sarah. Falklands Conservation; Fil: Cuthbert, Richard J.. Royal Society For The Protection Of Birds; Fil: Dallmeijer, Hanneke. Veda Consultancy; Fil: Davies, Delia. University Of Cape Town; Fil: Davies, Rachel. Marinelife; Fil: Dilley, Ben J.. University Of Cape Town; Fil: Dinis, Herculano Andrade. Parque Natural Do Fogo; Fil: Dossa, Justine. Birdlife International; Fil: Dunn, Michael J.. British Antarctic Survey; Fil: Efe, Marcio A.. Universidade Federal de Alagoas; Brasil Fil: Fayet, Annette L.. University of Oxford; Reino Unido Fil: Figueiredo, Leila. Universidade Federal de Alagoas; Brasil Fil: Frederico, Adelcides Pereira. Projecto Vitó; Fil: Gjerdrum, Carina. Environment Canada; Canadá Fil: Godley, Brendan J.. University of Exeter; Reino Unido Fil: Granadeiro, José Pedro. Faculdade de Ciências, Universidade de Lisboa; Portugal Fil: Guilford, Tim. University of Oxford; Reino Unido Fil: Hamer, Keith C.. University Of Leeds; Fil: Hazin, Carolina. Birdlife International; Fil: Hedd, April. Environment Canada; Canadá Fil: Henry, Leeann. Environmental And Natural Resources Directorate; Fil: Hernández-Montero, Marcos. Projeto Biodiversidade; Fil: Hinke, Jefferson. Noaa Fisheries Service; Fil: Kokubun, Nobuo. National Institute Of Polar Research; Fil: Leat, Eliza. Ascension Island Government; Fil: Tranquilla, Laura McFarlane. Royal Society For The Protection Of Birds; . Environmental And Natural Resources Directorate; Fil: Metzger, Benjamin. Birdlife Malta; Malta Fil: Militão, Teresa. Universidad de Barcelona; España Fil: Montrond, Gilson. Projecto Vitó; Fil: Mullié, Wim. Veda Consultancy; Fil: Padget, Oliver. University of Oxford; Reino Unido Fil: Pearmain, Elizabeth J.. Birdlife International; Fil: Pollet, Ingrid L.. Bird Studies Canada; Canadá Fil: Pütz, Klemens. Antarctic Research Trust; Fil: Quintana, Flavio Roberto. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Centro Nacional Patagónico. Instituto de Biología de Organismos Marinos; Argentina Fil: Ratcliffe, Norman. British Antarctic Survey; Fil: Ronconi, Robert A.. Environment Canada; Canadá Fil: Ryan, Peter G.. University Of Cape Town; Fil: Saldanha, Sarah. Universidad de Barcelona; España Fil: Shoji, Akiko. University of Oxford; Reino Unido Fil: Sim, Jolene. Ascension Island Government; Fil: Small, Cleo. Royal Society For The Protection Of Birds; Fil: Soanes, Louise. University of Liverpool; Reino Unido Fil: Takahashi, Akinori. National Institute Of Polar Research; Fil: Trathan, Phil. British Antarctic Survey; Fil: Trivelpiece, Wayne. Noaa Fisheries Service; Fil: Veen, Jan. Veda Consultancy; Fil: Wakefield, Ewan. University of Glasgow; Reino Unido Fil: Weber, Nicola. University of Exeter; Reino Unido. Ascension Island Government; Fil: Weber, Sam. Ascension Island Government; . University of Exeter; Reino Unido Fil: Zango, Laura. Universidad de Barcelona; España Fil: González-Solís, Jacob. Universidad de Barcelona; España Fil: Croxall, John. Birdlife International

Capsule Dive behaviour in Black Guillemots Cepphus grylle was close to that predicted for a benthic forager by allometry based on body mass.Aims To report the diving behaviour of Black Guillemots in Northern Ireland.Methods A time-depth-recorders and GPS logger were deployed on four chick-rearing breeding Black Guillemots.Results Dive shape implied most dives were benthic with a small number at the start of each bout associated with searching. Diving only occurred during daylight hours, but dive depth was unrelated to light availability outside of the twilight periods. Dive durations (max = 90 s; mean = 54 s) were shorter and dive depths (max = 15 m; mean = 9 m) were shallower than recorded for guillemots elsewhere. The birds dived a maximum of 1.8 km from the colony.Conclusions Black Guillemots' dive duration was similar to the value predicted from allometry, and is therefore likely representative of this species. Bathymetry likely influences the dive behaviour of this benthic-feeding species because mos...

In long-lived species, care-giving parents are expected to balance their own condition with that of their offspring. Many species of seabirds display a unique behavioural adaptation for managing these conflicting demands known as dual foraging, in which long trips, largely for self-maintenance, are alternated with short trips, which are primarily for offspring care. While dual foraging is a widely studied behaviour, it entails a complication that is seldom discussed: if parents independently employ a dual-foraging strategy, chicks might be abandoned for extended periods when the long trips of both partners coincide. Whether partners coordinate their dual-foraging strategies, however, is largely unknown. To investigate this possibility, we used radiofrequency identification readers coupled with passive integrated transponder tags to record extended sequences of foraging trips for breeding Manx shearwaters, Puffinus puffinus. Our results show a pattern of foraging trips that indicates a high level of coordination between parents, which facilitates consistent provisioning. Additionally, we show that the propensity for pairs to coordinate declines across the chick-rearing period. Given the potential costs of not coordinating, we expect this behaviour to be widely spread among dual-foraging species.

Global wind patterns affect flight strategies in many birds, including pelagic seabirds, many of which use wind-powered soaring to reduce energy costs during at-sea foraging trips and migration. Such long-distance movement patterns are underpinned by local interactions between wind conditions and flight behaviour, but these fine-scale relationships are far less well understood. Here we show that remotely sensed ocean wind speed and direction are highly significant predictors of soaring behaviour in a migratory pelagic seabird, the Manx shearwater ( Puffinus puffinus ). We used high-frequency GPS tracking data (10 Hz) and statistical behaviour state classification to identify two energetic modes in at-sea flight, corresponding to flap-like and soar-like flight. We show that soaring is significantly more likely to occur in tailwinds and crosswinds above a wind speed threshold of around 8 m s −1 , suggesting that these conditions enable birds to reduce metabolic costs by preferentially soaring over flapping. Our results suggest a behavioural mechanism by which wind conditions may shape foraging and migration ecology in pelagic seabirds, and thus indicate that shifts in wind patterns driven by climate change could impact this and other species. They also emphasize the emerging potential of high-frequency GPS biologgers to provide detailed quantitative insights into fine-scale flight behaviour in free-living animals.

Population-level estimates of species' distributions can reveal fundamental ecological processes and facilitate conservation. However, these may be difficult to obtain for mobile species, especially colonial central-place foragers (CCPFs; e.g., bats, corvids, social insects), because it is often impractical to determine the provenance of individuals observed beyond breeding sites. Moreover, some CCPFs, especially in the marine realm (e.g., pinnipeds, turtles, and seabirds) are difficult to observe because they range tens to ten thousands of kilometers from their colonies. It is hypothesized that the distribution of CCPFs depends largely on habitat availability and intraspecific competition. Modeling these effects may therefore allow distributions to be estimated from samples of individual spatial usage. Such data can be obtained for an increasing number of species using tracking technology. However, techniques for estimating population-level distributions using the telemetry data are poorly developed. This is of concern because many marine CCPFs, such as seabirds, are threatened by anthropogenic activities. Here, we aim to estimate the distribution at sea of four seabird species, foraging from approximately 5,500 breeding sites in Britain and Ireland. To do so, we GPS-tracked a sample of 230 European Shags Phalacrocorax aristotelis, 464 Black-legged Kittiwakes Rissa tridactyla, 178 Common Murres Uria aalge, and 281 Razorbills Alca torda from 13, 20, 12, and 14 colonies, respectively. Using Poisson point process habitat use models, we show that distribution at sea is dependent on (1) density-dependent competition among sympatric conspecifics (all species) and parapatric conspecifics (Kittiwakes and Murres); (2) habitat accessibility and coastal geometry, such that birds travel further from colonies with limited access to the sea; and (3) regional habitat availability. Using these models, we predict space use by birds from unobserved colonies and thereby map the distribution at sea of each species at both the colony and regional level. Space use by all four species' British breeding populations is concentrated in the coastal waters of Scotland, highlighting the need for robust conservation measures in this area. The techniques we present are applicable to any CCPF.

Individual foraging specializations, where individuals use a small component of the population niche width, are widespread in nature with important ecological and evolutionary implications. In long-lived animals, foraging ability develops with age, but we know little about the ontogeny of individuality in foraging. Here we use precision global positioning system (GPS) loggers to examine how individual foraging site fidelity (IFSF), a common component of foraging specialization, varies between breeders, failed breeders and immatures in a long-lived marine predator—the northern gannet Morus bassanus. Breeders (aged 5+) showed strong IFSF: they had similar routes and were faithful to distal points during successive trips. However, centrally placed immatures (aged 2–3) were far more exploratory and lacked route or foraging site fidelity. Failed breeders were intermediate: some with strong fidelity, others being more exploratory. Individual foraging specializations were previously thought to arise as a function of heritable phenotypic differences or via social transmission. Our results instead suggest a third alternative—in long-lived species foraging sites are learned during exploratory behaviours early in life, which become canalized with age and experience, and refined where possible—the exploration-refinement foraging hypothesis. We speculate similar patterns may be present in other long-lived species and moreover that long periods of immaturity may be a consequence of such memory-based individual foraging strategies.

Compass orientation is central to the control of animal movement from the scale of local food-caching movements around a familiar area in parids and corvids to the first autumn vector navigation of songbirds embarking on long-distance migration. In the study of diurnal birds, where the homing pigeon, Columba livia, has been the main model, a time-compensated sun compass is central to the two-step map-and-compass process of navigation from unfamiliar places, as well as guiding movement via a representation of familiar area landmarks. However, its use by an actively navigating wild bird is yet to be shown. By phase shifting an animal's endogenous clock, known as clock-shifting, sun-compass use can be demonstrated when the animal incorrectly consults the sun's azimuthal position while homing after experimental displacement. By applying clock-shift techniques at the nest of a wild bird during natural incubation, we show here that an oceanic navigator—the Manx shearwater, Puffinus puffinus—incorporates information from a time-compensated sun compass during homeward guidance to the breeding colony after displacement. Consistently with homing pigeons navigating within their familiar area, we find that the effect of clock shift, while statistically robust, is partial in nature, possibly indicating the incorporation of guidance from landmarks into movement decisions. Padget et al. show that a wild bird makes use of a time-compensated sun compass during active navigation. By measuring the minute-by-minute orientation of GPS-tracked Manx shearwaters homing under clock shift from distant release sites, the authors demonstrate that a sun compass is fundamental even once visual landmarks are available.

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The way animals move through space is likely to affect the way they learn and remember spatial information. For example, a pelagic fish, Astyanax fasciatus, moves freely in vertical and horizontal space and encodes information from both dimensions with similar accuracy. Benthic fish can also move with six degrees of freedom, but spend much of their time travelling over the substrate; hence they might be expected to prioritise the horizontal dimension. To understand how benthic fish encode and deploy three-dimensional spatial information we used a fully rotational Y-maze to test whether Corydoras aeneus (i) encode space as an integrated three-dimensional unit or as separate elements, by testing whether they can decompose a three-dimensional trajectory into its vertical and horizontal components, and (ii) whether they prioritise vertical or horizontal information when the two conflict. In contradiction to the expectation generated by our hypothesis, our results suggest that C. aeneus are better at extracting vertical information than horizontal information from a three-dimensional trajectory, suggesting that the vertical axis is learned and remembered robustly. Our results also showed that C. aeneus prioritise vertical information when it conflicts with horizontal information. From these results, we infer that benthic fish attend preferentially to a cue unique to the vertical axis, and we suggest that this cue is hydrostatic pressure.

Several laboratory studies have revealed functional hemispheric lateralization in birds performing visual tasks. However, the role of functional brain asymmetries in spatial behaviour in natural settings is still poorly investigated. We studied monocularly occluded homing pigeons, Columba livia, to investigate potential differences in the hemispheric control of navigational performance. We GPS-tracked monocularly occluded and control binocular homing pigeons during seven group training releases and a final solitary release from each of two sites. The pigeons were then given one last release from each site after a phase shift of the light-dark cycle under binocular conditions, to distinguish compass-based orientation from landmark-based pilotage. Overall, pigeons homing with the left eye/right hemisphere (RH) displayed a greater fidelity to the familiar space previously experienced than pigeons homing with the right eye/left hemisphere (LH). Another difference between the two monocular groups is that LH pigeons were more likely than RH pigeons to fly with other pigeons during the group training releases. The data support the hypothesis that the left eye/right hemisphere plays a more substantial role as pigeons develop fidelity to certain routes to home from familiar release sites, an enhanced fidelity that may be supported by superior memory for familiar landmarks.

Before visiting or leaving their remote island colonies, seabirds often engage in a behaviour termed ‘rafting’, where birds sit, often in groups, on the water close to the colony. Despite rafting being a widespread behaviour across many seabird taxa, the functional significance of rafting remains unknown. Here we combine global positioning system (GPS) tracks, observational and wind condition data to investigate correlates of rafting behaviour in Manx shearwaters (Puffinus puffinus) at a large colony on Skomer Island, Wales. We test (1) the influence of wind direction on rafting location and (2) whether raft size changes with respect to wind speed. Our approach further allows us to describe day-night trends in (3) raft distance from shore through time; (4) the number of birds present in the nearshore waters through time; and (5) spatial patterns of Manx shearwater rafts in marine waters adjacent to the breeding colony. We find no evidence that wind direction, for our study period, influences Manx shearwater rafting location, yet raft size marginally increases on windier days. We further find rafting birds closer to the shore at night than during the day. Thus, before sunset, birds form a “halo” around Skomer Island, but this halo disappears during the night as more individuals return from foraging trips and raft nearer the colony on Skomer Island. The halo pattern reforms before sunrise as rafts move away from land and birds leave for foraging. Our results suggest that wind conditions may not be as ecologically significant for rafting locations as previously suspected, but rafting behaviour may be especially important for avoiding predators and cleaning feathers.

World-wide, many seabirds are affected by fisheries in opposing ways: as a source of mortality from bycatch, but also by providing discards as a predictable and abundant food resource. This applies to the Balearic shearwater Puffinus mauretanicus, the most endangered European seabird, whose time to extinction was estimated at only ˜40 years a decade ago. Since the previous assessment, new data and more sophisticated demographic modelling have become available, and new fishing policies from the European Union (Common Fisheries Policy, CFP) will apply, posing different scenarios for the viability of the species. Thus, there is both an urgent need and an opportunity for a more reliable update of the conservation status of the species. Demographic data were collected between 1985 and 2014 at one of the world's largest colonies. Most demographic parameters were estimated using multievent capture–recapture modelling. Some parameters, such as bycatch rate, immature individual survival and recruitment, were estimated for the first time. We incorporated estimates into stochastic population models to forecast time to extinction and assess the viability under different management scenarios, accounting for upcoming fishing policies. Adult survival was much lower than expected (0·809, SE: 0·013) and largely influenced by bycatch, which accounted for a minimum of 0·455 (SE: 0·230) of total mortality. Breeding success was positively correlated with discard availability. Recruitment started at low rates in 3-year-old birds (0·030, SE: 0·0455), increasing in following age classes and was almost complete at 6 years. Under the present scenario, we predict a time to extinction of 61 years (95% CI: 55–69). Synthesis and applications. Population projections suggest that the actual impact of fisheries on Balearic shearwaters is unsustainable and the imminent discard ban under the new Common Fisheries Policy may accelerate the declining trend. This study demonstrates that reducing the bycatch rates of fisheries is an unavoidable and urgent conservation measure for avoiding the extinction of the species. We also advise setting up demographic long-term studies, to allow researchers to diagnose, with reliability, the effectiveness of management actions. These actions will also benefit many other marine top-predator species affected by this anthropogenic impact., Funds were provided by FEDER programme (Balearic Government), the Spanish Ministries of Economy and Competitiveness (ref. CGL2013-42203-R) and Agriculture, Food and Environment, the H2020 framework EU programme (MINOUW project) and Fundación Biodiversidad.

Despite decades of interest, the adaptive significance of the extraordinary diversity in the design of animal signals remains elusive. It is suggested that signal design consists of two components: ‘strategic design’ and ‘efficacy’. Strategic design is concerned with how a signal is constructed by natural selection to provide the information necessary to make a receiver respond (e.g. by being good at displaying underlying quality), whilst efficacy is concerned with how a signal is designed to get that information across to the receiver (e.g. by being easily measured). It is argued that an important but neglected evolutionary force on animal signals is therefore the psychology of the signal receiver, and that three aspects of receiver psychology (what a receiver finds easy to detect, easy to discriminate and easy to remember) constitute powerful selective forces in signal design. Greatest emphasis is given to memorability because this has been least considered by previous authors. It is argued that learning and memory are involved in a wide range of signals, and numerous hypotheses as to how signals may be adapted to be more memorable to receivers are suggested. The relationship of this analysis to earlier attempts at understanding signals is explored, particularly with reference to the concepts of honesty, manipulation and mind-reading.

Animal communication is studied both by neurobiologists and by evolutionary biologists, but in very different ways. The purpose of this article is to show how both groups could benefit from a greater appreciation of each other's approach. Evolutionary biologists should take more account of the role played by the sensory systems and brains of receivers in constraining the design of animal signals. Neurobiologists should be more aware of recent advances in the understanding of signal—receiver co-evolution and the evolutionary origins of animal signals. A series of recent examples are cited that illustrate how pre-existing neurophysiological or psychological properties of receiver organisms are essential to our understanding of the design characteristics of animal signals and of their origins. Also discussed are a number of other areas of signalling in which the study of ‘receiver psychology' is likely to be fruitful.

The warning signals of toxic insects are often 'multimodal', combining bright coloration with sounds or odours (or both). Pyrazine (a common insect warning odour) can elicit an intrinsic avoidance in domestic chicks Gallus gallus domesticus, both against novel coloured food, and also against food colours that are specifically associated with aposematism, namely yellow and red. In three experiments, we investigated the role of novelty in this innate bias against yellow coloured food in the presence of pyrazine. Naive chicks were familiarized either to pyrazine odour or to coloured food before being tested for a bias against yellow (warningly coloured) food as opposed to green (nonwarningly coloured) food. In experiment 1, pyrazine novelty was shown to be vital for eliciting a bias against yellow food. However, experiment 2 suggested that colour novelty was not important: chicks familiarized with coloured crumbs still avoided yellow crumbs when pyrazine was presented. In a third experiment that gave chicks an even greater degree of pre-exposure to coloured crumbs, the bias against yellow food eventually waned, although pyrazine continued to elicit an aversion to yellow even after birds had had experience of up to 24 palatable yellow crumbs. Pyrazine novelty has been an important pressure in the evolution of multimodal warning signals, and can continue to promote the avoidance of warningly coloured food, even when it is relatively familiar. The implications for warning signals are discussed. Copyright 1999 The Association for the Study of Animal Behaviour.