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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.

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

1. As more and more species face anthropogenic threats, understanding the causes of population declines in vulnerable taxa is essential. However, long-term datasets, ideal to identify lasting or indirect effects on fitness measures such as those caused by environmental factors, are not always available. 2. Here we use a single year but multi-population approach on populations with contrasting demographic trends to identify possible drivers and mechanisms of seabird population changes in the north-east Atlantic, using the Atlantic puffin, a declining species, as a model system. 3. We combine miniature GPS trackers with camera traps and DNA metabarcoding techniques on four populations across the puffins’ main breeding range to provide the most comprehensive study of the species' foraging ecology to date. 4. We find that puffins use a dual foraging tactic combining short and long foraging trips in all four populations, but declining populations in southern Iceland and north-west Norway have much greater foraging ranges, which require more (costly) flight, as well as lower chick-provisioning frequencies, and a more diverse but likely less energy-dense diet, than stable populations in northern Iceland and Wales. 5. Together, our findings suggest that the poor productivity of declining puffin populations in the north-east Atlantic is driven by breeding adults being forced to forage far from the colony, presumably because of low prey availability near colonies, possibly amplified by intraspecific competition. Our results provide valuable information for the conservation of this and other important North-Atlantic species and highlight the potential of multi-population approaches to answer important questions about the ecological drivers of population trends. biologging, diet, DNA metabarcoding, dual foraging, foraging ecology, intraspecific competition, population decline, seabirds

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.

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.

Domesticated animals have been culturally and economically important throughout history. Many of their ancestral lineages are extinct or genetically en dangered following hybridization with domesticated relatives. Consequently, they have been understudied compared to the ancestral lineages of domestic plants. The domestic pigeon Columba livia, which was pivotal in Darwin’s studies, has maintained outsized cultural significance. Its role as a model organism spans the fields of behavior, genetics, and evolution. Domestic pigeons have hybridized with their progenitor, the Rock Dove, rendering the latter of dubious genetic sta tus. Here, we use genomic and morphological data from the putative Rock Doves of the British Isles to identify relictual undomesticated populations. We reveal that Outer Hebridean Rock Doves have experienced minimal levels of introgres sion. Our results outline the contemporary status of these wild pigeons, high lighting the role of hybridization in the homogenization of genetic lineages.

The inverse optimality approach can allow us to learn about an animal's environment by assuming their behaviour is optimal. This approach has been applied to animals diving underwater for food to produce the index of patch quality (IPQ), which aims to provide a proxy for prey abundance or quality in a foraging patch based on the animal's diving behaviour. The IPQ has been used in several empirical studies but has never been evaluated theoretically. Here, we discuss the strengths and weaknesses of the IPQ approach from a theoretical angle and review the empirical evidence supporting its use. We highlight several potential issues, in particular with the gain function—the function describing the energetic gain of an animal during a dive—used to calculate the IPQ. We investigate an alternative gain function which is appropriate in some cases, provide a new model based on this function, and discuss differences between the IPQ model and ours. We also find that there is little supporting empirical evidence justifying the general use of the IPQ and suggest future empirical validation methods which could help strengthen the case for the IPQ. Our findings have implications for the field of diving ecology and habitat assessment.

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

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.

Documenting novel cases of tool use in wild animals can inform our understanding of the evolutionary drivers of the behavior’s emergence in the natural world. We describe a previously unknown tool-use behavior for wild birds, so far only documented in the wild in primates and elephants. We observed 2 Atlantic puffins at their breeding colonies, one in Wales and the other in Iceland (the latter captured on camera), spontaneously using a small wooden stick to scratch their bodies. The importance of these observations is 3-fold. First, while to date only a single form of body-care-related tool use has been recorded in wild birds (anting), our finding shows that the wild avian tool-use repertoire is wider than previously thought and extends to contexts other than food extraction. Second, we expand the taxonomic breadth of tool use to include another group of birds, seabirds, and a different suborder (Lari). Third, our independent observations span a distance of more than 1,700 km, suggesting that occasional tool use may be widespread in this group, and that seabirds’ physical cognition may have been underestimated.

In Focus: Campioni L., Dias M. P., Granadeiro J. P., & Catry P. (2019). An ontogenetic perspective on migratory strategy of a long-lived pelagic seabird: Timings and destinations change progressively during maturation. Journal of Animal Ecology, 89, 29-43. https://doi.org/10.1111/1365-2656.13044 How migration routes develop in long-lived migratory animals remains poorly understood. Campioni et al. (2019) tracked the migration of immature and breeding Cory's shearwaters and found that the timings and routes gradually changed with age. Young birds followed more exploratory routes at first, but gradually shortened and defined them as they matured, and they returned to the breeding grounds progressively earlier. These findings suggest that long-lived species start by exploring potentially suitable areas for the non-breeding season and progressively refine their migration route as they age.

International audience; Inter-individual variation in behavioural traits has important implications for evolutionary and ecological processes. Site fidelity, where individuals consistently use the same foraging site, is common among marine predators. Sex differences in foraging are also well studied in marine vertebrates, but the extent to which consistent inter-individual differences in foraging vary between the sexes is poorly known. Here we quantify the effects of sex on individual foraging site fidelity (IFSF), both within and between years, in chick brooding Campbell albatross (Thalassarche impavida). Using bird-borne Global Positioning System loggers, we calculated route fidelity (nearest neighbour distance), repeatability of site fidelity (terminal latitude and longitude), and foraging effort (total distance travelled and trip duration) during two to ten repeat trips. Overall, Campbell albatrosses showed a high degree of site fidelity. Birds travelled to similar sites not only within the same year, but also between two consecutive years, suggesting that the within-year consistency is not simply in response to short-term patches of food. Moreover, within the same year there were differences in terms of IFSF between the sexes. Females that foraged closer to the colony in neritic and shelf waters were more likely to follow similar routes on repeated foraging trips and were more consistent in their foraging effort than males. Males that foraged further offshore in pelagic waters had more repeatable foraging longitudes than females. Our study provides further evidence of the importance of IFSF among marine vertebrates. However, it also reveals that the strength of such specialisations may vary with sex.

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.

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.

Socially learned signals and behaviors are found throughout the animal kingdom and often show striking variation within species over space and time. However, the extent to which this cultural diversity is generated by demographic factors such as immigration and dispersal has proved difficult to investigate and remains largely untested in a natural setting. Focusing on 14 different local neighborhoods within a wild population of great tits Parus major, we examined the relationship between individual movements over a 7-year period and patterns of cultural diversity at a neighborhood level, assessed using 2 indices: Song repertoire size and repertoire novelty, measured at the neighborhood (but not individual) level. We found that the repertoire size of a neighborhood is positively correlated with its levels of immigration, but that repertoire novelty is constrained by dispersal and geographic proximity among neighborhoods, both of which promote song sharing. In parallel, we show that the acoustic structure of some songs changes subtly over distance, in line with the view that learning errors can lead to the establishment of novel social traits (cultural differentiation). Our findings demonstrate that spatial variation in cultural diversity in animal societies can partly be explained as the outcome of dispersal and immigration transferring traits between neighborhoods and also suggest that trait transmission declines in fidelity over distance.

When individuals invest in a common good, an efficient outcome is hard to achieve, because each can free ride on others’ efforts. This problem can lead parents that raise their young together to reduce their investment in care, with negative consequences for offspring. Here, we present a mathematical model to show that a strategy of conditional cooperation, in which parents take turns feeding their young, can help to resolve this problem. To test this idea, we studied the behavior of great tit parents raising chicks together. We found that parents alternated visits to the nest more than would be expected by chance, speeding up their feeding rate after their partner had visited the chicks, but slowing down again once they had visited in turn. This effect was not mediated by visit-to-visit changes in offspring begging intensity, although females (but not males) were influenced by mean begging levels across broods. We conclude that conflict over parental investment in this species is partly ameliorated by a simple form of reciprocity.

Summary Long‐lived migratory animals must balance the cost of current reproduction with their own condition ahead of a challenging migration and future reproduction. In these species, carry‐over effects, which occur when events in one season affect the outcome of the subsequent season, may be particularly exacerbated. However, how carry‐over effects influence future breeding outcomes and whether (and how) they also affect behaviour during migration and wintering is unclear.Here we investigate carry‐over effects induced by a controlled, bidirectional manipulation of the duration of reproductive effort on the migratory, wintering and subsequent breeding behaviour of a long‐lived migratory seabird, the Manx shearwater Puffinus puffinus. By cross‐fostering chicks of different age between nests, we successfully prolonged or shortened by ∼25% the chick‐rearing period of 42 breeding pairs. We tracked the adults with geolocators over the subsequent year and combined migration route data with at‐sea activity budgets obtained from high‐resolution saltwater‐immersion data. Migratory behaviour was also recorded during non‐experimental years (the year before and/or two years after manipulation) for a subset of birds, allowing comparison between experimental and non‐experimental years within treatment groups.All birds cared for chicks until normal fledging age, resulting in birds with a longer breeding period delaying their departure on migration; however, birds that finished breeding earlier did not start migrating earlier. Increased reproductive effort resulted in less time spent at the wintering grounds, a reduction in time spent resting daily and a delayed start of breeding with lighter eggs and chicks and lower breeding success the following breeding season. Conversely, reduced reproductive effort resulted in more time resting and less time foraging during the winter, but a similar breeding phenology and success compared with control birds the following year, suggesting that ‘positive’ carry‐over effects may also occur but perhaps have a less long‐lasting impact than those incurred from increased reproductive effort.Our results shed light on how carry‐over effects can develop and modify an adult animal's behaviour year‐round and reveal how a complex interaction between current and future reproductive fitness, individual condition and external constraints can influence life‐history decisions.

Competition and, ultimately, adaptive specialization are the major ecological forces behind spatial segregation in foraging distributions, and are commonly driven by size-related differences in competitiveness between individuals of different sex, age or social status. However, such segregation can also be observed in long-lived monomorphic species, often between immature and breeding individuals. In many of these species, individuals often forage in patchy and potentially unpredictable environments in which resources can be spread over large scales and be difficult to find, and efficient foraging may require advanced cognitive skills (for example in navigation and memory). Particularly in species with deferred breeding, experience rather than size may be an important driver of segregation and may lead to differences in competitiveness between young and old, but whether there is a relationship between age, foraging efficiency and spatial segregation has never been properly investigated. Here we tested this hypothesis by simultaneously tracking individuals at different life stages in a long-lived seabird, the Manx shearwater, Puffinus puffinus, during a period of central-place foraging around the colony, to investigate spatial segregation, and by measuring foraging efficiency by combining an ethoinformatics approach and mass gain. We found substantial spatial segregation between immature and breeding adults. Compared with adults, immatures gained less mass per unit of time spent foraging and foraged in less productive waters, suggesting lower foraging efficiency, probably because of inexperience.

In order to maximize foraging efficiency in a varying environment, predators are expected to optimize their search strategy. Environmental conditions are one important factor affecting these movement patterns, but variations in breeding constraints (self-feeding vs. feeding young and self-feeding) during different breeding stages (incubation vs. chick-rearing) are often overlooked, so that the mechanisms responsible for such behavioral shifts are still unknown. Here, to test how search patterns are affected at different breeding stages and to explore the proximate causes of these variations, we deployed data loggers to record both position (global positioning system) and dive activity (time–depth recorders) of a colonial breeding seabird, the razorbill Alca torda. Over a period of 3 years, our recordings of 56 foraging trips from 18 breeders show that while there is no evidence for individual route fidelity, razorbills exhibit higher foraging flexibility during incubation than during chick rearing, when foraging becomes more focused on an area of high primary productivity. We further show that this behavioral shift is not due to a shift in search patterns, as reorientations during foraging are independent of breeding stage. Our results suggest that foraging flexibility and search patterns are unlinked, perhaps because birds can read cues from their environment, including conspecifics, to optimize their foraging efficiency. Electronic supplementary material The online version of this article (doi:10.1007/s00227-016-2826-x) contains supplementary material, which is available to authorized users.

Many passerine birds lay white eggs with reddish brown speckles produced by protoporphyrin pigment. However, the function of these spots is contested. Recently, the sexually selected eggshell coloration (SSEC) hypothesis proposed that eggshell color is a sexually selected signal through which a female advertises her quality (and hence the potential quality of her future young) to her male partner, thereby encouraging him to contribute more to breeding attempts. We performed a test of the SSEC hypothesis in a common passerine, the great tit Parus major. We used a double cross-fostering design to determine whether males change their provisioning behavior based on eggshell patterns they observe at the nest. We also tested the assumption that egg patterning reflects female and/or offspring quality. Because birds differ from humans in their color and pattern perception, we used digital photography and models of bird vision to quantify egg patterns objectively. Neither male provisioning nor chick growth was related to the pattern of eggs males observed during incubation. Although heavy females laid paler, less speckled eggs, these eggs did not produce chicks that grew faster. Therefore, we conclude that the SSEC hypothesis is an unlikely explanation for the evolution of egg speckling in great tits.

Using combined miniature archival light and salt-water immersion loggers, we characterise the year-round individual at-sea movements of Europe's only critically endangered seabird, the Balearic shearwater Puffinus mauretanicus, for the first time. Focusing on the non-breeding period, we show that all of the 26 breeding birds tracked from their breeding site on Mallorca in the Mediterranean Sea successfully made a 2–4 month migration into the Atlantic Ocean, where they utilised well-defined core areas off Portuguese and French coasts. As well as identifying high-risk areas in the Atlantic, our results confirm that breeding birds spend most of the year concentrated around productive waters of the Iberian shelf in the western Mediterranean. Migration phenology appeared largely unrelated to the subsequent (distinctly synchronous) breeding attempt, suggesting that any carry-over effects were compensated for during a long pre-laying period spent over winter in the Mediterranean. Using the light and salt-water immersion data alone we were also able to characterise the pattern of pre-laying visits to the colony in considerable detail, demonstrating that breeding pairs appear to coordinate their over-day visits using a high frequency of night-time visits throughout the winter. Our study shows that geolocation technology is a valuable tool for assessing the spatial distribution of risks to this critically endangered species, and also provides a low-impact method for remotely observing the detailed behaviour of seabird species that may be sensitive to disturbance from traditional study methods.