Seabird populations are declining globally, in part due to a combination of climate change and fishery-related pressures. While the impacts of bycatch are well documented, empirical evidence for population effects of prey depletion by commercial fisheries is limited, particularly in systems where forage fish are managed conservatively. Therefore, evaluating whether existing management frameworks are effective at maintaining prey availability within ecologically safe limits presents an important, but underdeveloped, component of ecosystem-based fisheries management. We assessed the effectiveness of Barents Sea capelin (Mallotus villosus) fishery management in sustaining seabird populations, by modelling relationships between fisheries landings, prey availability, climate variability, and seabirds' adult survival. Using a path modelling framework, we integrated spatio-temporal data on capelin landings, prey abundance, and sea surface temperature across seabirds' distributions with long-term demographic data from eight breeding populations of four pelagic seabird species. Direct negative associations between capelin landings and adult survival were only detected in one population, corresponding to the strongest spatial overlap between this population's pre-breeding distribution and areas with high capelin harvest. Survival responses to prey availability and ocean temperature otherwise varied among species and populations, reflecting differences in their foraging ecology and space use. Krill abundance was positively associated with survival in two populations, while warmer autumn temperatures were linked to reduced capelin abundance, indicating climate-driven bottom-up effects. By focussing on adult survival as a response, a demographic rate to which seabird population trajectories are particularly sensitive, our results suggest that current capelin management has limited adverse effects on seabirds under contemporary environmental conditions. More broadly, this study demonstrates that seabird survival has the potential to be used as an indicator of management effectiveness due to its sensitivity to multiple environmental factors and its potential impact on population trends. This study highlights the potential for integrating seabird survival into ecosystem-based fisheries management through long-term monitoring and high-resolution fisheries data.

Climate-driven ocean warming is profoundly reshaping marine ecosystems, with cascading effects on biodiversity and trophic interactions. For migratory marine predators such as seabirds, demographic responses to warming depend on when and where populations are exposed across the annual cycle. Therefore, integrating demographic monitoring and tracking data, across broad geographic and temporal scales, is essential, given the spatial and seasonal variability in ocean warming. Here, we integrated long-term demographic data, seasonal distributions, and sea surface temperatures (SSTs) for 26 populations of five seabird species breeding in the North-East Atlantic to assess the effects of SSTs on reproduction, survival, and population growth trajectories. Demographic responses varied widely among populations and seasons, but negative effects were most consistently associated with warming during the autumn period postbreeding, particularly in the Barents and East Greenland Seas. Winter warming also corresponded to reduced survival, while breeding-season SSTs showed fewer significant effects on reproductive rates. Populations with dual responses to warming in both the breeding and nonbreeding seasons had the lowest projected population growth rates under future SSTs given a high emissions scenario. These results demonstrate that population vulnerability reflects the interaction between seabirds' year-round distributions and regional ocean warming. This underlines the need to integrate year-round tracking and long-term monitoring to inform conservation strategies and marine spatial planning to ensure climate-resilient marine ecosystems.

Migration is a common trait among many animals allowing the exploitation of spatiotemporally variable resources. It often implies high energetic costs to cover large distances, for example between breeding and wintering grounds. For flying or swimming animals, the adequate use of winds and currents can help reduce the associated energetic costs. Migratory seabirds are good models because they dwell in habitats characterized by strong winds while undertaking very long migrations. We tested the hypothesis that seabirds migrate through areas with favourable winds. To that end, we used the SEATRACK dataset, a multi‐colony geolocator tracking dataset, for two North Atlantic seabirds with contrasting flight capabilities, the black‐legged kittiwake Rissa tridactyla and the Atlantic puffin Fratercula arctica , and wind data from the ERA5 climate reanalysis model. Both species had on average positive wind support during migration. Their main migratory routes were similar and followed seasonally prevailing winds. The general migratory movement had a loop‐shape at the scale of the North Atlantic, with an autumn route (southward) along the east coast of Greenland, and a spring route (northward) closer to the British Isles. While migrating, both species had higher wind support in spring than in autumn. Kittiwakes migrated farther and benefited from higher wind support than puffins on average. The variation in wind conditions encountered while migrating was linked to the geographical location of the colonies. Generally, northernmost colonies had a better wind support in autumn while the southernmost colonies had a better wind support in spring, with some exceptions. Our study helps understanding how the physical environment shapes animal migration, which is crucial to further predict how migrants will be impacted by ongoing environmental changes.

Mercury (Hg) is a heterogeneously distributed toxicant affecting wildlife and human health. Yet, the spatial distribution of Hg remains poorly documented, especially in food webs, even though this knowledge is essential to assess large-scale risk of toxicity for the biota and human populations. Here, we used seabirds to assess, at an unprecedented population and geographic magnitude and high resolution, the spatial distribution of Hg in North Atlantic marine food webs. To this end, we combined tracking data of 837 seabirds from seven different species and 27 breeding colonies located across the North Atlantic and Atlantic Arctic together with Hg analyses in feathers representing individual seabird contamination based on their winter distribution. Our results highlight an east-west gradient in Hg concentrations with hot spots around southern Greenland and the east coast of Canada and a cold spot in the Barents and Kara Seas. We hypothesize that those gradients are influenced by eastern (Norwegian Atlantic Current and West Spitsbergen Current) and western (East Greenland Current) oceanic currents and melting of the Greenland Ice Sheet. By tracking spatial Hg contamination in marine ecosystems and through the identification of areas at risk of Hg toxicity, this study provides essential knowledge for international decisions about where the regulation of pollutants should be prioritized.

IntroductionOffshore wind energy development (OWED) has been identified as a major contributor to the aspired growth in Norwegian renewable energy production. Spatially explicit vulnerability assessments are necessary to select sites that minimize the harm to biodiversity, including seabird populations. Distributional data of seabirds in remote areas are scarce, and to identify vulnerable areas, species, and seasons it is necessary to combine data sets and knowledge from different sources.MethodsIn this study, we combined seabird tracking data, data from dedicated coastal and seabird at-sea surveys, and presence-only data from citizen science databases to develop habitat suitability maps for 55 seabird species in four seasons throughout the Norwegian exclusive economic zone; in total 1 million km2 in the Northeast Atlantic. The habitat suitability maps were combined with species-specific vulnerability indicators to yield maps of seabird vulnerability to offshore wind farms (OWFs). The resulting map product can be used to identify the relative vulnerability of areas prospected for OWED with respect to seabird collision and habitat displacement. More detailed assessments can be done by splitting the spatial indicators into seasonal and species-specific components.Results and discussionAssociated with higher diversity of seabirds near the coast, the cumulative vulnerability indicator showed a strong declining gradient from the coast to offshore waters while the differences in vulnerability between ocean areas and seasons were negligible. Although the present map product represents the best currently available knowledge, the indicators are associated with complex uncertainties related to known and unknown sampling biases. The indicators should therefore be used cautiously, they should be updated regularly as more data become available, and we recommend that more detailed environmental impact assessments based on dedicated seabird surveys, tracking of birds from potentially affected populations and population viability analyses are conducted in areas ultimately selected for OWED.

With the projected increases in shipping activity and hydrocarbon extraction globally, there is an increased risk of negative ecological impacts from oil pollution on the marine environment, including seabirds. Oil Vulnerability Indices (OVIs) are a common approach to assess seabird species vulnerability to oil pollution and to identify where species are most at risk, typically across regional spatial scales and for a relatively limited number of species. This approach generally requires comprehensive data on at-sea distributions and densities; however, for many regions, these data are limited. We present a simplified OVI to assess seabird species vulnerability to oil pollution. To create the spatial component of the OVI, we used a predictive foraging radius approach, using existing colony size and foraging range data, to project at-sea distributions of seabird populations during the breeding season. We demonstrate this approach over a large spatial scale, the eastern North Atlantic, which includes areas where seabird at-sea data are lacking. Our results reveal areas off west Greenland, Iceland, and Norway where seabirds are most vulnerable to oil pollution during the breeding season, largely driven by large colonies of auks (Alcidae). We also identify locations along the coast of mainland Norway, Iceland, and Scotland, where seabirds are particularly at risk to oil pollution associated with major shipping routes. Identifying areas where species are most at risk can help inform where, and which, measures should be put in place to mitigate the impacts of oil pollution, such as protecting and avoiding high risk areas, for example, through adopting dynamic Areas to be Avoided (ATBAs). Our simplified OVI combined with the predictive foraging radius approach can be adapted to other regions globally that lack seabird-at-sea distribution data, to other marine wildlife, and to assess risk from hydrocarbon extraction and other anthropogenic threats, including fishing activities and offshore renewable developments.

Abstract Although nesting in colonies can offer substantial reproductive benefits for many seabird species, increased visibility to predators remains a significant disadvantage for most colony‐breeders. To counteract this, some seabird species have evolved aggressive nest defense strategies to protect vulnerable eggs and chicks. Here, we used an experimental approach to test whether colony inhabitance by breeding gulls Larus spp. in western Norway impacts visitation rates of a native, mammalian predator, the Eurasian otter Lutra lutra during the breeding season. Camera traps were placed inside of and on the periphery of seabird colonies prior to the breeding season and left to run for one continuous year. Sighting frequency of otters on these cameras was compared to a control region free of gull nesting. We found that otter activity was significantly reduced in the colonies when gulls were incubating and rearing chicks, compared to time periods when gulls were building nests and absent from the colonies. Rhythmic activity patterns did not seem to be significantly impacted by the presence of gulls. This study provides clear evidence that certain colony‐nesting species can have a direct, negative impact on visitation rates of a native carnivore. Seasonal carnivore activity patterns are likely to be highly dependent on differing nesting strategies and level of nest defense by seabirds.

Two subspecies of the Great Cormorant breed in Norway, the continental Phalacrocorax carbo sinensis in the south, along the Skagerrak coast, and the marine P. c. carbo from central Norway and northwards. Here we review the information existing until 2017 on population status and trends, breeding performance and diet of these two subspecies in Norway. The most recent national population estimates are approximately 2500 (in 2012) and 19,000 (in 2012–2014) breeding pairs of sinensis and carbo, respectively. The sinensis population established itself in 1996 in Rogaland at the south-western tip of Norway, and in 1997 in Østfold close to the Swedish border; in both areas it increased for about ten years. Since then, the numbers have stabilised. For carbo, the population increased from 21,000 pairs in the early 1980s to 27,000 in 1995, and then decreased to the current number of 19,000 pairs. Significant annual variations in clutch size and reproductive output have been observed, but the drivers of these changes have not been identified. Unidentified gadoids and Atlantic Cod Gadus morhua were the most common prey of carbo, whereas inshore species such as Corkwing Wrasse Symphodus melops, Rockcook Centrolabrus exoletus, Goldsinny Wrasse Ctenolabrus rupestris and Black Goby Gobius niger were the most common prey in the eastern Skagerrak caught by sinensis. Carbo took very large numbers of 1–3-year-old gadoids during the year, and we cannot exclude the possibility this can have local effects on fish mortality rates.

Migratory seabirds are exposed to various pollutants throughout their annual cycle. Among them, mercury (Hg) is of particular concern given its large impact on animal health. Recent studies suggest that winter is a critical period for seabirds when contamination by Hg can be higher than at other times of year. However, individuals within and between species can have different migration strategies and winter distributions that could affect their exposure. Here, we combined multi-year individual tracking data and Hg measurements from 6 Arctic seabird species. We investigated whether inter-annual variations in individual winter contamination with Hg was related to seabird fidelity to a wintering site over years. First, our results show that Hg concentrations above the toxicity threshold (i.e. 5 µg g-1dry weight in feathers) were observed in variable proportions according to species (from 2% of northern fulmars to 37% of Brünnich’s guillemots). Second, individuals with high fidelity to a wintering ground had more similar Hg concentrations among years compared to individuals with low fidelity, suggesting an effect of their migratory strategy on Hg contamination. Further, we found that the directional change in wintering areas among years influenced seabird Hg contamination, highlighting an additional effect of seabirds’ winter distribution. More specifically, individuals migrating to the northwest direction of a previous wintering ground tended to be more contaminated compared to those moving to eastern directions. These results confirm spatial differences in Hg concentration throughout the North Atlantic-Arctic and an east-west gradient increase in Hg concentrations. Verifying this trend will require more large-scale ecotoxicological studies at smaller spatial resolution.

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.

Bird migration is commonly defined as a seasonal movement between breeding and non-breeding grounds. It generally involves relatively straight and directed large-scale movements, with a latitudinal change, and specific daily activity patterns comprising less or no foraging and more traveling time. Our main objective was to describe how this general definition applies to seabirds. We investigated migration characteristics of 6 pelagic seabird species (little auk Alle alle, Atlantic puffin Fratercula arctica, common guillemot Uria aalge, Brünnich’s guillemot U. lomvia, black-legged kittiwake Rissa tridactyla and northern fulmars Fulmarus glacialis). We analysed an extensive geolocator positional and saltwater immersion dataset from 29 colonies in the North-East Atlantic and across several years (2008-2019). We used a novel method to identify active migration periods based on segmentation of time series of track characteristics (latitude, longitude, net-squared displacement). Additionally, we used the saltwater immersion data of geolocators to infer bird activity. We found that the 6 species had, on average, 3 to 4 migration periods and 2 to 3 distinct stationary areas during the non-breeding season. On average, seabirds spent the winter at lower latitudes than their breeding colonies and followed specific migration routes rather than non-directionally dispersing from their colonies. Differences in daily activity patterns were small between migratory and stationary periods, suggesting that all species continued to forage and rest while migrating, engaging in a ‘fly-and-forage’ migratory strategy. We thereby demonstrate the importance of habitats visited during seabird migrations as those that are not just flown over, but which may be important for re-fuelling.

Aim In migratory species, individuals often use fixed and individual-specific migration strategies, which we term individual migration strategy fidelity (IMSF). Our goal was to test if guillemots have flexible or fixed individual migration strategies (i.e. IMSF), if this behaviour is consistent across large parts of the genus' range and if they were philopatric to geographical sites or a habitat feature. Location North Atlantic. Taxon Uria spp. Methods We quantified consistent individual differences in inter-annual spatial distribution and habitat occupied throughout the non-breeding period using a large geolocator tracking dataset of 729 adult seabirds breeding at 13 colonies across the Northeast Atlantic and repeatedly tracked up to 7 years over a 9-year period. Additionally, we used a similarity index to calculate relative fidelity to either geographical sites or habitats and linear mixed-effects models to assess persistence of spatial site fidelity over multiple years. Results Both guillemot species exhibited IMSF across a large part of the genus' range which persisted over multiple years. Individuals of both species and almost all colonies showed fidelity to geographical sites and not to specific habitats. Main conclusions Guillemots show IMSF that is best explained by site familiarity (fidelity to specific sites) rather than habitat specialization (fidelity to specific habitats). In the context of rapidly changing environments, favourable habitats may permanently shift locations and hence species displaying IMSF driven by site familiarity-such as the genus Uria-may not be able to adjust their migration strategies sufficiently fast to sustain individual fitness and ensure population persistence.

Identifying drivers of population trends in migratory species is difficult, as they can face many stressors while moving through different areas and environments during the annual cycle. To understand the potential of migrants to adjust to perturbations, it is critical to study the connection of different areas used by different populations during the annual cycle (i.e. migratory connectivity). Using a large-scale tracking data set of 662 individual seabirds from 2 sympatric auk meta-populations (common guillemots Uria aalge and Brünnich’s guillemots U. lomvia) breeding in 12 colonies throughout the Northeast Atlantic, we estimated migratory connectivity in seasonal space use as well as occupied environmental niches. We found strong migratory connectivity, within and between species. This was apparent through a combination of seasonal space use and occupied environmental niches. Brünnich’s guillemot populations grouped into 2 and common guillemot populations into 5 previously undescribed spatiotemporal clusters. Common guillemot populations clustered in accordance with the variable population trends exhibited by the species, while Brünnich’s guillemot populations are declining everywhere where known within the study area. Individuals from different breeding populations in both species were clustered in their space and environmental use, utilising only a fraction of the potential species-wide range. Further, space use varied among seasons, emphasising the variable constraints faced by both species during the different stages of their annual cycle. Our study highlights that considering spatiotemporal dynamics, not only in space but also in occupied environmental niches, improves our understanding of migratory connectivity and thus population vulnerability in the context of global change.

Tracking data of marine predators are increasingly used in marine spatial management. We developed a spatial data set with estimates of the monthly distribution of 6 pelagic seabird species breeding in the Northeast Atlantic. The data set was based on year-round global location sensor (GLS) tracking data of 2356 adult seabirds from 2006-2019 from a network of seabird colonies, data describing the physical environment and data on seabird population sizes. Tracking and environmental data were combined in monthly species distribution models (SDMs). Cross-validations were used to assess the transferability of models between years and breeding locations. The analyses showed that birds from colonies close to each other (<500 km apart) used the same nonbreeding habitats, while birds from distant colonies (>1000 km) used colony-specific and, in many cases, non-overlapping habitats. Based on these results, the SDM from the nearest model colony was used to predict the distribution of all seabird colonies lying within a species-specific cut-off distance (400-500 km). Uncertainties in the predictions were estimated by cluster bootstrap sampling. The resulting data set consisted of 4692 map layers, each layer predicting the densities of birds from a given species, colony and month across the North Atlantic. This data set represents the annual distribution of 23.5 million adult pelagic seabirds, or 87 % of the Northeast Atlantic breeding population of the study species. We show how the data set can be used in population and spatial management applications, including the detection of population-specific non-breeding habitats and identifying populations influenced by marine protected areas.

Seabirds are undergoing drastic declines globally and spend the non-breeding season at sea, making it challenging to study the drivers of their survival. Harsh weather and changes in climate conditions can have large impacts on seabird population dynamics through increased mortality. The intensity and persistence of extreme events are forecasted to increase with global warming. As shared conditions can induce population synchrony, multi-population studies of key demographic parameters are imperative to explore the influence of climate change. We used long-term mark-recapture data and position data to determine non-breeding stop-over areas of 5 Atlantic puffin (Fratercula arctica) populations over a latitudinal gradient in the Northeast Atlantic (56°11’-70°23’N). We investigated synchrony in adult survival in relation to shared stop-over areas. We quantified effects of extreme extra-tropical cyclones (ETCs) specific to populations’ stop-over areas and the North Atlantic Oscillation on adult survival. Populations with overlapping stop-over areas exhibited temporal synchrony in survival rates. Winter ETCs negatively influenced survival in 1 population, which was the one most exposed to extreme weather, but did not directly influence adult survival in the other 4 populations. Synchrony among populations with shared stop-over areas highlights the importance of these areas for adult survival, a key life-history rate. However, extreme weather was not identified as a driving factor for the majority of study populations. This suggests other factors in these areas, likely related to bottom-up trophic interactions, as environmental drivers of synchrony in the survival of Atlantic puffins.

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.''(1-3) During these, thousands of emaciated seabird carcasses are washed ashore along European and North American coasts. Winter cyclones can therefore shape seabird population dynamics(4,5) 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.(6) We performed the first ocean-basin scale assessment of cyclone exposure in a seabird community by coupling winter tracking data for similar to 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(7) 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.(8)

Mercury (Hg) is a natural trace element found in high concentrations in top predators, including Arctic seabirds. Most current knowledge about Hg concentrations in Arctic seabirds relates to exposure during the summer breeding period when researchers can easily access seabirds at colonies. However, the few studies focused on winter have shown higher Hg concentrations during the non-breeding period than breeding period in several tissues. Hence, improving knowledge about Hg exposure during the non-breeding period is crucial to understanding the threats and risks encountered by these species year-round. We used feathers of nine migratory alcid species occurring at high latitudes to study bird Hg exposure during both the breeding and non-breeding periods. Overall, Hg concentrations during the non-breeding period were similar to 3 times higher than during the breeding period. In addition, spatial differences were apparent within and between the Atlantic and Pacific regions. While Hg concentrations during the non-breeding period were similar to 9 times and similar to 3 times higher than during the breeding period for the West and East Atlantic respectively, Hg concentrations in the Pacific during the non-breeding period were only similar to 1.7 times higher than during the breeding period. In addition, individual Hg concentrations during the non-breeding period for most of the seabird colonies were above 5 mu g g(-1) dry weight (dw), which is considered to be the threshold at which deleterious effects are observed, suggesting that some breeding populations might be vulnerable to non-breeding Hg exposure. Since wintering area locations, and migration routes may influence seasonal Hg concentrations, it is crucial to improve our knowledge about spatial ecotoxicology to fully understand the risks associated with Hg contamination in Arctic seabirds. Crown Copyright (C) 2020 Published by Elsevier B.V. All rights reserved.

We explored the implications of reaching the Paris Agreement Objective of limiting global warming to <2°C for the future winter distribution of the North Atlantic seabird community. We predicted and quantified current and future winter habitats of five North Atlantic Ocean seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia and Rissa tridactyla) using tracking data for ~1500 individuals through resource selection functions based on mechanistic modeling of seabird energy requirements, and a dynamic bioclimate envelope model of seabird prey. Future winter distributions were predicted to shift with climate change, especially when global warming exceed 2°C under a “no mitigation” scenario, modifying seabird wintering hotspots in the North Atlantic Ocean. Our findings suggest that meeting Paris agreement objectives will limit changes in seabird selected habitat location and size in the North Atlantic Ocean during the 21st century. We thereby provide key information for the design of adaptive marine‐protected areas in a changing ocean.

Europe is commonly occupied by Black Kites of the nominal subspecies Milvus migrans migrans. In recent years, there has been an increase in the number of sightings of Black Kites with morphological features of Milvus migrans lineatus suggesting the spreading of M. m. lineatus and a subsequent hybridization zone between M. m. migrans and M. m. lineatus in a westerly direction from Siberia across continental Europe. Furthermore, it appears that the emergence of Black Kites with lineatus features in Europe coincides with new observations of wintering Black Kites in some areas of southern Europe.