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The intensifcation of coastal development poses potential threats for coastal seabirds, and understanding their habitat use is a key factor to guide conservation and management. In sub-arctic areas, black guillemots (Cepphus grylle) use coastal habitats year-round, which makes them vulnerable to the increasing human activities in these areas. In mainland Norway, one of the species’ strongholds, black guillemots are red-listed after substantial population declines. However, their fnescale foraging behaviour has received little attention to date. We collected and analysed GPS tracking data from adult black guillemots at three sites located over a latitudinal gradient of 250 km in central/northern Norway. Maximum foraging ranges of 33 km at Sklinna (65°12′N) for incubating birds, and 18 km at both Vega (65°34′N) and Sklinna for chick-rearing birds, are among the longest reported for this species. At all three sites, foraging probability was highest in shallow waters (

Density-dependent feedback is recognized as important regulatory mechanisms of population size. Considering the spatial scales over which such feedback operates has advanced our theoretical understanding of metapopulation dynamics. Yet, metapopulation models are rarely fit to time-series data and tend to omit details of the natural history and behavior of long-lived, highly mobile species such as colonial mammals and birds. Seabird metapopulations consist of breeding colonies that are connected across large spatial scales, within a heterogeneous marine environment that is increasingly affected by anthropogenic disturbance. Currently, we know little about the strength and spatial scale of density-dependent regulation and connectivity between colonies. Thus, many important seabird conservation and management decisions rely on outdated assumptions of closed populations that lack density-dependent regulation. We investigated metapopulation dynamics and connectivity in an exemplar seabird species, the Northern gannet (Morus bassanus), using more than a century of census data of breeding colonies distributed across the Northeast Atlantic. We developed and fitted these data to a novel hierarchical Bayesian state-space model, to compare increasingly complex scenarios of metapopulation regulation through lagged, local, regional, and global density dependence, as well as different mechanisms for immigration. Models with conspecific attraction fit the data better than the equipartitioning of immigrants. Considering local and regional density dependence jointly improved model fit slightly, but importantly, future colony size projections based on different mechanistic regulatory scenarios varied widely: a model with local and regional dynamics estimated a lower metapopulation capacity (645,655 Apparently Occupied Site [AOS]) and consequently higher present saturation (63%) than a model with local density dependence (1,367,352 AOS, 34%). Our findings suggest that metapopulation regulation in the gannet is more complex than traditionally assumed, and highlight the importance of using models that consider colony connectivity and regional dynamics for conservation management applications guided by precautionary principles. Our study advances our understanding of metapopulation dynamics in long-lived colonial species and our approach provides a template for the development of metapopulation models for colonially living birds and mammals. connectivity, conspecific attraction, dispersal, immigration, latent process, long-term population monitoring, marine conservation, metapopulation dynamics, Monte Carlo Markov Chain, Morus bassanus, Northern gannet, regulatory feedback

A recent study on the diving behaviour of European shags (Gulosus aristotelis (L.)) foraging in kelp forests off rocky coasts of Norway suggests surface durations are related only to the duration of the preceding dive, and hence are being used for respiratory recovery. These results contrast with earlier reports concerning shags foraging in highly tidal estuarine waters off the coast of Lundy Island, SW England, where there was a stronger relationship between dive durations and preceding pre-dive surface durations, suggesting the use of preparatory variation in oxygen loading. These two datasets were collected using different methods, and statistically analysed in quite different ways, so the contrasting results here could be due to different methodologies rather than the ecological differences between the two foraging environments. Here, we re-analyse the two datasets using similar statistical methods, and we confirm the contrasting results produced by the two datasets. We, therefore, conclude that shag breathing strategies do differ between these two marine environments, presumably reflecting adaptive facultatively plastic responses to differences in predictability of foraging dive durations. Off the Norwegian coast, unpredictable variation in the depth and availability of pelagic prey in complex environments may require more responsive post-dive respiratory recovery on the surface after each dive. In the more uniform English near-shore environment, however, pre-dive preparatory oxygen loading customised to match predictable dive-to- dive variation in benthic prey depths and foraging durations, may be more time and energy efficient. dive behaviour, dive physiology, dive preparation, dive recovery, foraging behaviour, foraging ecology, marginal value theorem

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. Environmental niche · Inter-population mixing · Large-scale spatiotemporal dynamics · Light-level geolocation · Murres · Population spread · Seasonality © The authors 2021. Open Access under Creative Commons by Attribution Licence. Use, distribution and reproduction are un - restricted. Authors and original publication must be credited.

Foraging dives in birds and mammals involve complex physiological and behavioural adaptations to cope with the breaks in normal respiration. Optimal dive strategies should maximize the proportion of time spent under water actively foraging versus the time spent on the surface. Oxygen loading and carbon dioxide dumping carried out on the surface could involve recovery from the consequences of the last dive and/or preparation in anticipation of the next dive depth and duration. However, few studies have properly explored the causal pattern of effects within such dive cycles, which is crucial prior to any assessment of optimal dive strategies. Using time depth recorders and global positioning system loggers, we recorded over 42 000 dives by 39 pairs of male and female European shags, Phalacrocorax aristotelis. Dives either involved a straight descent and ascent, presumably reflecting an unsuccessful search for prey, or a descent followed by horizontal movement followed by an ascent, presumably reflecting active hunting pursuit of pelagic prey. Males were larger than females, but we were unable to distinguish between sex effects and the nonlinear effects of body mass on dive behaviour. Path analysis showed that within-individual dive-to-dive variation in surface times can best be explained as recovery from the previous dive. As expected in a pelagic hunter with unpredictable dive durations, there was no evidence of anticipatory preparation of oxygen stores in predive surface durations. Among-individual variation in dives showed that body mass directly affected descent durations, but individual variation in all other dive and surface durations was driven by variation in descent duration, suggesting a critical role for dive depth in overcoming body mass-dependent effects of hydrodynamic/wave drag and buoyancy. Our analyses test for the first time certain critical assumptions for studies assessing optimal dive strategies in birds and mammals, thereby revealing new details and avenues for research concerning adaptive diving behaviour. body mass dive behaviour dive preparation dive recovery European shag foraging behaviour Phalacrocorax aristotelis sex differences TDR telemetry

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. Great Cormorant, Norway, sinensis, carbo, population changes, breeding performance, diet

Piscivorous wildlife is often perceived as competitors by humans. Great cormorants of the continental subspecies (Phalacrocorax carbo sinensis) in the Baltic and North Sea increase, while local cod (Gadus morhua) stocks decline. In contrast, numbers of the Atlantic subspecies (Phalacrocorax carbo carbo), breeding along the Norwegian and Barents Seas, have been relatively stable. We investigated the diet of both great cormorant subspecies in breeding colonies along the Norwegian Coast from Lofoten to the Skagerrak and estimated the biomass of fish consumed annually by great cormorants in Norwegian waters. The birds’ consumption was compared with estimated fish stock sizes and fishery catches. Cod and saithe (Pollachius virens) dominated the diet in the Norwegian Sea and wrasses in the North Sea and Skagerrak. Estimated total fish consumption of cod and saithe by great cormorants was

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

Climate warming is driving changes in species distributions, although many species show a so-called climatic debt, where their range shifts lag behind the fast shift in temperature isoclines. Protected areas (PAs) may impact the rate of distribution changes both positively and negatively. At the cold edges of species distributions, PAs can facilitate species distribution changes by increasing the colonization required for distribution change. At the warm edges, PAs can mitigate the loss of species, by reducing the local extinction of vulnerable species. To assess the importance of PAs to affect species distribution change, we evaluated the changes in a non-breeding waterbird community as a response to temperature increase and PA status, using changes of species occurrence in the Western-Palearctic over 25 years (97 species, 7,071 sites, 39 countries, 1993– 2017). We used a community temperature index (CTI) framework based on species thermal affinities to investigate the species turn-over induced by temperature increase. In addition, we measured whether the thermal community adjustment was led by cold-dwelling species extinction and/or warm-dwelling species colonization, by modelling the change in standard deviation of the CTI (CTIsd). Using linear mixed-effects models, we investigated whether communities within PAs had lower climatic debt and different patterns of community change regarding the local PA surface. Thanks to the combined use of the CTI and CTIsd, we found that communities inside PAs had more species, higher colonization, lower extinction and the climatic debt was 16% lower than outside PAs. The results suggest the importance of PAs to facilitate warm-dwelling species colonization and attenuate cold-dwelling species extinction. The community adjustment was however not sufficiently fast to keep pace with the strong temperature increase in central and northeastern Western-Palearctic regions. Our study underlines the potential of the combined CTI and CTIsd metrics to understand the colonization-extinction patterns driven by climate warming.

International audience; Abstract: Tracking data of marine predators are increasingly used in marine spatial management. We developed a spatial dataset with estimates of the monthly distribution of six pelagic seabird species breeding in the Northeast Atlantic. The dataset is 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 (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). The uncertainties in predictions were estimated by cluster bootstrap sampling. The resulting dataset consists of 4692 map layers, each layer predicting the densities of birds from a given species, colony and month across the North Atlantic. The dataset 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 dataset can be used in population and spatial management applications, including the detection of population-specific nonbreeding habitats and identifying populations influenced by marine protected areas.

Shore-based observation of the diving behaviour of European shags (and possibly other birds)

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.

Species breeding at high latitudes face a significant challenge of surviving the winter. Such conditions are particularly severe for diurnal marine endotherms such as seabirds. A critical question is therefore what behavioural strategies such species adopt to maximise survival probability. We tested 3 hypotheses: (1) they migrate to lower latitudes to exploit longer day length (‘sun-chasing’), (2) they forage at night (‘night-feeding’), or (3) they target high-quality food patches to minimise foraging time (‘feasting’). We studied the winter migration and foraging strategies of European shags Phalacrocorax aristotelis from 6 colonies across a latitudinal gradient from temperate regions to north of the Arctic Circle using geolocators deployed over 11 winters. We found evidence for ‘sun-chasing’, whereby average southerly movements were greatest from colonies at higher latitudes. However, a proportion of individuals from higher latitudes remained resident in winter and, in the absence of daylight, they foraged during twilight and only very occasionally during the night. At lower latitudes, there was little evidence that individuals migrated south, nocturnal feeding was absent, and twilight feeding was infrequent, suggesting that there was sufficient daylight in winter. There was no evidence that winter foraging time was lowest at higher latitudes, as predicted by the ‘feasting’ hypothesis. Our results suggest that shags adopt different behavioural strategies to survive the winter across their latitudinal range, dictated by the differing light constraints. Our study highlights the value of multi-colony studies in testing key hypotheses to explain population persistence in seabird species that occur over large latitudinal ranges.

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.

Coastal kelp forest ecosystems create dynamic and productive habitats, supporting a wide range of epiphytic flora, invertebrates, fish and seabirds. Worldwide, kelp is harvested commercially, affecting kelp-associated animal communities. There is, however, limited knowledge of how fish and seabird respond to kelp harvest, highlighting the need to evaluate the ecological impact of harvest on all ecosystem levels. Using 6 years of GPS-tracking data, we examined the effects of kelp harvest on foraging behaviour of breeding European shags (Phalacrocorax aristotelis) from a colony in central Norway. We determined the spatial overlap between kelp harvest and foraging areas of shags and assessed the immediate, short- and long-term impacts of harvest on shag foraging behaviour. Our results demonstrated large spatial and temporal overlap in areas used by foraging shags and kelp harvest. We could not detect any clear alterations in the diving activity of shags due to kelp harvest. However, the broad temporal and spatial scale of our study constrained the detection of fine scale changes in shag behaviour in response to kelp harvest. Our study, nonetheless, identifies several issues that should be addressed before concluding on the effects of kelp harvest on seabird populations. This includes the need for experimental studies using directed and controlled harvest to investigate the effects of kelp harvest through the different trophic levels, including top predators. This is essential for ecosystem-based management of coastal resources, considering the many species composed in the coastal ecosystem.

International audience; Aim: In migratory species, individuals often use fixed and individual-specific migra-tion 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 geo-graphical 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 colo-nies showed fidelity to geographical sites and not to specific habitats.Main conclusions: Guillemots show IMSF that is best explained by site familiarity (fi-delity 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.

Ship and boat traffic are increasing sources of disturbance to marine wildlife. During moult, sea ducks are flightless and rely on productive and shallow feeding areas. However, this period coincides with the peak of the recreational boating season. This is the first study to investigate the escape behaviour of moulting common eiders (Somateria mollissima) to the approach of small boats. We quantified flight initiation distances (flock-to-boat distance at which an energy-demanding escape occurred), displacement distances (distance between the pre- and post-disturbance position of the flock) and the time it took flocks to return to pre-disturbance (foraging- or resting-) behaviour. Moulting common eiders showed average flight initiation distances of 177 m and displacement distances of 771 m. Displacement distances decreased with flock size, under higher wind speeds and when previous foraging habitat was shallower. Time-to-return to pre-disturbance behaviour decreased with flock size but increased with wind speed and accessibility of foraging habitat at the previous location. Most (75%) of flocks returned to pre-disturbance behaviour within 10 min after the disturbance, while three flocks kept disturbed even 45 min after the approach. Finally, flocks encountered less accessible (deeper) habitats after disturbance than before. Our results suggest that approaching boats imply considerable disturbance effects for moulting common eiders through increased locomotion costs, displacement from accessible foraging habitat and/or time lost for foraging or resting. We provide valuable information for policy makers and marine spatial planning and highlight the need for awareness among recreational boat drivers on their impact on wildlife.

A global analysis recently showed that seabird breeding phenology (as the timing of egg-laying and hatching) does not, on average, respond to temperature changes or advance with time (Keogan et al. 2018 Nat. Clim. Change 8, 313–318). This group, the most threatened of all birds, is therefore prone to spatio-temporal mismatches with their food resources. Yet, other aspects of the breeding phenology may also have a marked influence on breeding success, such as the arrival date of adults at the breeding site following winter migration. Here, we used a large tracking dataset of two congeneric seabirds breeding in 14 colonies across 18° latitudes, to show that arrival date at the colony was highly variable between colonies and species (ranging 80 days) and advanced 1.4 days/year while timing of egg-laying remained unchanged, resulting in an increasing pre-laying duration between 2009 and 2018. Thus, we demonstrate that potentially not all components of seabird breeding phenology are insensitive to changing environmental conditions.

Aim: Many species are showing distribution shifts in response to environmental change. We explored (a) the effects of inter-annual variation in winter weather conditions on non-breeding distributional abundance of waterbirds exploiting different habitats (deep-water, shallow water, farmland) and (b) the long-term shift in the population centroid of these species and investigate its link to changes in weather conditions. Location: Europe. Methods: We fitted generalized additive mixed Models to a large-scale, 24-year dataset (1990–2013) describing the winter distributional abundance of 25 waterbird species. We calculated the annual and long-term (3-year periods) population centroid of each species and used the winter North Atlantic Oscillation (NAO) index to explain the inter-annual and long-term shifts in their location. Results: (a) Year-to-year southwestwards shifts in the population centroids of deep- and shallow-water species were linked to negative NAO values. Shallow-water species shifted northeastwards associated with positive NAO values and the distance shifted increased with increasing NAO. Deep-water species shifted northeastwards up to zero NAO indices, but showed no further increase at higher NAO values. (b) Deep-water species showed long-term northeastwards shifts in distributional abundance throughout the 1990s and the 2000s. Shallow-water species, on the other hand, shifted northeastwards during the 1990s and early 2000s, but southwestwards thereafter. There were no significant links between the NAO and year-to-year movements or long-term shifts in farmland species’ population centroid. Main Conclusions: We provide evidence for a link between both year-to-year and long-term changes in waterbird winter distributional abundances at large geographical scales to short- and long-term changes in winter weather conditions. We also show that species using shallow water, deep-water and farmland habitats responded differently, especially at high NAO values. As well as important ecological implications, these findings contribute to the development of future conservation measures for these species under current and future climate change. (Less)

Accepted manuscript. Final version published in Biology Letters, 15(10), is available at https://doi.org/10.1098/rsbl.2019.0634. A global analysis recently showed that seabird breeding phenology (as the timing of egg-laying and hatching) does not, on average, respond to temperature changes or advance with time (Keogan et al. 2018 Nat. Clim. Change 8, 313–318). This group, the most threatened of all birds, is therefore prone to spatio-temporal mismatches with their food resources. Yet, other aspects of the breeding phenology may also have a marked influence on breeding success, such as the arrival date of adults at the breeding site following winter migration. Here, we used a large tracking dataset of two congeneric seabirds breeding in 14 colonies across 18° latitudes, to show that arrival date at the colony was highly variable between colonies and species (ranging 80 days) and advanced 1.4 days/year while timing of egg-laying remained unchanged, resulting in an increasing pre-laying duration between 2009 and 2018. Thus, we demonstrate that potentially not all components of seabird breeding phenology are insensitive to changing environmental conditions.

Seabird populations have declined worldwide, and several of the potential threats are of anthropogenic origin. To understand how changes in seabird populations relate to environmental conditions it is important to know the functional relationships between prey availability and foraging behaviour, prey choice and breeding performance over several years. This was studied by linking breeding success of European shag Phalacrocorax aristotelis to variation in diet composition, and investigating the underlaying mechanism driving this variation, primarily based on be - havioural costs associated with foraging, such as foraging range and diving effort. We obtained demographic data and foraging trip metrics (using GPS-loggers and time−depth recorders) from a shag colony at Sklinna, Central Norway, during the 2011−2016 breeding seasons. Breeding population size was closely and positively correlated with breeding success, which in turn was positively correlated with the proportion of saithe Pollachius virens in the diet. When the dietary proportion of uncommon prey species increased, breeding success decreased. Breeding success was negatively influenced by increasing distance travelled and accumulated dive depths on an annual basis. Summed dive depths were greatest when prey species other than saithe dominated the diet. We found that in years with low availability of saithe fewer shags bred, and those that did had lower breeding success. This indicates that in years with poor feeding conditions, there might not be sufficient resources in the foraging area to support the whole breeding population of shags.

A global analysis recently showed that seabird breeding phenology (as the timing of egg-laying and hatching) does not, on average, respond to temperature changes or advance with time (Keogan et al. 2018 Nat. Clim. Change 8, 313–318). This group, the most threatened of all birds, is therefore prone to spatio-temporal mismatches with their food resources. Yet, other aspects of the breeding phenology may also have a marked influence on breeding success, such as the arrival date of adults at the breeding site following winter migration. Here, we used a large tracking dataset of two congeneric seabirds breeding in 14 colonies across 18° latitudes, to show that arrival date at the colony was highly variable between colonies and species (ranging 80 days) and advanced 1.4 days/year while timing of egg-laying remained unchanged, resulting in an increasing pre-laying duration between 2009 and 2018. Thus, we demonstrate that potentially not all components of seabird breeding phenology are insensitive to changing environmental conditions.

The timing of annual events such as reproduction is a critical component of how free-living organisms respond to ongoing climate change. This may be especially true in the Arctic, which is disproportionally impacted by climate warming. Here, we show that Arctic seabirds responded to climate change by moving the start of their reproduction earlier, coincident with an advancing onset of spring and that their response is phylogenetically and spatially structured. The phylogenetic signal is likely driven by seabird foraging behavior. Surface-feeding species advanced their reproduction in the last 35 years while diving species showed remarkably stable breeding timing. The earlier reproduction for Arctic surface-feeding birds was significant in the Pacific only, where spring advancement was most pronounced. In both the Atlantic and Pacific, seabirds with a long breeding season showed a greater response to the advancement of spring than seabirds with a short breeding season. Our results emphasize that spatial variation, phylogeny, and life history are important considerations in seabird phenological response to climate change and highlight the key role played by the species' foraging behavior.

Clausen, Preben/0000-0001-8986-294X WOS: 000536149100018 Migratory waterbirds require an effectively conserved cohesive network of wetland areas throughout their range and life-cycle. Under rapid climate change, protected area (PA) networks need to be able to accommodate climate-driven range shifts in wildlife if they are to continue to be effective in the future. Thus, we investigated geographical variation in the relationship between local temperature anomaly and the abundance of 61 waterbird species during the wintering season across Europe and North Africa during 1990-2015. We also compared the spatio-temporal effects on abundance of sites designated as PAs, Important Bird and Biodiversity Areas (IBAs), both, or neither designation (Unlisted). Waterbird abundance was positively correlated with temperature anomaly, with this pattern being strongest towards north and east Europe. Waterbird abundance was higher inside IBAs, whether they were legally protected or not. Trends in waterbird abundance were also consistently more positive inside both protected and unprotected IBAs across the whole study region, and were positive in Unlisted wetlands in southwestern Europe and North Africa. These results suggest that IBAs are important sites for wintering waterbirds, but also that populations are shifting to unprotected wetlands (some of which are IBAs). Such IBAs may therefore represent robust candidate sites to expand the network of legally protected wetlands under climate change in north-eastern Europe. These results underscore the need for monitoring to understand how the effectiveness of site networks is changing under climate change. Academy of FinlandAcademy of Finland [312579, 275606]; Tiina and Antti Herlin Foundation, Helsinki (Finland); Research Council of NorwayResearch Council of Norway [280952/E20]; project Future Bird Scenarios through the 2017-2018 Belmont Forum; Research Council of Norway (Forskningsradet)Research Council of Norway [NINA: 295767]; Academy of Finland (AKA, Univ. Helsinki) [326338]; Swedish Research Council (Formas, Lund Univ.) [2018-02441] We thank the thousands of volunteer birdwatchers that carry out the International Waterbird Census (IWC) across Europe and Africa. We also thank Wetlands International (www.wetlands.org) for coordinating such a large-scale monitoring scheme and for compiling all these data. We acknowledge all the funding sources that allow the continuation of the IWC. This work was supported by the Academy of Finland (projects 312579 and 275606), the Tiina and Antti Herlin Foundation, Helsinki (Finland), the Research Council of Norway (project 280952/E20, cofounded by Statnett and the Norwegian Water Resources and Energy Directorate). In addition, this study is framed within the project Future Bird Scenarios funded through the 2017-2018 Belmont Forum and BiodivERsA joint call for research proposals, under the BiodivScen ERANet COFUND programme, with the following funding organisations: the Research Council of Norway (Forskningsradet, NINA: 295767), the Academy of Finland (AKA, Univ. Helsinki: 326338), and the Swedish Research Council (Formas, Lund Univ.: 2018-02441).

To identify priorities for management of seabirds during the breeding season, it is important to understand the ecological mechanisms driving chick growth and survival. In this study, we examined the effects of diet and prevailing weather on the growth and survival of chicks of black-legged kittiwakes Rissa tridactyla over a 10 yr period at Anda, a seabird colony in Northern Norway. We show that across all years, there was a significant effect of diet composition delivered to chicks on their growth and survival. A higher proportion of sandeel Ammodytes spp. in the chick diet was associated with an increase in daily growth rates, a pattern that was especially pronounced for the youngest chick in 2-chick broods. A high proportion of mesopelagic fish in the chick diet was associated with a decrease in survival, again, especially for the youngest chick in 2-chick broods. Periods of strong southerly winds also led to reduced survival, probably linked to nests being washed down from the colony. Growth rates of kittiwake chicks were negatively affected by wind speed, likely due to adults having to work more in the exposed habitats in strong winds, causing a reduction in the amount of food supplied to the chicks. Our results emphasise the importance of conservation of specific marine habitats shown to be important foraging areas in ensuring the reproductive success of seabirds. This might prove increasingly important if future climate regimes make ecological conditions more challenging for seabirds. Foraging effort · Mesopelagic fish · Nestling development · Prey availability · Rissa tridactyla · Sandeel · Wind conditions © The authors 2018. Open Access under Creative Commons by Attribution Licence. Use, distribution and reproduction are un - restricted. Authors and original publication must be credited.

Reproductive timing in many taxa plays a key role in determining breeding productivity , and is often sensitive to climatic conditions . Current climate change may alter the timing of breeding at different rates across trophic levels, potentially resulting in temporal mismatch between the resource requirements of predators and their prey . This is of particular concern for highertrophic-level organisms, whose longer generation times confer a lower rate of evolutionary rescue than primary producers or consumers. However, the disconnection between studies of ecological change in marine systems makes it difficult to detect general changes in the timing of reproduction5 . Here, we use a comprehensive meta-analysis of 209 phenological time series from 145 breeding populations to show that, on average, seabird populations worldwide have not adjusted their breeding seasons over time (−0.020 days yr−1 ) or in response to sea surface temperature (SST) (−0.272 days °C−1 ) between 1952 and 2015. However, marked between-year variation in timing observed in resident species and some Pelecaniformes and Suliformes (cormorants, gannets and boobies) may imply that timing, in some cases, is affected by unmeasured environmental conditions. This limited temperature-mediated plasticity of reproductive timing in seabirds potentially makes these top predators highly vulnerable to future mismatch with lower-trophic-level resources Fil: Keogan, Katharine. University of Edinburgh; Reino Unido Fil: Daunt, Francis. Centre for Ecology & Hydrology; Reino Unido Fil: Wanless, Sarah. Centre for Ecology & Hydrology; Reino Unido Fil: Phillips, Richard A.. British Antarctic Survey; Reino Unido Fil: Walling, Craig A.. University of Edinburgh; Reino Unido Fil: Agnew, Philippa. Oamaru Blue Penguin Colony; . Oamaru Blue Penguin Colony; Nueva Zelanda Fil: Ainley, David G.. HT Harvey and Associates; Estados Unidos Fil: Anker-Nilssen, Tycho. Norwegian Institute For Nature Research; Noruega Fil: Ballard, Grant. Point Blue Conservation Science; Estados Unidos Fil: Barrett, Robert T.. Uit The Arctic University Of Norway; Noruega Fil: Barton, Kerry J.. Landcare Research; Nueva Zelanda Fil: Bech, Claus. Norges Teknisk-naturvitenskapelige Universitet; Noruega Fil: Becker, Peter. Institut Fur Vogelforschung "vogelwarte Helgoland"; Alemania Fil: Berglund, Per-Arvid. University of Agricultural Sciences. Institute of Marine Research, Department of Aquatic Resources; Suecia Fil: Bollache, Loïc. Universite de Bourgogne; Francia Fil: Bond, Alexander L.. Memorial University Of Newfoundland; Canadá Fil: Bouwhuis, Sandra. Institut Fur Vogelforschung "vogelwarte Helgoland"; Alemania Fil: Bradley, Russell W.. Point Blue Conservation Science; Estados Unidos Fil: Burr, Zofia M.. The University Centre In Svalbard; Noruega Fil: Camphuysen, Kees. University of Utrecht; Países Bajos Fil: Catry, Paulo. Ispa - Instituto Universitário; Portugal Fil: Chiaradia, Andre. Monash University; Australia. Phillip Island Nature Park; Australia Fil: Christensen-Dalsgaard, Signe. Norwegian Institute for Nature Research; Noruega Fil: Cuthbert, Richard. Royal Society For The Protection Of Birds; Reino Unido Fil: Dehnhard, Nina. Universiteit Antwerp; Bélgica Fil: Descamps, Sébastien. Norwegian Polar Institute; Noruega Fil: Diamond, Tony. University Of New Brunswick; Canadá Fil: Divoky, George. Friends of Cooper Island; Estados Unidos Fil: Drummond, Hugh. Universidad Nacional Autónoma de México; México Fil: Dugger, Katie M.. United States Geological Survey; Estados Unidos Fil: Dunn, Michael J.. British Antarctic Survey; Reino Unido Fil: Emmerson, Louise. Australian Antarctic Division; Australia Fil: Erikstad, Kjell Einar. Norwegian Polar Institute; Noruega Fil: Fort, Jérôme. Université de la Rochelle; Francia Fil: Fraser, William. Polar Oceans Research Group; Estados Unidos Fil: Genovart, Meritxell. Consejo Superior de Investigaciones Cientificas. Instituto Mediterraneo de Estudios Avanzados; España Fil: Gilg, Olivier. Universite de Bourgogne; Francia. Groupe de Recherches En Ecologie Arctique; Fil: González-Solís, Jacob. Universidad de Barcelona; España Fil: Granadeiro, José Pedro. Universidade de Lisboa; Portugal Fil: Grémillet, David. University Of Cape Town; Sudáfrica Fil: Hansen, Jannik. University Aarhus; Dinamarca Fil: Hanssen, Sveinn A.. Norwegian Institute For Nature Research; Noruega Fil: Harris, Mike. Centre for Ecology & Hydrology; Reino Unido Fil: Hedd, April. Environment Canada; Canadá Fil: Hinke, Jefferson. National Oceanic And Atmospheric Administration; Estados Unidos Fil: Igual, José Manuel. Consejo Superior de Investigaciones Cientificas. Instituto Mediterraneo de Estudios Avanzados; España Fil: Jahncke, Jaime. Point Blue Conservation Science; Estados Unidos Fil: Jones, Ian. Memorial University Of Newfoundland; Canadá Fil: Kappes, Peter J.. Oregon Cooperative Fish And Wildlife Research Unit; Estados Unidos Fil: Lang, Johannes. Groupe de Recherche en Ecologie Arctique; Francia Fil: Langset, Magdalene. Norwegian Institute For Nature Research; Noruega Fil: Lescroël, Amélie. Géosciences Montpellier; Francia Fil: Lorentsen, Svein-Håkon. Norwegian Institute For Nature Research; Noruega Fil: Lyver, Phil O'B.. Landcare Research; Nueva Zelanda Fil: Mallory, Mark. Acadia University; Canadá Fil: Moe, Børge. Norwegian Institute For Nature Research; Noruega Fil: Montevecchi, William A.. Memorial University Of Newfoundland; Canadá Fil: Monticelli, David. University Of Coimbra, Marine And Environmental Sciences Center; Portugal Fil: Mostello, Carolyn. Massachusetts Division Of Fisheries And Wildlife; Estados Unidos Fil: Newell, Mark. Centre for Ecology & Hydrology; Reino Unido Fil: Nicholson, Lisa. Halfmoon Biosciences; Australia Fil: Nisbet, Ian. I. C. T. Nisbet & Company; Estados Unidos Fil: Olsson, Olof. Stockholms Universitet; Suecia Fil: Oro, Daniel. Consejo Superior de Investigaciones Cientificas. Instituto Mediterraneo de Estudios Avanzados; España Fil: Pattison, Vivian. Laskeek Bay Conservation Society; Canadá Fil: Poisbleau, Maud. Universiteit Antwerp; Bélgica Fil: Pyk, Tanya. Deakin University; Australia Fil: Quintana, Flavio Roberto. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Ramos, Jaime A.. University Of Coimbra, Marine And Environmental Sciences Center; Portugal Fil: Ramos, Raül. Universidad de Barcelona; España Fil: Reiertsen, Tone Kirstin. Norwegian Polar Institute; Noruega Fil: Rodríguez, Cristina. Universidad Nacional Autónoma de México; México Fil: Ryan, Peter. University Of Cape Town; Sudáfrica Fil: Sanz-Aguilar, Ana. Consejo Superior de Investigaciones Cientificas. Instituto Mediterraneo de Estudios Avanzados; España Fil: Schmidt, Niels M.. University Aarhus; Dinamarca Fil: Shannon, Paula. National Audubon Society Seabird Restoration Program; Estados Unidos Fil: Sittler, Benoit. Groupe de Recherche en Ecologie Arctique; Francia Fil: Southwell, Colin. Australian Antarctic Division; Australia Fil: Surman, Christopher. Halfmoon Biosciences; Australia Fil: Svagelj, Walter Sergio. Universidad Nacional de Mar del Plata; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Mar del Plata. Instituto de Investigaciones Marinas y Costeras. Universidad Nacional de Mar del Plata. Facultad de Ciencias Exactas y Naturales. Instituto de Investigaciones Marinas y Costeras; Argentina Fil: Trivelpiece, Wayne. Bolinas; Estados Unidos Fil: Warzybok, Pete. Point Blue Conservation Science; Estados Unidos Fil: Watanuki, Yutaka. Hokkaido University; Japón Fil: Weimerskirch, Henri. Centre National de la Recherche Scientifique; Francia Fil: Wilson, Peter R.. Landcare Research; Nueva Zelanda Fil: Wood, Andrew G.. British Antarctic Survey; Reino Unido Fil: Phillimore, Albert B.. University of Edinburgh; Reino Unido Fil: Lewis, Sue. University of Edinburgh; Reino Unido

We review the current and future threats to duck populations that breed, stage, moult and/or winter in the Nordic countries. Migratory duck species are sensitive indicators of their changing environment, and their societal value confirms the need to translate signals from changes in their distribution, status and abundance into a better understanding of changes occurring in their wetland environments. We used expert opinion to highlight 25 major areas of anthropogenic change (and touch briefly on potential mitigation measures through nature restoration and reserve management projects) that we consider key issues likely to influence Nordic duck populations now and in the near future to stimulate debate, discussion and further research. We believe such reviews are essential in contributing to development of successful management policy as well as stimulating specific research to support the maintenance of duck species in favourable future conservation status in the face of multiple population pressures and drivers.

Polar Regions are experiencing environmental changes at unprecedented rates. These changes can spread throughout entire food webs from lower trophic levels to apex predators. As many top predators forage over large areas, these indirect effects may be associated with large-scale patterns of climate variability. Using global climate indices that are known to impact the Southern Ocean ecosystem (the El Niño Southern Oscillation and Antarctic Oscillation Indices) we assessed their efficacy to predict variation in the demographic parameters of Antarctic seabirds. First, we used a long-term dataset on adult survival (estimated from capture–mark–recapture data) and reproduction of Antarctic petrel Thalassoica antarctica, from the largest known breeding colony (Svarthamaren, Dronning Maud Land) and examined whether large-scale oceanographic fluctuations impact survival and reproduction. Second, we conducted an exhaustive literature review to determine whether the effects of large-scale environmental variability on Antarctic seabirds have a coherent fingerprint across the Antarctic continent and nearby islands. We found that most of the variation in both reproductive success, timing of hatching, and survival of Antarctic petrels can be accurately modeled using the two modes of large-scale climate variability in Antarctica. The literature review, combined with the results from our field study, suggests that while the anticipated trends in the global patterns of climatic variability will generally have detrimental effects on populations of top predators in the Southwest Atlantic, these conclusions cannot be extrapolated to all seabird populations in Antarctica without additional data.

Because of the fast rate of wind-energy development it will become a challenge to verify impacts on birdlife and construe ways to minimise these. Birds colliding with wind turbines are generally perceived as one of the major conflict issues for wind-energy development. Development of effective and practical measures to reduce bird mortality related to offshore and onshore wind energy is therefore paramount to avoid any delay in consenting processes. The expected efficacy of post-construction mitigation measures for wind-turbine induced avian mortality can be expected to be species-specific with regard to audible, optical and biomechanical constraints and options. Species-specific sensory faculties limit the ability to observe a wind turbine in a given circumstance. Their consequent cognitive perception may depend on the possibilities for associating wind turbines with risk, and discriminating these from other sources. Last but not least, perceived risks may only be evaded when their aerodynamic, locomotive physiology enables them to do so in due time. In order to be able to identify and construe functional mitigation measures these aspects need to be taken into account. Measures eliciting a series of intermittent strong stimuli that are variable in frequency may limit habituation effects; these should only be elicited specifically to mitigate imminent collision. Thus measures either adjusting turbine operation or warning/deterring birds approaching turbines are expected to be most functional. Warning signals may either be based on optical or audible stimuli; however, birds׳ hearing is inferior to humans while their visual acuity and temporal resolution is higher, but with great differences among species. Implementing effective mitigation measures could reduce the general level of conflicts with birdlife and thus enable both the development at new sites, at sites that have been declared having too high conflict levels, and utilise the wind resources better at specific sites without increasing the conflict levels.

Current and future climate alterations might impact ecological processes like timing of breeding. We used multivariate linear models to assess the importance of food availability and climate on timing of breeding and hatching brood size of European shags Phalacrocorax aristotelis in 2 colonies, Sklinna (65° N) and Røst (67° N), in the Norwegian Sea over 21 yr (1989 to 2009). Predictor variables were ICES abundance estimates of young saithe Pollachius virens, the staple food of shags in these colonies, the previous winter’s North Atlantic Oscillation index (wNAO), and sea surface and air temperatures. Annual population size was included to control for density-dependent effects. The best model for hatching date explained 15% (Sklinna) and 70% (Røst) of the variation and showed that shags bred earlier when 1 yr old saithe were more abundant. The model for Røst also included 2 yr old saithe and wNAO. The best model for hatching brood size explained 40% (Sklinna) and 48% (Røst) and included a positive effect of population size at both sites. The model for Røst also included a positive effect of 1 yr old saithe. Our study demonstrates that abundance of 1 yr old saithe is a key factor for the breeding performance of shags in both colonies, whereas the climate variables were of less importance. Delayed breeding and low brood size were not associated with a high population of breeders, indicating breeding performance was density-independent and that the positive relationship between breeding numbers and saithe abundance mainly reflected variation in non-breeding among established breeders. KEY WORDS: Phalacrocorax aristotelis · Norwegian Sea · Timing of breeding · Brood size · Pollachius virens · NAO

Norwegian Northern Gannet Morus bassanus populations exhibit contrasting trends on a regional scale, with several colony extinctions having occurred in recent decades. In an attempt to understand the ecological drivers of such variability, we tested whether resource availability is a factor limiting the current development of gannetries in the Lofoten/Vesteralen area. Between 2007 and 2010, we recorded arrival and departure times of breeding Northern Gannets from two colonies from regions showing contrasting population growth rates during the past two decades. We also recorded the duration of joint attendances by Northern Gannet parents at the nest, performed opportunistic diet sampling and counted numbers of occupied nests. Finally, we compiled ring recoveries over a 30-year period to assess inter-colony movements. Norwegian Gannet parents spent more time together, attending their chick, and performed shorter foraging trips than those in British and French colonies of similar size. This suggests that, despite some annual variations, their foraging effort was relatively low. Diet samples from both colonies mainly constituted fish of high energetic value, such as large herring Clupea harengus, mackerel Scomber scombrus, and saithe Pollachius virens, prey that are relatively abundant within the study area. Data from ringed birds revealed a northward movement of adults ringed as breeding birds and chicks from extinct Lofoten colonies that established a growing colony close to the North Cape. Recorded foraging features (trip duration, joint attendance and prey quality) during our study does not indicate food availability as a limiting factor explaining successive extinctions and re-colonisations of breeding sites in Lofoten/Vesteralen. White-tailed Eagles Haliaeetus albicilla are known to predate opportunistically on Northern Gannet adults or chicks and their populations are growing in the Lofoten area. Their potential impact on the Norwegian Northern Gannet population dynamics should be further investigated.

Impacts of anthropogenic activities on coastal seabirds might be extensive, especially in the breeding season. Identifying important foraging areas and associated habitats is important for a proper management of seabirds. To identify habitat characteristics driving the selection of foraging sites of breeding European shags Phalacrocorax aristotelis, this study used tracking data (GPS- and TDR-loggers), from 282 individual birds comprising 905 foraging trips and 27,303 dives with known locations, to create habitat selection models. To explore possible effects of regional differences in habitat on foraging behavior, the study was performed at two Norwegian colonies, Sklinna in the Norwegian Sea (65°N, 11°E) and Hornoya in the Barents Sea (70°N, 31°E), with distinct differences in seascape structure and habitat availability. Shags at Sklinna foraged further away from the colony than those at Hornoya but diving depth and duration were similar at the two colonies. In both colonies, sea depth was an important predictor of habitats selected by chick-rearing shags during foraging, with birds preferring shallow depths. At Sklinna, shags also selected for flat areas with high probability of kelp forest occurrence. There was no difference in trip length and duration between sexes, but males dived deeper than females in both colonies. This suggests that males and females might utilize different microhabitats within the same foraging area. The study discusses the application of habitat selection modeling to identify important foraging areas for coastal seabirds, and how this may contribute to the management, conservation and assessment of impacts of human activities.

For marine top predators like seabirds, the oceans represent a multitude of habitats regarding oceanographic conditions and food availability. Worldwide, these marine habitats are being altered by changes in climate and increased anthropogenic impact. This is causing a growing concern on how seabird populations might adapt to these changes. Understanding how seabird populations respond to fluctuating environmental conditions and to what extent behavioral flexibility can buffer variations in food availability can help predict how seabirds may cope with changes in the marine environment. Such knowledge is important to implement proper long-term conservation measures intended to protect marine predators. We explored behavioral flexibility in choice of foraging habitat of chick-rearing black-legged kittiwakes Rissa tridactyla during multiple years. By comparing foraging behavior of individuals from two colonies with large differences in oceanographic conditions and distances to predictable feeding areas at the Norwegian shelf break, we investigated how foraging decisions are related to intrinsic and extrinsic factors. We found that proximity to the shelf break determined which factors drove the decision to forage there. At the colony near the shelf break, time of departure from the colony and wind speed were most important in driving the choice of habitat. At the colony farther from the shelf break, the decision to forage there was driven by adult body condition. Birds furthermore adjusted foraging behavior metrics according to time of the day, weather conditions, body condition, and the age of the chicks. The study shows that kittiwakes have high degree of flexibility in their behavioral response to a variable marine environment, which might help them buffer changes in prey distribution around the colonies. The flexibility is, however, dependent on the availability of foraging habitats near the colony. central-place foragers, foraging decisions, GPS tracking, resource allocation, seabird This is an open access article under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in any medium, provided the original work is properly cited.© 2017 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.

This is the peer reviewed version of the following article: Descamps S, Anker‐Nilssen T, Barrett RT, et al. Circumpolar dynamics of a marine top‐predator track ocean warming rates. Glob Change Biol. 2017;23:3770–3780. https://doi.org/10.1111/gcb.13715, which has been published in final form at https://doi.org/10.1111/gcb.13715 . This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Self-Archiving. Global warming is a nonlinear process, and temperature may increase in a stepwise manner. Periods of abrupt warming can trigger persistent changes in the state of ecosystems, also called regime shifts. The responses of organisms to abrupt warming and associated regime shifts can be unlike responses to periods of slow or moderate change. Understanding of nonlinearity in the biological responses to climate warming is needed to assess the consequences of ongoing climate change. Here, we demonstrate that the population dynamics of a long-lived, wide-ranging marine predator are associated with changes in the rate of ocean warming. Data from 556 colonies of black-legged kittiwakes Rissa tridactyla distributed throughout its breeding range revealed that an abrupt warming of sea-surface temperature in the 1990s coincided with steep kittiwake population decline. Periods of moderate warming in sea temperatures did not seem to affect kittiwake dynamics. The rapid warming observed in the 1990s may have driven large-scale, circumpolar marine ecosystem shifts that strongly affected kittiwakes through bottom-up effects. Our study sheds light on the nonlinear response of a circumpolar seabird to large-scale changes in oceanographic conditions and indicates that marine top predators may be more sensitive to the rate of ocean warming rather than to warming itself.

The black-legged kittiwake Rissa tridactyla is a pelagic seabird whose population has recently declined in most parts of the North Atlantic and which is red-listed in most bordering countries. To investigate a possible cause for this decline, we analysed the population dynamics of 5 kittiwake colonies along the Norwegian coast, ranging from 62° to 71° N, over the last 20 to 35 yr. By quantifying the importance of sea surface temperatures (SST) in relevant areas of the North Atlantic, we tested the importance of climatic conditions throughout the populations’ annual cycles. We found no synchrony among colonies; however, SST affected population dynamics, explaining between 6% and 37% (average 18%) of the variation in annual population growth rate. While dynamics of the southerly colonies were mainly affected by winter conditions in the Grand Banks area, dynamics of the northernmost colonies were dominated by autumn conditions off Svalbard. Negative slopes indicated stronger population decline under warmer ocean conditions. Population dynamics were affected both via adult survival and offspring recruitment, as evidenced by the presence of unlagged effects as well as effects lagged by the age at recruitment. Finally, we performed population viability analyses taking into account the projected warming trends for the future. The median time to extinction of the Norwegian colonies was 52 to 181 yr without considering covariates; 45 to 94 yr when considering the effects of SST but ignoring future warming; and 10 to 48 yr when ocean warming, based on a ‘business as usual’ scenario, was taken into account. Global warming · Non-breeding distribution · Population dynamics · Population viability analysis · Rissa tridactyla · Sea surface temperature

In seasonal environments, organisms are expected to optimally schedule reproduction within an annual range of environmental conditions. Latitudinal gradients generate a range of seasonality to which we can expect adaptations to have evolved, and can be used to explore drivers of timing strategies across species’ distribution ranges. This study compares the timing of egg hatching in four seabird species (Atlantic puffin Fratercula arctica, black-legged kittiwake Rissa tridactyla, common guillemot Uria aalge, and Brünnich's guillemot U. lomvia) covering a subarctic to Arctic latitudinal gradient along the Norwegian coast to Svalbard (65–79°N). Hatching was significantly delayed by an estimated 1.7, 2.3, and 1.9 d per latitudinal degree for puffins, kittiwakes, and common guillemots, respectively, but was not delayed for Brünnich's guillemots. Hatching distributions revealed an increase in intra-annual breeding synchronicity along a latitudinal gradient for kittiwakes only, whereas the two guillemots exhibited high hatching synchronicity at all colonies. We used this large-scale, multispecies timing data series to discuss constraints, adaptations, and mechanisms affecting breeding timing, a necessary step to recognize risks to populations and predict future ecosystem change. Copyright: © 2016 Burr et al. This is an open access article under the terms of the Creative Commons Attribution License,which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

Colonial breeding is an evolutionary puzzle, as the benefits of breeding in high densities are still not fully explained. Although the dynamics of existing colonies are increasingly understood, few studies have addressed the initial formation of colonies, and empirical tests are rare. Using a high-resolution larval drift model, we here document that the distribution of seabird colonies along the Norwegian coast can be explained by variations in the availability and predictability of fish larvae. The modelled variability in concentration of fish larvae is, in turn, predicted by the topography of the continental shelf and coastline. The advection of fish larvae along the coast translates small-scale topographic characteristics into a macroecological pattern, viz. the spatial distribution of top-predator breeding sites. Our findings provide empirical corroboration of the hypothesis that seabird colonies are founded in locations that minimize travel distances between breeding and foraging locations, thereby enabling optimal foraging by central-place foragers., Seabirds breed in high density colonies, but the factors determining colony position aren't clear. Here, Sandvik et al. show that small-scale coastal topography is related to likely variation in fish larval abundance, which predicts the distribution of seabird colonies along the Norwegian coast.

International audience; Commercial fisheries may impact marine ecosystems and affect populations of predatorslike seabirds. In the Southern Ocean, there is an extensive fishery for Antarctic krill Euphausiasuperba that is projected to increase further. Comparing distribution and prey selectionof fishing operations versus predators is needed to predict fishery-related impacts on krilldependentpredators. In this context, it is important to consider not only predators breedingnear the fishing grounds but also the ones breeding far away and that disperse during thenon-breeding season where they may interact with fisheries. In this study, we first quantifiedthe overlap between the distribution of the Antarctic krill fisheries and the distribution of akrill dependent seabird, the Antarctic petrel Thalassoica antarctica, during both the breedingand non-breeding season. We tracked birds from the world biggest Antarctic petrel colony(Svarthamaren, Dronning Maud Land), located >1000 km from the main fishing areas, duringthree consecutive seasons. The overall spatial overlap between krill fisheries and Antarcticpetrels was limited but varied greatly among and within years, and was high in someperiods during the non-breeding season. In a second step, we described the length frequencydistribution of Antarctic krill consumed by Antarctic petrels, and compared thiswith results from fisheries, as well as from diet studies in other krill predators. Krill taken byAntarctic petrels did not differ in size from that taken by trawls or from krill taken by most Antarctickrill predators. Selectivity for specific Antarctic krill stages seems generally low in Antarcticpredators. Overall, our results show that competition between Antarctic petrels andkrill fisheries is currently likely negligible. However, if krill fisheries are to increase in thefuture, competition with the Antarctic petrel may occur, even with birds breeding thousandsof kilometers away.

The diet of breeding European Shag (Phalacrocamx aristotelis) was studied at Sklinna, one of the world's largest colonies, during 2007–2010. Regurgitated pellets were collected regularly throughout the chick rearing period and hard parts used to identify prey items to the lowest possible taxonomic level. The length and width of otoliths were used to estimate fish body length and mass. Shags' diet was varied and contained at least 17 different species of fish. Gadoid species, including saithe (Pollachius virens), cod (Gadus morhua) and Poor Cod (Trisopterus minutus), were the main prey and contributed approximately 90% of the diet during the study period. Saithe dominated the diet by numerical frequency, biomass and frequency of occurrence in all study years. Sandeels (Ammodytidae) contributed little to shag diet. The numerical frequency and percentage saithe biomass was lower in 2010 than in the other years. However, this was compensated by a higher frequency of cod and poor cod, making the tota...

Seabird movements outside the breeding season are generally poorly known, but can cover thousands of square km and a multitude of habitats, feeding conditions and potential threats. During the last decades, many seabird species in the North Atlantic have experienced large reductions in population size and breeding success, probably caused by reduced prey abundance caused by climate alterations and overfishing. One of these seabird species is the common guillemot. We used global location sensors (geolocators) to identify inter-breeding movements of 10 individuals breeding at Sklinna, a colony off the coast of Central Norway during July 2009–July 2010. All individuals moved northwards after breeding, and eight of them (80 %) entered the Barents Sea where they probably completed their moult. Three individuals moved southwards before the winter, but in total, half of the individuals stayed in the Barents Sea during winter. The other half wintered off the coast of Central Norway–Lofoten. The fact that all individuals moved northwards to winter was surprising as ringing recoveries suggest they also moves southwards (to the Skagerrak area) to winter. This suggests variation (individual or annual) in wintering movements and calls for a multi-year geolocator study at a number of colonies. Much of the area in the Barents Sea–Lofoten area is classified as vulnerable with respect to specific environmental pressures such as oil pollution and other anthropogenic factors, and the importance of the Barents Sea as a major wintering area for common guillemots from central Norway certainly has implications for the management authorities.

Aim: An understanding of the non-breeding distribution and ecology of migratory species is necessary for successful conservation. Many seabirds spend the non-breeding season far from land, and information on their distribution during this time is very limited. The black-legged kittiwake, Rissa tridactyla, is a widespread and numerous seabird in the North Atlantic and Pacific, but breeding populations throughout the Atlantic range have declined recently. To help understand the reasons for the declines, we tracked adults from colonies throughout the Atlantic range over the non-breeding season using light-based geolocation. Location: North Atlantic. Methods: Geolocation data loggers were deployed on breeding kittiwakes from 19 colonies in 2008 and 2009 and retrieved in 2009 and 2010. Data from 236 loggers were processed and plotted using GIS. Size and composition of wintering populations were estimated using information on breeding population size. Results: Most tracked birds spent the winter in the West Atlantic, between Newfoundland and the Mid-Atlantic Ridge, including in offshore, deep-water areas. Some birds (mainly local breeders) wintered in the North Sea and west of the British Isles. There was a large overlap in winter distributions of birds from different colonies, and colonies closer to each other showed larger overlap. We estimated that 80% of the 4.5 million adult kittiwakes in the Atlantic wintered west of the Mid-Atlantic Ridge, with only birds from Ireland and western Britain staying mainly on the European side. Main conclusions: The high degree of mixing in winter of kittiwakes breeding in various parts of the Atlantic range implies that the overall population could be sensitive to potentially deteriorating environmental conditions in the West Atlantic, e.g. owing to lack of food or pollution. Our approach to estimating the size and composition of wintering populations should contribute to improved management of birds faced with such challenges.

Coastal kelp forest ecosystems provide important habitats for a diverse assemblage of invertebrates, fish and marine top-predators such as seabirds and sea mammals. Although kelp is harvested industrially on a worldwide scale little is known about the multi-trophic consequences of this habitat removal. We investigated how kelp fisheries, which remove feeding and nursery grounds of coastal fish, influence local food webs and the availability of food to a marine top predator, the great cormorant (Phalacrocorax carbo). We conducted experimental harvesting of the canopy-forming kelp (Laminaria hyperborea) during a 3 year period (2001–2003) in an area at the coast of Central Norway while synoptically monitoring fish occurrence and cormorant foraging parameters. Our results demonstrate that cormorants preferentially foraged within kelp-forested areas and performed significantly more dives when feeding in harvested versus un-harvested areas suggesting lower foraging yield in the former case. In kelp areas that were newly harvested the number of small (