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Knowledge of seabird distributions plays a key role in seabird conservation and sustainable marine management, underpinning efforts to designate protected areas or assess the impact of human developments. Technological advances in animal tracking devices increasingly allow researchers to acquire information on the movement of birds from specific colonies. Nevertheless, most seabird colonies have not been subject to such tracking and another means must be found to assess their likely foraging distribution. Consequently, foraging range data collated and summarized across other tracking studies has often been used to estimate species-level foraging distances for use within applied settings. However, generic species-specific foraging ranges must be used with caution due to the amount of variation in seabird foraging behaviour at both the individual and colony level. Specifically, while current reviews of seabird foraging ranges provide summary estimates of maximum foraging range, they typically do not assess the extent of among-colony or among-individual variation around such estimates. To address this, we conducted a variance component analysis of the maximum distance reached from the breeding colony per foraging trip (foraging range) using multi-colony tracking datasets to estimate the degree of between-individual, between-year, and between-colony variation in foraging range in four UK breeding seabirds (Black-legged Kittiwake Rissa tridactyla, Common Guillemot Uria aalge, Razorbill Alca torda & European Shag Gulosus aristotelis). We also provide updated estimates of typical foraging ranges for each species and quantified the influence of breeding stage and colony size. Overall, between-colony variation was typically the largest variance component, explaining 20% - 30% of the observed variation in foraging range across the four species. Individual-level variation was also relatively large among Shags. In Kittiwake, Guillemot, and Shag, but not Razorbill, average forag

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.

The non-breeding season presents significant energetic challenges to birds that breed in temperate or polar regions, with clear implications for population dynamics. In seabirds, the environmental conditions at non-breeding sites drive food availability and the energetic cost of regulatory processes, resulting in variation in diet, behaviour and energetics; however, very few studies have attempted to understand if and how these aspects vary between populations. We investigated whether non-breeding location influenced diet, behaviour and energetics in the common guillemot Uria aalge. We studied guillemots from four UK breeding colonies, two located on the west coast of Scotland and two on the east. We quantified non-breeding distribution, foraging behaviour and activity budgets of 39 individuals from July to March, using geolocation–immersion loggers and time-depth recorders, and used feather stable isotope signatures to infer diet during the post-breeding moult. We calculated energy expenditure and investigated whether the peak (an indicator of the potential vulnerability to marine threats) varied between colonies. Individuals were spatially segregated according to the coastline they breed on, with west coast guillemots distributed off the west coast of the UK and east coast guillemots distributed off the east coast. Diet and behaviour were more similar in guillemots that shared a breeding coastline than those that did not, as west coast guillemots foraged at a lower trophic level, spent less time diving and engaged in more pelagic foraging than east coast guillemots. However, energy expenditure was remarkably similar between colonies, peaking during late February/early March, indicating that, during our study period, there was high synchrony between colonies in the timing of potential vulnerability to marine threats. Therefore, any anthropogenic changes that result in decreased food availability or increased energy expenditure during late winter may have greater im

Animal tracking has afforded insights into patterns of space use in numerous species and thereby informed area-based conservation planning. A crucial consideration when estimating spatial distributions from tracking data is whether the sample of tracked animals is representative of the wider population. However, it may also be important to track animals in multiple years to capture changes in distribution in response to varying environmental conditions. Using GPS-tracking data from 23 seabird species, we assessed the importance of multi-year sampling for identifying important sites for conservation during the chick-rearing period, when seabirds are most spatially constrained. We found a high degree of spatial overlap among distributions from different years in most species. Multi-year sampling often captured a significantly higher portion of reference distributions (based on all data for a population) than sampling in a single year. However, we estimated that data from a single year would on average miss only 5 % less of the full distribution of a population compared to equal-sized samples collected across three years (min: −0.3 %, max: 17.7 %, n = 23). Our results suggest a key consideration for identifying important sites from tracking data is whether enough individuals were tracked to provide a representative estimate of the population distribution during the sampling period, rather than that tracking necessarily take place in multiple years. By providing an unprecedented multi-species perspective on annual spatial consistency, this work has relevance for the application of tracking data to informing the conservation of seabirds.

Migration is a widespread strategy for escaping unfavourable conditions during winter, but the extent to which populations that segregate during the breeding season aggregate during the non-breeding season is poorly understood. Low non-breeding season aggregation may be associated with higher likelihood of overlap with threats, but with fewer populations affected, whereas high aggregation may result in a lower probability of exposure to threats, but higher overall severity. We investigated non-breeding distributions and extent of population aggregation in 2 sympatrically breeding auks. We deployed geolocation-immersion loggers on common guillemots Uria aalge and razorbills Alca torda at 11 colonies around the northern UK and tracked their movements across 2 non-breeding seasons (2017-18 and 2018-19). Using 290 guillemot and 135 razorbill tracks, we mapped population distributions of each species and compared population aggregation during key periods of the non-breeding season (post-breeding moult and mid-winter), observing clear interspecific differences. Razorbills were largely distributed in the North Sea, whereas guillemot distributions were spread throughout Scottish coastal waters and the North, Norwegian and Barents Seas. We found high levels of aggregation in razorbills and a strong tendency for colony-specific distributions in guillemots. Therefore, razorbills are predicted to have a lower likelihood of exposure to marine threats, but more severe potential impact due to the larger number of colonies affected. This interspecific difference may result in divergent population trajectories, despite the species sharing protection at their breeding sites. We highlight the importance of taking whole-year distributions into account in spatial planning to adequately protect migratory species.

Europe has set ambitious green energy targets, to which offshore renewable developments (ORDs) will make a significant contribution. Governments are legally required to deliver ORDs sustainably; however, they may have detrimental impacts on wildlife, especially those already experiencing declines due to climate change. Population viability analysis (PVA) is the standard method for forecasting population change in ORD assessments, but PVAs do not currently account for climate effects. We quantified climate effects on seabird breeding success for 8 UK species breeding in the North Sea. We assessed the potential for seabirds to mitigate climate-driven changes in breeding success by accessing wider resources through increased foraging ranges around colonies. We demonstrate strong links between breeding success and climate in 5 species. In 4 of these species, future climate projections indicated large declines in breeding success relative to current rates. Only one species was predicted to increase breeding success under future climate. In all 5 species, there was limited opportunity for species to increase breeding success by expanding foraging ranges to access more suitable future climatic conditions. Climate change will have significant ramifications for future breeding success of seabirds breeding in the North Sea, an area undergoing extensive and rapid offshore renewable energy development. We recommend 3 methods for including climate-driven changes to seabird breeding success within ORD assessments: development of predictive climate-driven habitat use models to estimate ORD-wildlife interactions; delivery of a new ORD assessment framework that includes dynamic predictions of climate-driven habitat use and demography of wildlife populations; and consideration of climate-driven changes in the implementation of compensatory measures.

1. The ability of individual animals to balance their energy budgets throughout the annual cycle is important for their survival, reproduction and population dynamics. However, the annual cycles of many wild, mobile animals are difficult to observe and our understanding of how individuals balance their energy budgets throughout the year therefore remains poor. 2. We developed a hierarchical Bayesian state-space model to investigate how key components of animal energy budgets (namely individual energy gain and storage) varied in space and time. Our model used biologger-derived estimates of time-activity budgets, locations and energy expenditure to infer year-round time series of energy income and reserves. The model accounted for seasonality in environmental drivers such as sea surface temperature and daylength, allowing us to identify times and locations of high energy gain. 3. Our study system was a population of common guillemots Uria aalge breeding at a western North Sea colony. These seabirds manage their energy budgets by adjusting their behaviour and accumulating fat reserves. However, typically during severe weather conditions, birds can experience an energy deficit over a sustained period, leading to starvation and large-scale mortality events. 4. We show that guillemot energy gain varied in both time and space. Estimates of guillemot body mass varied throughout the annual cycle and birds periodically experienced losses in mass. Mass losses were likely to have either been adaptive, or due to energetic bottlenecks, the latter leading to increased susceptibility to mortality. Guillemots tended to be lighter towards the edge of their spatial distribution. 5. We describe a framework that combines biologging data, time-activity budget analysis and Bayesian state-space modelling to identify times and locations of high energetic reward or potential energetic bottlenecks in a wild animal population. Our approach can be extended to address ecological and conservation

1. Understanding how ecological processes combine to shape population dynamics is crucial in a rapidly changing world. Evidence has been emerging for how fundamental drivers of density dependence in mobile species are related to two differing types of environmental variation—temporal variation in climate, and spatiotemporal variation in food resources. However, to date, tests of these hypotheses have been largely restricted to mid-trophic species in terrestrial environments and thus their general applicability remains unknown. 2. We tested if these same processes can be identified in marine upper trophic level species. We assembled a multi-decadal data set on population abundance of 10 species of colonial seabirds comprising a large component of the UK breeding seabird biomass, and covering diverse phylogenies, life histories and foraging behaviours. 3. We tested for evidence of density dependence in population growth rates using discrete time state-space population models fit to long time-series of observations of abundance at seabird breeding colonies. We then assessed if the strength of density dependence in population growth rates was exacerbated by temporal variation in climate (sea temperature and swell height), and attenuated by spatiotemporal variation in prey resources (productivity and tidal fronts). 4. The majority of species showed patterns consistent with temporal variation in climate acting to strengthen density dependent feedbacks to population growth. However, fewer species showed evidence for a weakening of density dependence with increasing spatiotemporal variation in prey resources. 5. Our findings extend this emerging theory for how different sources of environmental variation may shape the dynamics and regulation of animal populations, demonstrating its role in upper trophic marine species. We show that environmental variation leaves a signal in long-term population dynamics of seabirds with potentially important consequences for their demograph

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.

Capsule: A data-thinning approach was used to assess the effects of reducing the frequency of nest-checks on estimates of breeding success of Common Guillemots Uria aalge. Inter-year and inter-colony differences in fledging age and their implications for setting a minimum age after which a chick could be assumed to have fledged were evaluated. Aims: To assess the consequences of reducing sampling frequency on the estimation of breeding success, and on the robustness of the assumption that breeding has been successful if a chick survives to 15 days old. Methods: Breeding success, ages at fledging and loss of chicks were estimated from daily checks at two Scottish colonies over a six-year period. Data-thinning was used to assess the consequences of reducing checks from daily to every two or three days. Breeding success was recalculated assuming that all chicks surviving to 15 days fledged. Results: Reducing the frequency of checks from daily to every second or third day resulted in a small but statistically significant increase in the estimate of success. Between 20% and 25% of chick losses occurred when chicks were ≥15 days old. Assuming that these chicks had fledged resulted in significant increases in breeding success. Conclusion: Assumptions about whether or not a chick fledged had a greater impact on the estimate of breeding success than reducing the frequency of nest-checks from daily to every second or third day. There was no threshold age after which a chick could be assumed to have fledged. These findings are relevant to other monitoring schemes where there is pressure to reduce input. Sampling methods used in monitoring schemes need to be clearly stated and changes in protocols documented so that sampling effects can be incorporated into future analyses.

Marine spatial planning aims to deliver sustainable use of marine resources by minimizing environmental impacts of human activities and designating Marine Protected Areas. This poses a challenge where species’ distributions show spatio-temporal heterogeneity. However, due to logistic constraints and challenging timescales many studies of distribution are undertaken over few years or on a restricted subset of the population. Long-term studies can help identify the degree of uncertainty in those less comprehensive in space and time. We quantify inter-annual and sub-colony variation in the summer foraging distribution of a population of European shags Phalacrocorax aristotelis, using a tracking data set comprising 320 individuals and 1106 foraging trips in 15 years from 1987 to 2010. Foraging distribution over the study period was concentrated in three areas. Data from one and two years captured an average of 54% and 64% of this distribution, respectively, but it required 8 years’ data to capture over 90% of the distribution. Foraging range increased with population size when breeding success was low, suggesting interplay between extrinsic and intrinsic effects. Furthermore, females had foraging ranges on average 36% greater than males. Finally, sub-colony segregation occurred in foraging areas up to 4 km from the colony and in the most distant locations (>10 km), whilst there was considerable overlap at intermediate distances (6–10 km). Our study highlights important considerations for marine spatial planning in particular, and species conservation in general, notably the proportion of the population distribution identified, the prevailing conditions experienced and the need for balanced sampling across the population.

Miniature geolocator loggers (Global Location Sensing, GLS) that provide daily locations of birds have revolutionised the study of winter ecology and migration patterns of seabirds. A long-term study of ringing recoveries and analyses of heavy metals and pollutants in tissues of Atlantic puffins Fratercula arctica from the Isle of May, south-east Scotland, suggested that this population wintered mainly within the North Sea. However, deployment of GLS devices over the 2007/2008 winter showed that many breeding birds made major excursions into the east Atlantic. This winter was the second of two when survival was extremely low (survival in 2006/2007 and 2007/2008 was 0.696 and 0.695, respectively, compared to the average of 0.922 over the period 1984/1985–2005/2006). These low rates of survival suggested that the unexpected use of the Atlantic might have been associated with unusually poor conditions in the North Sea as indicated by very low breeding success in 2007. Survival rate returned to previous levels in 2008/2009 providing the opportunity to test whether higher survival was associated with birds remaining in the North Sea, or whether movements into the Atlantic are a feature of this population unrelated to survival. Accordingly, geolocators were deployed over the 2009/2010 winter when adult survival was subsequently established to be high (0.913). We found greater support for the hypothesis that winter distribution is not associated with survival. Thus, 8 (40 %) of 20 individuals followed in 2009/2010 went into the Atlantic, a rate not significantly different from 11 (58 %) of the 19 followed in the 2007/2008 winter. Indeed, birds actually spent longer in the Atlantic and used a wider variety of areas in 2009/2010, although the time spent away from the colony was significantly shorter than in 2007/2008. Since our data were from individuals that survived, remaining in or moving out of the North Sea can both be successful strategies during winters when the popul

Large areas of sea around Britain have been identified as potential sites for renewable energy development, heightening the need for information about important areas for seabirds. The Atlantic Puffin Fratercula arctica is one of the commonest seabirds in northeast Britain yet little is known about where individuals feed. We used back-mounted GPS loggers to track Atlantic Puffins breeding at a colony close to where wind farms are proposed. During chick rearing, birds made two types of feeding trip: long absences that included an overnight stay at distant (38–66 km) feeding areas and short daytime excursions to areas much nearer the colony (9–17 km). There was considerable overlap of the distant feeding area with the proposed wind farms. However, Atlantic Puffins are known to be sensitive to disturbance and comparison of individuals with and without loggers showed that the birds' behaviour had been disrupted by some aspect of the procedure. While the areas used by the birds carrying GPS loggers accorded with expectations based on other methods, it is possible that results from these birds represent a worst-case scenario and overestimate the degree of overlap with the proposed wind farms.

In species that breed and winter in different areas, migration is a key period of the annual cycle with important effects on subsequent events in life. In order to investigate relationships between migration and other events in the annual cycle or environmental factors, we first need to acquire a basic knowledge of the dynamics (pattern and timing) of migration of various species. Our current knowledge of migration dynamics of European passerine birds is based mainly on studies of populations passing via the Western flyway. In contrast, relatively little is known about populations migrating via the Eastern flyway. We examined the seasonal pattern and timing of migration of several abundant passerine species flying across the eastern part of the Balkan Peninsula. We tested whether these were consistent between years within species and also among species within two ecological groups (long- and short-distance migrants). For most species the dynamics of migration were consistent between years. The pattern of migration was consistent among species of long- and of short-distance migrants. The two groups differed in timing of passage in autumn, with long-distance migrants passing significantly earlier than short distance ones.

This report summarises the results of a Knowledge Transfer Research Project that was undertaken by Dr Jane Reid (University of Aberdeen), Professor Pat Monaghan, (University of Glasgow), Dr Eric and Mrs Sue Bignal (Scottish Chough Study Group) and Dr Davy McCracken (Scottish Agricultural College). Dr Maria Bogdanova was employed as the postdoctoral research assistant on the project. The work was carried out in partnership with Scottish Natural Heritage (SNH) and the Royal Society for the Protection of Birds (RSPB). Funding was provided by a Knowledge Transfer Grant from the Natural Environment Research Council (NERC, PIs P. Monaghan & J. Reid), with matching partnership funding and in-kind support from SNH and RSPB. The overall aims of the project were to develop the scientific understanding of the population ecology of choughs on Islay, and to use this understanding to inform the development of appropriate conservation strategies and policies. The project built on existing long-term research on Islay’s choughs. It involved further analysis of long-term data, plus two years of intensive fieldwork designed to answer specific questions. The work aimed primarily to understand the ecology of choughs in their sub-adult years (ie, from fledging to breeding age). Survival from fledging to breeding is a key factor in causing population change. However, relatively little was previously known about the behaviour and ecology of choughs during this time. This report provides an overview of the results of the scientific study and focuses on presenting the scientific evidence on which resulting recommendations for chough conservation management on Islay are based. The report is written with the intention of presenting the results of the data analyses, and the rationale underlying those analyses, in a way that is accessible to non-specialists. Further details of analyses and technicalities are provided in published, peer-reviewed papers and/or are available on request. The report