Long-term individual foraging site fidelitywhy some gannets don't change their spots. Ecology, 96(11), There may be differences between this version and the published version. You are advised to consult the publisher's version if you wish to cite from it.http://eprints.gla.ac.uk/113930/ shown that individual foraging site fidelity (IFSF -when individuals consistently forage in only a 19 small part of their population's home range) occurs in some colonial breeders. Short-term IFSF 20 could result from animals using a win-stay, lose-shift (WSLS) foraging strategy. Alternatively, it 21 may be a consequence of other forms of individual specialisation. Pelagic seabirds are colonial 22 central-place foragers, classically assumed to use flexible foraging strategies to target widely 23 2 dispersed, spatiotemporally patchy prey. However, tracking has shown that IFSF occurs in many 24 seabirds, although it is not known whether this persists across years. To test for long-term IFSF 25 and to examine alternative hypotheses concerning its cause, we repeatedly tracked 55 northern 26 gannets from a large colony in the North Sea within and across three successive breeding 27 seasons. Gannets foraged in neritic waters, predictably structured by tidal mixing and thermal 28 stratification but subject to stochastic wind-induced overturning. Both within and across years, 29 coarse to mesoscale (10s km) IFSF was significant but not absolute and foraging birds departed 30 the colony in individually consistent directions. Carbon stable isotope ratios in gannet blood 31 tissues were repeatable within years and nitrogen ratios were also repeatable across years, 32 suggesting long-term individual dietary specialisation. Individuals were also consistent across 33 years in habitat use with respect to relative sea surface temperature and in some dive metrics yet 34 none of these factors accounted for IFSF. Moreover, at the scale of weeks, IFSF did not decay 35 over time and the magnitude of IFSF across years was similar to that within years, suggesting 36 that IFSF is not primarily the result of WSLS foraging. Rather, we hypothesise that site 37 familiarity, accrued early in life, causes IFSF by canalising subsequent foraging decisions. 38Evidence from this and other studies suggests that IFSF may be common in colonial central-39 place foragers, with far-reaching consequences for our attempts to understand and conserve these 40 animals in a rapidly changing environment. 41 42

The use of biologging devices continues to increase with technological advances yielding remarkable ecological insights and generating new research questions. However, as devices develop and are deployed more widely, there is a need to update our knowledge of the potential ethical impacts to allow scientists to balance these against the knowledge gained. We employed a suite of phylogenetically controlled meta‐analyses on a dataset comprising more than 450 published effect sizes across 214 different studies to examine the effects of biologger tagging on five key traits in birds. Overall, we found small but significant negative effects of tagging on survival, reproduction, parental care. In addition, tagging was positively associated with foraging trip duration, but had no effect on body mass. Meta‐regressions revealed that flying style, migration distance and proportional tag mass were significant influences producing these deleterious effects, with attachment type and position additionally important covariates influencing survival‐ and reproduction‐based effect sizes. There was a positive correlation between the effects of tagging on survival and reproduction, highlighting that effects may be cumulative, with the full effects of tagging not necessarily apparent in studies focused on single traits. We discuss the tradeoff between these negative effects and the advances gained through the use of biologgers. Finally, given the number of studies from our initial literature search that lacked sufficient data for inclusion in analyses, we provide recommendations on the essential information that all biologging studies should report in order to facilitate future assessments of impacts on animals.

Shearwaters and petrels (hereafter petrels) are highly adapted seabirds that occur across all the world's oceans. Petrels are a threatened seabird group comprising 124 species. They have bet-hedging life histories typified by extended chick rearing periods, low fecundity, high adult survival, strong philopatry, monogamy and long-term mate fidelity and are thus vulnerable to change. Anthropogenic alterations on land and at sea have led to a poor conservation status of many petrels with 52 (42%) threatened

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

Mid-latitude (~30-60 o) seasonally stratifying shelf-seas support a high abundance and diversity of marine predators such as marine mammals and seabirds. However, anthropogenic activities and climate change impacts are driving changes in the distributions and population dynamics of these animals, with negative consequences for ecosystem functioning. Across mid-latitude shelf-seas, marine mammals and seabirds are known to forage at a number of oceanographic habitats that structure the spatio-temporal distributions of prey. Knowledge of these and the bio-physical mechanisms driving such associations are needed to improve marine management and policy. Here, we provide a concise and easily accessible guide for both researchers and managers of marine systems on the predominant oceanographic habitats that are favoured for foraging by marine mammals and seabirds across mid-latitude shelf seas. We (1) identify and describe key discrete physical features present across the continental shelf, working inshore from the shelf-edge to the shore line, (2) provide an overview of findings relating to associations between these habitats and marine mammals and

Background: State-space models are important tools for quality control and analysis of error-prone animal movement data. The near real-time (within 24 h) capability of the Argos satellite system can aid dynamic ocean management of human activities by informing when animals enter wind farms, shipping lanes, and other intensive use zones. This capability also facilitates the use of ocean observations from animal-borne sensors in operational ocean forecasting models. Such near real-time data provision requires rapid, reliable quality control to deal with error-prone Argos locations. Methods: We formulate a continuous-time state-space model to filter the three types of Argos location data (Least-Squares, Kalman filter, and Kalman smoother), accounting for irregular timing of observations. Our model is deliberately simple to ensure speed and reliability for automated, near real-time quality control of Argos location data. We validate the model by fitting to Argos locations collected from 61 individuals across 7 marine vertebrates and compare model-estimated locations to contemporaneous GPS locations. We then test assumptions that Argos Kalman filter/smoother error ellipses are unbiased, and that Argos Kalman smoother location accuracy cannot be improved by subsequent state-space modelling. Results: Estimation accuracy varied among species with Root Mean Squared Errors usually <5 km and these decreased with increasing data sampling rate and precision of Argos locations. Including a model parameter to inflate Argos error ellipse sizes in the north-south direction resulted in more accurate location estimates. Finally, in some cases the model appreciably improved the accuracy of the Argos Kalman smoother locations, which should not be possible if the smoother is using all available information. Conclusions: Our model provides quality-controlled locations from Argos Least-Squares or Kalman filter data with accuracy similar to or marginally better than Argos Kalman smoother data that are only available via fee-based reprocessing. Simplicity and ease of use make the model suitable both for automated quality control of near real-time Argos data and for manual use by researchers working with historical Argos data.

Search behavior is often used as a proxy for foraging effort within studies of animal movement, despite it being only one part of the foraging process, which also includes prey capture. While methods for validating prey capture exist, many studies rely solely on behavioral annotation of animal movement data to identify search and infer prey capture attempts. However, the degree to which search correlates with prey capture is largely untested. This study applied seven behavioral annotation methods to identify search behavior from GPS tracks of northern gannets (Morus bassanus), and compared outputs to the occurrence of dives recorded by simultaneously deployed time–depth recorders. We tested how behavioral annotation methods vary in their ability to identify search behavior leading to dive events. There was considerable variation in the number of dives occurring within search areas across methods. Hidden Markov models proved to be the most successful, with 81% of all dives occurring within areas identified as search. k‐Means clustering and first passage time had the highest rates of dives occurring outside identified search behavior. First passage time and hidden Markov models had the lowest rates of false positives, identifying fewer search areas with no dives. All behavioral annotation methods had advantages and drawbacks in terms of the complexity of analysis and ability to reflect prey capture events while minimizing the number of false positives and false negatives. We used these results, with consideration of analytical difficulty, to provide advice on the most appropriate methods for use where prey capture behavior is not available. This study highlights a need to critically assess and carefully choose a behavioral annotation method suitable for the research question being addressed, or resulting species management frameworks established.

Most of the Great Skuas (Stercorarius skua) breeding at Hermaness, Shetland, exhibit dietary specialization: a small proportion feed almost exclusively upon seabird prey, a small proportion feed as generalists, and most feed on fishery discards. We investigated the foraging dynamics, reproductive performance, and survival of Great Skuas that specialized in depredating other seabirds compared with those feeding predominantly on fish. Around half of the specialist bird predators defended combined breeding and feeding territories that included a section of seabird colony; the remainder of the predatory skuas foraged away from breeding territories. Specialist bird predators retained their feeding habit and, if present, feeding territory, across years. Time budgets revealed that specialist bird predators spent less time foraging than skuas feeding predominantly on fish. Results of radio-telemetry indicated that bird-specialist skuas have smaller home ranges than other birds. In a comparison of reproductive performance, specialist bird predators consistently hatched earlier among years. They also showed larger clutch volumes and improved chick condition, but these were subject to annual variations. Hatching success and fledging success for specialist bird predators and specialist fish predators were similar. Specialist bird predators showed similar annual survival compared with fish-feeders over the same period. Specializing as a bird predator may be limited to the best birds in the population, but their poorer than predicted breeding success reveals the need for further study into the relationship between diet and reproductive success in this species.Consecuencias Reproductivas para Individuos de Stercorarius skua que se Especializan como Depredadores de Aves MarinasResumen. La mayoría de los individuos de Stercorarius skua que se reproducen en Hermaness, Shetland, presentan una especialización de la dieta: una proporción pequeña se alimenta casi exclusivamente de aves marinas, otra proporción pequeña generalista y la mayoría se alimenta de desechos de pesqueros. Se investigó la dinámica de forrajeo, el desempeño reproductivo y la supervivencia de individuos de S. skua que se especializan en la depredación de otras aves marinas comparándolo con otros que se alimentan predominantemente de pescado. Cerca de la mitad de los depredadores especialistas de aves defienden de manera combinada territorios para reproducción y para alimentación que incluyen una sección de la colonia: el resto de los individuos depredadores forrajean lejos de los territorios de reproducción. Los aves depredadores especialistas de aves mantienen sus há bitos alimenticios y, en caso de presentarse, el territorio para alimentación, a través de los años. Los presupuestos de tiempo revelaron que los depredadores especialistas de aves pasan menos tiempo forrajeando que los que se alimentan predominantemente de pescado. Los resultados de radio-telemetría indicaron que las aves especialistas tienen rangos de hogar menores que otras aves. En una comparación de desempeño reproductivo, los depredadores especialistas de aves presentaron fechas de eclosión más tempranas, las cuales se repitieron a traves de los años. Así mismo, mostraron volúmenes de puesta más grandes y mejor condición de las crías, pero éstos estuvieron sujetos a variaciones anuales. El éxito de eclosión y el éxito de emplumamiento de los depredadores especialistas de aves y de los depredadores especialistas de pescado fueron similares. Los depredadores especialistas de aves mostraron una supervivencia anual similar a la de los que se alimentan de pescado durante el mismo periodo. La especialización como depredador de aves puede limitarse a las mejores aves de la población, pero el bajo éxito reproductivo encontrado, contrario a lo predicho, revela la necesidad de estudios adicionales sobre la relación entre la dieta y el éxito reproductivo en esta especie.