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Genetic diversity can affect population viability and can be reduced by both acute and chronic mechanisms. Using the history of the establishment and management of two invasive rat species on Tetiaroa atoll, French Polynesia, we investigated the intensity and longevity of contrasting population bottleneck mechanisms on genetic diversity and bottleneck signal. Using microsatellite loci we show how both a chronic reduction over approximately 50 years of a Rattus exulans population caused by the arrival of its competitor R. rattus, and an acute reduction in a R. rattus population caused by a failed eradication approximately 10 years ago, caused similar magnitudes of genetic diversity loss. Furthermore, these strong bottleneck signals were in addition to the lasting signal from initial colonisation by each species many decades to centuries earlier, characterising a genetic paradox of biological invasion. These findings have implications for the study of population genetics of invasive species, and underscore how important historical context of population dynamics is when interpreting snapshots of genetic diversity.

All data presented are available via the invacost R package. Related code and supplementary data available at github.com/cjabradshaw/InvaCostVersionTrends.; International audience; Biological invasions are a global challenge that has received insufficient attention. Recently available cost syntheses have provided policy and decision makers with reliable and up-to-date information on the economic impacts of invasive alien species, aiming to motivate effective management. The resultant InvaCost database is now publicly and freely accessible and enables rapid extraction of monetary cost information globally. This has facilitated knowledge sharing, developed a more integrated and multidisciplinary network of researchers, and forged multidisciplinary collaborations among diverse organisations and stakeholders. Over 50 scientific publications so far have used the database and provided detailed assessments of invasion costs across geographic, taxonomic, and spatio-temporal scales. These studies have been instrumental in guiding policy and legislative decisions, while attracting public and media attention. We provide an overview of the improved availability, reliability, and defragmentation of monetary costs, how this has enhanced invasion science as a discipline, and outline directions for future development

Biological invasions represent a key threat to insular systems and have pronounced impacts across environments and economies. The ecological impacts have received substantial focus, but the socioeconomic impacts are poorly synthesized across spatial and temporal scales. We used the InvaCost database, the most comprehensive assessment of published economic costs of invasive species, to assess economic impacts on islands worldwide. We analyzed socioeconomic costs across differing expenditure types and examined temporal trends across islands that differ in their political geography-island nation states, overseas territories, and islands of continental countries. Over US$36 billion in total costs (including damages and management) has occurred on islands from 1965 to 2020 due to invasive species' impacts. Nation states incurred the greatest total and management costs, and islands of continental countries incurred costs of similar magnitude, both far higher than those in overseas territories. Damage-loss costs were significantly lower, but with qualitatively similar patterns across differing political geographies. The predominance of management spending differs from the pattern found for most countries examined and suggests important knowledge gaps in the extent of many damage-related socioeconomic impacts. Nation states spent the greatest proportion of their gross domestic products countering these costs, at least 1 order of magnitude higher than other locations. Most costs were borne by authorities and stakeholders, demonstrating the key role of governmental and nongovernmental bodies in addressing island invasions. Temporal trends revealed cost increases across all island types, potentially reflecting efforts to tackle invasive species at larger, more socially complex scales. Nevertheless, the already high total economic costs of island invasions substantiate the role of biosecurity in reducing and preventing invasive species arrivals to reduce strains on limited financial resources and avoid threats to sustainable development goals.Costos económicos de proteger a las islas de las especies invasoras Resumen Las invasiones biológicas representan una amenaza importante para los sistemas insulares, además de tener impactos pronunciados en el ambiente y en la economía. Los impactos ecológicos han recibido atención sustancial, mientras que los impactos socioeconómicos se encuentran pobremente sintetizados en las escalas temporales y espaciales. Usamos la base de datos InvaCost, el análisis más completo de los costos económicos de las especies invasoras, para evaluar los impactos económicos sobre las islas a nivel mundial. Analizamos los costos socioeconómicos en varios tipos de gastos y examinamos las tendencias temporales en las islas que difieren en su geografía política - islas estado-nación, territorios ultramarinos e islas de países continentales. En las islas han ocurrido gastos de más de $36 mil millones de dólares entre 1965 y 2020 debido a los impactos de las especies invasoras. Las islas estado-nación produjeron los mayores costos de manejo y el mayor total, mientras que las islas de los países continentales produjeron costos de una magnitud similar, ambas con gastos mucho más elevados que los de los territorios ultramarinos. Los costos de las pérdidas por daños fueron significativamente más bajas, aunque con patrones cualitativamente similares entre las diferentes geografías políticas. El predominio del gasto en el manejo difiere del patrón hallado en la mayoría de los países analizados y sugiere que hay vacíos importantes en el conocimiento del alcance de muchos de los impactos socioeconómicos relacionados con los daños. Las islas estado-nación gastaron la mayor proporción de su producto interno bruto en contrarrestar estos costos, al menos una orden de magnitud mayor que las otras localidades. La mayoría de los costos fueron asumidos por las autoridades y los accionistas, lo que demuestra el papel clave que tienen los organismos gubernamentales y no gubernamentales en cómo se atienden las invasiones insulares. Las tendencias temporales revelaron incrementos en el costo en todos los tipos de islas, lo que potencialmente refleja los esfuerzos por combatir a las especies invasoras a escalas más grandes y socialmente más complejas. Aun así, el elevado costo económico total de las invasiones insulares fundamenta la función que tiene la bioseguridad en la reducción y prevención de la llegada de especies invasoras para reducir presiones sobre los recursos financieros limitados y evitar amenazas para las metas de desarrollo sustentable.生物入侵是岛屿系统面临的主要威胁, 对环境和经济都产生了明显的影响。生态方面的影响已得到很多关注, 但社会经济方面的影响仍缺少时空尺度上的综合分析。本研究使用 InvaCost 数据库, 包含已发表的入侵物种经济成本最全面的评估数据, 来评估全球岛屿在生物入侵中面临的经济影响。我们分析了不同支出类型的社会经济成本, 并研究了不同政治地理的岛屿 (岛屿民族国家、海外领地和大陆国家的岛屿) 中社会经济成本随时间变化的趋势。结果显示, 从 1965 年到 2020 年, 岛屿由于入侵物种的影响产生的总成本 (包括破坏和管理) 超过 360 亿美元。民族国家的总成本和管理成本最高, 大陆国家的岛屿产生的成本规模相近, 都远远高于海外领地的成本。破坏导致的成本明显较低, 在不同政治地理的岛屿中有相似的情况。而管理支出占主导地位的情况则在我们研究的大多数国家中有所不同, 表明在许多与破坏有关的社会经济影响程度方面仍存在重要的知识空缺。民族国家用于这些成本的国内生产总值比例最高, 至少比其他地区高一个数量级。大多数费用由当局和利益相关者承担, 反映了政府和非政府机构在应对岛屿入侵问题中的关键作用。时间趋势分析表明, 所有类型的岛屿承担的成本都随时间推移而增加, 可能反映了在更大、更复杂的社会范围内应对生物入侵的努力。然而, 岛屿入侵问题的总经济成本已经很高, 这强调了应关注生物安全来减少和预防入侵物种的引入, 以减缓对有限财政资源的压力, 并避免对可持续发展目标的威胁。 【翻译: 胡怡思; 审校: 聂永刚】.

Sex-specific niche differentiation is common in marine vertebrates, but how this varies long-term is poorly understood. Here we investigated interannual variation in sexual segregation among breeding northern gannets Morus bassanus, wide-ranging central-place foragers with slight sexual dimorphism. Over 11 breeding seasons, we used GPS tracking and/or stable isotopes to test for sex differences in foraging trip characteristics (range, duration and timing); spatial distribution; habitat selection; and carbon and nitrogen isotopes in blood. When combining data from all years, females foraged further and for longer than males, yet despite this, the foraging areas of the sexes almost completely overlapped. Males and females selected foraging habitats that differed in terms of oceanography but not fishing vessel density. We also detected temporal segregation: females were more likely to be at sea during the day than at night, while males were more likely to be at sea during the night. However, foraging behaviour quantified by all GPS analyses varied interannually, with sex differences detected in some years but not others. Finally, males had consistently higher red blood cell δ13C and δ15N than females across all years, which was not driven by size dimorphism, instead likely by prey choice or very fine-scale habitat selection. We conclude that environmental variation influenced short-term sex differences in movement, but sex differences in stable isotopes that integrate behaviour over longer periods reveal more consistent differences. Our results suggest that inferences drawn from single-year studies may not relate to general patterns, highlighting the importance of long-term studies and combining methods.

Physiological processes, including those that disrupt oxidative balance, have been proposed as key to understanding fundamental life-history trade-offs. Yet, examination of changes in oxidative balance within wild animals across time, space and major life-history challenges remains uncommon. For example, migration presents substantial physiological challenges for individuals, and data on migratory individuals would provide crucial context for exposing the importance of relationships between oxidative balance and fitness outcomes. Here we examined the consistency of commonly used measures of oxidative balance in longitudinally sampled free-living individuals of a long-lived, long-distance migrant, the Brent goose Branta bernicla hrota over periods of months to years. Although inter-individual and temporal variation in measures of oxidative balance were substantial, we found high consistency in measures of lipid peroxidation and circulating non-enzymatic antioxidants in longitudinally sampled individuals. This suggests the potential for the existence of individual oxidative phenotypes. Given intra-individual consistency, we then examined how these physiological measures relate to survival and reproductive success across all sampled individuals. Surprisingly, lower survival was predicted for individuals with lower levels of damage, with no measured physiological metric associated with reproductive success. Our results demonstrate that snapshot measurements of a consistent measure of oxidative balance can inform our understanding of differences in a key demographic trait. However, the positive relationship between oxidative damage and survival emphasises the need to investigate the relationships between the oxidative system and fitness outcomes in other species undergoing similar physiologically challenging life cycles. This would highlight the extent to which variation in such traits and resource allocation trade-offs is a result of adaptation to different life-history strategies.

Highlights: • Since 1960, management for biological invasions totalled at least $95.3 billion. • Damage costs from invasions were substantially higher ($1130.6 billion). • Pre-invasion management spending is 25-times lower than post-invasion. • Management and damage costs are increasing rapidly over time. • Proactive management substantially reduces future costs at the trillion-$ scale. Abstract: The global increase in biological invasions is placing growing pressure on the management of ecological and economic systems. However, the effectiveness of current management expenditure is difficult to assess due to a lack of standardised measurement across spatial, taxonomic and temporal scales. Furthermore, there is no quantification of the spending difference between pre-invasion (e.g. prevention) and post-invasion (e.g. control) stages, although preventative measures are considered to be the most cost-effective. Here, we use a comprehensive database of invasive alien species economic costs (InvaCost) to synthesise and model the global management costs of biological invasions, in order to provide a better understanding of the stage at which these expenditures occur. Since 1960, reported management expenditures have totalled at least US$95.3 billion (in 2017 values), considering only highly reliable and actually observed costs — 12-times less than damage costs from invasions ($1130.6 billion). Pre-invasion management spending ($2.8 billion) was over 25-times lower than post-invasion expenditure ($72.7 billion). Management costs were heavily geographically skewed towards North America (54%) and Oceania (30%). The largest shares of expenditures were directed towards invasive alien invertebrates in terrestrial environments. Spending on invasive alien species management has grown by two orders of magnitude since 1960, reaching an estimated $4.2 billion per year globally (in 2017 values) in the 2010s, but remains 1–2 orders of magnitude lower than damages. National management spending increased with incurred damage costs, with management actions delayed on average by 11 years globally following damage reporting. These management delays on the global level have caused an additional invasion cost of approximately $1.2 trillion, compared to scenarios with immediate management. Our results indicate insufficient management — particularly pre-invasion — and urge better investment to prevent future invasions and to control established alien species. Recommendations to improve reported management cost comprehensiveness, resolution and terminology are also made.

International audience; In addition to being a major threat to biodiversity and ecosystem functioning, biological invasions also have profound impacts on economies and human wellbeing. However, the threats posed by invasive species often do not receive adequate attention and lack targeted management. In part, this may result from different or even ambivalent perceptions of invasive species which have a dual effect for stakeholders—being simultaneously a benefit and a burden. For these species, literature that synthesizes best practice is very limited, and analyses providing a comprehensive understanding of their economics are generally lacking. This has resulted in a critical gap in our understanding of the underlying trade-offs surrounding management efforts and approaches. Here, we explore qualitative trends in the literature for invasive species with dual effects, drawing from both the recently compiled InvaCost database and international case studies. The few invasive species with dual roles in InvaCost provide evidence for a temporal increase in reporting of costs, but with benefits relatively sporadically reported alongside costs. We discuss methods, management, assessment and policy frameworks dedicated to these species, along with lessons learned, complexities and persisting knowledge gaps. Our analysis points at the need to enhance scientific understanding of those species through inter- and cross-disciplinary efforts that can help advance their management.

Biological invasions represent a key threat to island ecosystems, with pronounced impacts across environments and economies. The ecological impacts have received substantial focus, but the economic costs have lacked synthesis at spatial and temporal scales. Here we utilise the InvaCost database, the most comprehensive global assessment of published economic costs of invasive species, to assess reported spend by cost types and socioeconomic sectors, and to examine temporal trends in spending, across islands that differ in their political geography - nation states, overseas territories or offshore islands of continental countries. We based this assessment on 1473 unique cost entries comprising 2914 annual costs totalling almost US$100 million in area-corrected costs between 1965-2020. We find that offshore islands of continental countries incur the greatest total and management costs. However, nation states incurred the greatest damage costs whilst substantially financing management actions, and spent an overall greater proportion of their GDP. In contrast, spending within overseas territories was significantly lower in all respects. The most impacted sector was authorities and stakeholders, demonstrating the key role of government in addressing island invasions. Temporal trends revealed continual increases in spending across all island types. This likely reflects ongoing introduction rates globally alongside an increased recognition of the importance of islands as biodiversity hotspots, and an appetite to tackle invasive species at larger and more socially complex scales. However, the high economic costs of invasions on islands substantiates the need to prevent them in order to avoid this dire threat to biodiversity and its burden on limited conservation resources.

The work was funded by the Natural Environment Research Council (Standard Research Grant NE/H007466/1) and donations from Evan Fountain (In Memoriam donations for Issy Fountain) and Mike Matthewson. Marine vertebrates show a diversity of migration strategies, including sex differences. This may lead to differential demography, but the consequences of such between-sex variation are little understood. Here, we studied the migration of known-sex northern gannets Morus bassanus — a partial migrant with females ~8 % heavier than males. We used geolocators to determine wintering areas of 49 breeding adults (19 females and 30 males during 2010 to 2014) from 2 colonies in the northeast Atlantic (Bass Rock and Grassholm, UK). We also tested for sex-specific survival probabilities using capture−mark−recapture methods (n = 72 individuals Bass Rock, n = 229 individuals Grassholm; 2010−2018) and applied sex-specific population projection matrices (PPMs) to quantify population-level effects. Tracked gannets wintered in a range of large marine ecosystems (LMEs): Canary Current LME (CCLME; 69 %), Celtic-Biscay Shelf LME (16 %), Iberian Coastal LME (8 %), North Sea LME (4 %) or Mediterranean LME (2 %). Migratory destination differed between the sexes: 90 % of females vs. 57 % of males wintered in the CCLME. Survival was similar between the sexes at Bass Rock (mean ± 95 % CI = 0.951 ± 0.053 and 0.956 ± 0.047 for females and males, respectively). At Grassholm, there was evidence of slight sex differences in breeder survival: females had lower annual survival (0.882 ± 0.040) than males (0.946 ± 0.026). At Bass Rock, PPMs with no sex effect best fitted the observed population increase (1994−2014). Sex-specific PPMs fitted the population estimates for Grassholm (1995−2015). Our results reveal that female gannets are more likely to travel further than males to winter in the CCLME. This difference is unlikely due to morphological differences, unlike in other bird species. However, the reason for slightly higher over-winter female mortality at Grassholm is unclear. Publisher PDF

Animals across vertebrate taxa form social communities and often exist as fission–fusion groups. Central place foragers (CPF) may form groups from which they will predictably disperse to forage, either individually or in smaller groups, before returning to fuse with the larger group. However, the function and stability of social associations in predatory fish acting as CPFs is unknown, as individuals do not need to return to a shelter yet show fidelity to core areas. Using dynamic social networks generated from acoustic tracking data, we document spatially structured sociality in CPF grey reef sharks at a Pacific Ocean atoll. We show that sharks form stable social groups over multiyear periods, with some dyadic associations consistent for up to 4 years. Groups primarily formed during the day, increasing in size throughout the morning before sharks dispersed from the reef at night. Our simulations suggest that multiple individuals sharing a central place and using social information while foraging (i.e. local enhancement) will outperform non-CPF social foragers. We show multiyear social stability in sharks and suggest that social foraging with information transfer could provide a generalizable mechanism for the emergence of sociality with group central place foraging.

Individual specialisations in animals are important contributors to a wide range of ecological and evolutionary processes, and have been particularly documented in relation to multiple aspects of foraging behaviours. Central-place foragers, such as seabirds, frequently exhibit pronounced specialisations and individual differences in a variety of foraging traits. In particular, the availability of fisheries discards alongside natural prey resources provides additional potential for differentiation and specialisation for opportunistically scavenging seabird species. However, the consequences of such specialisations for at-sea distributions and intraspecific interactions are not well known. Here, we investigated the links between the degree of dietary specialisation on natural or discarded prey and the foraging movements and spatial occupancy of northern gannets Morus bassanus in relation to differing intraspecific competition at 6 colonies of differing sizes. We found that, at most colonies, individuals with different dietary strategies concentrated foraging at differing levels of intraspecific competition. In addition, individuals pursuing different strategies were frequently, but not consistently, spatially separated, distinctions that were most acutely seen in females. However, this variation in individual strategy had no significant impact on current body condition. These analyses demonstrate how foraging-associated metrics need not covary within an unconstrained system. They also reveal that specialisation can have important consequences for the competitive regimes individuals experience, highlighting the complexity of examining interacting consequences at large spatial scales.

Spatial separation within predator communities can arise via territoriality but also from competitive interactions among and within species. However, linking competitive interactions to predator distribution patterns is difficult and theoretical models predict different habitat selection patterns dependent on habitat quality and how competition manifests itself. While models generally consider competitors to be either equal in ability, or for one phenotype to have a fixed advantage over the other, few studies consider that an animal may only have a competitive advantage in specific habitats. We used  10 years of telemetry data, habitat surveys and behavioral experiments, to show spatial partitioning between and within two species of reef shark (grey reef Carcharhinus amblyrhinchos and blacktip reef sharks C. melanopterus) at an unfished Pacific atoll. Within a species, sharks remained within small ‘sub-habitats’ with very few movements of individuals between sub-habitats, which previous models have suggested could be caused by intra-specific competition. Blacktip reef sharks were more broadly distributed across habitat types but a greater proportion used lagoon and backreef habitats, while grey reef sharks preferred forereef habitats. Grey reef sharks at a nearby atoll where blacktip reef sharks are absent, were distributed more broadly between habitat types than when both species were present. A series of individual-based models predict that habitat separation would only arise if there are competitive interactions between species that are habitat-specific, with grey reefs having a competitive advantage on the forereefs and blacktips in the lagoons and backreef. We provide compelling evidence that competition helps drive distribution patterns and spatial separation of a marine predator community, and highlight that competitive advantages may not be constant but rather dependent on habitats.

1.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. 2.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. 3.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. 4.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. 5.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. This article is protected by copyright. All rights reserved.

Publisher's version (útgefin grein), Fisheries produce large amounts of waste, providing food subsidies for scavengers. Discards influence seabird movement, demography and community structure, but little is known about seabird-fishery interactions where discarding is banned. Here, we investigate how northern gannets Morus bassanus respond to fishing vessels in Iceland, where discarding commercial species is illegal, but birds may still access bait, offal, or catch. We GPS-tracked 82 foraging trips for 36 breeding gannets from two colonies (Skrúður and Hellisey) and obtained time-matched vessel locations. We classified bird behaviour using Hidden Markov Models and then tested the effect of vessel distance on behavioural state-switching using multi-state Markov models. Fishing vessels were present during 94% of foraging trips. However, the likelihood of gannets switching from travelling to foraging was unaffected by vessel proximity, regardless of gear type or activity. When encountering vessels, gannets rarely foraged but instead were more likely to continue travelling. When controlling for population size, gannet foraging trips at both colonies were shorter than expected, suggesting favourable conditions. The lack of behavioural responses to vessels among Icelandic gannets is likely driven by the discard ban and availability of pelagic fishes. Our findings have implications for understanding bycatch risk and the consequences of discard reforms., BLC was supported by a NERC GW4+ Doctoral Training Partnership studentship from the Natural Environment Research Council [NE/L002434/1]. We thank Ólafur Torfason, Niall Tierney, and Rachel Stroud for fieldwork assistance in Skrúður, and Mamma-Rósa for food and housing in Vestmannaeyjar. We thank the Hellisey hunting club for the use of cabin and assistance with boat trips to Hellisey. We thank Filipa Samarra, Miguel Neves, Gary Haskins, and team members in the Icelandic Orca Project for boat trips to Hellisey. We thank Lucy Hawkes, David Pascall, Alice Williams, Richard Phillips, Brendan Godley and all reviewers for constructive comments on the manuscript. The GPS tracking data are available through the BirdLife International Seabird Tracking Database (http://www.seabirdtracking.org).

Abstract Identifying and understanding patterns in movement data are amongst the principal aims of movement ecology. By quantifying the similarity of movement trajectories, inferences can be made about diverse processes, ranging from individual specialisation to the ontogeny of foraging strategies. Movement analysis is not unique to ecology however, and methods for estimating the similarity of movement trajectories have been developed in other fields but are currently under-utilised by ecologists. Here, we introduce five commonly used measures of trajectory similarity: dynamic time warping (DTW), longest common subsequence (LCSS), edit distance for real sequences (EDR), Fréchet distance and nearest neighbour distance (NND), of which only NND is routinely used by ecologists. We investigate the performance of each of these measures by simulating movement trajectories using an Ornstein-Uhlenbeck (OU) model in which we varied the following parameters: (1) the point of attraction, (2) the strength of attraction to this point and (3) the noise or volatility added to the movement process in order to determine which measures were most responsive to such changes. In addition, we demonstrate how these measures can be applied using movement trajectories of breeding northern gannets (Morus bassanus) by performing trajectory clustering on a large ecological dataset. Simulations showed that DTW and Fréchet distance were most responsive to changes in movement parameters and were able to distinguish between all the different parameter combinations we trialled. In contrast, NND was the least sensitive measure trialled. When applied to our gannet dataset, the five similarity measures were highly correlated despite differences in their underlying calculation. Clustering of trajectories within and across individuals allowed us to easily visualise and compare patterns of space use over time across a large dataset. Trajectory clusters reflected the bearing on which birds departed the colony and highlighted the use of well-known bathymetric features. As both the volume of movement data and the need to quantify similarity amongst animal trajectories grow, the measures described here and the bridge they provide to other fields of research will become increasingly useful in ecology. Significance statement As the use of tracking technology increases, there is a need to develop analytical techniques to process such large volumes of data. One area in which this would be useful is the comparison of individual movement trajectories. In response, a variety of measures of trajectory similarity have been developed within the information sciences. However, such measures are rarely used by ecologists who may be unaware of them. To remedy this, we apply five common measures of trajectory similarity to both simulated data and real ecological dataset comprising of movement trajectories of breeding northern gannets. Dynamic time warping and Fréchet distance performed best on simulated data. Using trajectory similarity measures on our gannet dataset, we identified distinct foraging clusters centred on different bathymetric features, demonstrating one application of such similarity measures. As new technology and analysis techniques proliferate across ecology and the information sciences, closer ties between these fields promise further innovative analysis of movement data.

Impacts of generalist predators on declining prey populations are a major conservation issue, but management of this situation is constrained by limited knowledge of the factors influencing predator distribution and activity. In many declining populations of ground-nesting waders, high levels of nest and chick predation are preventing population recovery. Red foxes, Vulpes vulpes, are the main predator but their primary prey is small mammals. On wet grasslands managed for breeding waders, small mammals are concentrated in tall vegetation outside of fields, and nests closer to these patches are less likely to be predated. To assess whether these patterns result from fox attraction to small mammals, and thus the potential for management of tall vegetation to influence nest predation rates, we quantify seasonal and spatial variation in fox and small mammal activity in relation to tall vegetation patches. Across wet grassland sites, tall vegetation patches of any size (> 0.05 ha) supported small mammals and small mammal activity increased throughout the wader breeding season, while the use of fox track plots within fields declined seasonally. Although within field fox track plot use did not vary with distance to tall vegetation, over the 1064 nights of trail camera recording, foxes were seen in areas with tall vegetation on 13 nights compared with short vegetation on only two nights. These findings suggest that lower predation rates of Lapwing, Vanellus vanellus, nests close to tall vegetation could reflect fox attraction to areas with small mammal activity, but any such effects would primarily operate later in the breeding season, and may therefore primarily influence late nests and chicks.

Funding: UK Natural Environment Research Council (NERC Standard Grant NE/H007466/1). SB is currently funded by an ERC consolidator’s grant (310820: STATEMIG). EW is funded by the NERC (Independent Research Fellowship NE/M017990/1). Migration is a fundamental behavioral process prevalent among a wide variety of animal taxa. As individuals are increasingly shown to present consistent responses to environmental cues for breeding or foraging, it may be expected that approaches to migration would present similar among-individual consistencies. Seabirds frequently show consistent individual differences in a range of traits related to foraging and space-use during both the breeding and non-breeding seasons, but the causes and consequences of this consistency are poorly understood. In this study, we combined analysis of geolocation and stable isotope data across multiple years to investigate individual variation in the non-breeding movements and diets of northern gannets Morus bassanus, and the consequences for changes in body condition. We found that individuals were highly repeatable in their non-breeding destination over consecutive years even though the population-level non-breeding distribution spanned >35° of latitude. Isotopic signatures were also strongly repeatable, with individuals assigned to one of two dietary clusters defined by their distinct trophic (δ15N) and spatial (δ13C) position. The only non-breeding destination in which the two dietary clusters co-occurred was off the coast of northwest Africa. The majority of individuals adopted a consistent foraging strategy, as they remained within the same dietary cluster across years, with little variation in body mass corrected for size among these consistent individuals. In contrast, the few individuals that switched clusters between years were in better condition relative to the rest of the population, suggesting there may be benefits to flexibility during the non-breeding period. Our results indicate that a consistent migratory strategy can be effective regardless of wintering region or diet, but that there may be benefits to those individuals able to display flexibility. This appears to be an important behavioral strategy that may enhance individual condition. Publisher PDF

The manner in which patterns of variation and interactions among demographic rates contribute to population growth rate (λ) is key to understanding how animal populations will respond to changing climatic conditions. Migratory species are likely to be particularly sensitive to climatic conditions as they experience a range of different environments throughout their annual cycle. However, few studies have provided fully integrated demographic analyses of migratory populations in response to changing climatic conditions. Here, we employed integrated population models to demonstrate that the environmental conditions experienced during a short but critical period play a central role in the demography of a long-distance migrant, the light-bellied Brent goose (Branta bernicla hrota). Female survival was positively associated with June North Atlantic Oscillation (NAO) values, whereas male survival was not. In contrast, breeding productivity was negatively associated with June NAO, suggesting a trade-off between female survival and reproductive success. Both adult female and adult male survival showed low temporal variation, whereas there was high temporal variation in recruitment and breeding productivity. In addition, while annual population growth was positively correlated with annual breeding productivity, a sensitivity analysis revealed that population growth was most sensitive to changes in adult survival. Our results demonstrate that the environmental conditions experienced during a relatively short-time window at the start of the breeding season play a critical role in shaping the demography of a long-distant Arctic migrant. Crucially, different demographic rates responded in opposing directions to climatic variation, emphasising the need for integrated analysis of multiple demographic traits when understanding population dynamics.

Many established models of animal foraging assume that individuals are ecologically equivalent. However, it is increasingly recognized that populations may comprise individuals who differ consistently in their diets and foraging behaviors. For example, recent studies have shown that individual foraging site fidelity (IFSF, when individuals consistently forage in only a small part of their population's home range) occurs in some colonial breeders. Short-term IFSF could result from animals using a win-stay, lose-shift foraging strategy. Alternatively, it may be a consequence of individual specialization. Pelagic seabirds are colonial central-place foragers, classically assumed to use flexible foraging strategies to target widely dispersed, spatiotemporally patchy prey. However, tracking has shown that IFSF occurs in many seabirds, although it is not known whether this persists across years. To test for long-term IFSF and to examine alternative hypotheses concerning its cause, we repeatedly tracked 55 Northern Gannets (Morus bassanus) from a large colony in the North Sea within and across three successive breeding seasons. Gannets foraged in neritic waters, predictably structured by tidal mixing and thermal stratification, but subject to stochastic, wind-induced overturning. Both within and across years, coarse to mesoscale (tens of kilometers) IFSF was significant but not absolute, and foraging birds departed the colony in individually consistent directions. Carbon stable isotope ratios in gannet blood tissues were repeatable within years and nitrogen ratios were also repeatable across years, suggesting long-term individual dietary specialization. Individuals were also consistent across years in habitat use with respect to relative sea surface temperature and in some dive metrics, yet none of these factors accounted for IFSF. Moreover, at the scale of weeks, IFSF did not decay over time and the magnitude of IFSF across years was similar to that within years, suggesting that IFSF is not primarily the result of win-stay, lose-shift foraging. Rather, we hypothesize that site familiarity, accrued early in-life, causes IFSF by canalizing subsequent foraging decisions. Evidence from this and other studies suggests that IFSF may be common in colonial central-place foragers, with far-reaching consequences for our attempts to understand and conserve these animals in a rapidly changing environment.

A large increase in offshore wind turbine capacity is anticipated within the next decade, raising concerns about possible adverse impacts on birds as a result of collision risk. Birds' flight heights greatly influence this risk, yet height estimates are currently available only using methods such as radar- or ship-based observations over limited areas. 2. Bird-borne data-loggers have the potential to provide improved estimates of collision risk and here, we used data from Global Position System (GPS)-loggers and barometric pressure loggers to track the three-dimensional movements of northern gannets rearing chicks at a large colony in south-east Scotland (Bass Rock), located

Current fishing extraction methods often generate huge quantities of dead or dying biomass that is returned to the sea in the form of discards. This practice produces a readily available clumped resource for many scavengers such as seabirds, but in the face of declining stocks and via policy change, the amount of discards produced is set to decline in the future. To understand how discards have influenced seabird foraging in the past and how birds may respond to future change requires studies examining consistent individual foraging choices. There is increasing evidence that populations may be made up of generalist or specialist foragers and this is key to the population's ability to adapt to change. Here we test for consistent individual foraging behaviour of northern gannets Morus bassanus in relation to fishing vessels and examine consequences of scavenging behaviour in terms of foraging effort and body condition. Using a combination of bird-borne bio-logging devices (GPS and Time Depth Recorders) with high resolution GPS data acquired through vessel monitoring systems on fishing boats, we examined the overlap between birds and fisheries. We found that during repeat foraging trips in the same breeding season, gannets regularly foraged at fishing boats but there were also clear among individual differences in the extent of fisheries overlap. Furthermore, we show for the first time that these differences represent consistent strategies – individual differences in scavenging were highly repeatable across multiple trips within a period of several weeks. However, despite this finding, we found no differences in foraging effort or body condition between scavengers and non-scavengers. Moreover, scavenging strategy did not appear to influence diving behaviour or vary by sex. Scavenging on discards appears to be a strategy employed consistently by a subsection of the population and future work should examine whether these specialisations persist throughout and between years and what causes these individual differences, exploring possible demographic and fitness consequences in light of global changes to fish stocks and fisheries management.

Sexual segregation, common in many species, is usually attributed to intra-specific competi- tion or habitat choice. However, few studies have simultaneously quantified sex-specific foraging be- haviour and habitat use. We combined movement, diving, stable isotope and oceanographic data to test whether sexual segregation in northern gannets Morus bassanus results from sex-specific habitat use. Breeding birds foraging in a seasonally stratified shelf sea were tracked over 3 consecutive breeding seasons (2010�2012). Females made longer trips, foraged far- ther offshore and had lower δ 13 C values than males. Male and female foraging areas overlapped only slightly. Males foraged more in mixed coastal waters, where net primary production (NPP) was relatively high (>3 mg C m �2 d �1 ) and sea-surface temperature (SST) was relatively low ( 15°C) more than females, possibly as a consequence of foraging in productive mixed waters over offshore banks. Females foraged most frequently in stratified offshore waters, of intermediate SST (12�15°C), but exhibited no consistent response to NPP. Sex-specific differences in diving behaviour corresponded with differences in habitat use: males made more long and deep U- shaped dives. Such dives were characteristic of inshore foraging, whereas shorter and shallower V-shaped dives occurred more often in offshore waters. Heavier birds attained greater depths during V-shaped dives, but even when controlling for body mass, females made deeper V-shaped dives than males. Together

Individual foraging specializations, where individuals use a small component of the population niche width, are widespread in nature with important ecological and evolutionary implications. In long-lived animals, foraging ability develops with age, but we know little about the ontogeny of individuality in foraging. Here we use precision global positioning system (GPS) loggers to examine how individual foraging site fidelity (IFSF), a common component of foraging specialization, varies between breeders, failed breeders and immatures in a long-lived marine predator—the northern gannet Morus bassanus. Breeders (aged 5+) showed strong IFSF: they had similar routes and were faithful to distal points during successive trips. However, centrally placed immatures (aged 2–3) were far more exploratory and lacked route or foraging site fidelity. Failed breeders were intermediate: some with strong fidelity, others being more exploratory. Individual foraging specializations were previously thought to arise as a function of heritable phenotypic differences or via social transmission. Our results instead suggest a third alternative—in long-lived species foraging sites are learned during exploratory behaviours early in life, which become canalized with age and experience, and refined where possible—the exploration-refinement foraging hypothesis. We speculate similar patterns may be present in other long-lived species and moreover that long periods of immaturity may be a consequence of such memory-based individual foraging strategies.

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.

Summary 1.Interspecific competitive interactions typically result in niche differentiation to alleviate competition through mechanisms including character displacement. However, competition is not the sole constraint on resource partitioning, and its effects are mediated by factors including the environmental context in which species coexist. 2.Colonial seabirds provide an excellent opportunity to investigate the importance of competition in shaping realized niche widths because their life histories lead to variation in intra- and interspecific competition across the annual cycle. Dense breeding aggregations result in intense competition for prey in surrounding waters, whereas non-breeding dispersal to larger geographical areas produces lower densities of competitors. 3.Bayesian hierarchical models of the isotopic niche, closely aligned to the trophic niche, reveal the degree of segregation between species and functional groups during both time periods. Surprisingly, species explained far more of the variance in the isotopic niche during the non-breeding than the breeding period. 4.Our results underline the key role of non-breeding dynamics in alleviating competition and promoting distinctions between species through the facilitation of resource partitioning. Such situations may be common in a diverse range of communities sustained by ephemeral but abundant food items. 5.This highlights how consideration of the hierarchical grouping of competitive interactions alongside consideration of abiotic constraints across the complete annual cycle allows a full understanding of the role of competition in driving patterns of character displacement.

International audience; Consistent intra-population variability in foraging behaviour is found among a wide range of taxa. Such foraging specialisations are common among marine vertebrates, yet it is not clear how individuals repeatedly locate prey or foraging sites at ocean-wide scales. Using GPS and time-depth loggers we studied the fine-scale foraging behaviour of central-place northern gannets Morus bassanus at two large colonies. First, we estimated the degree of consistency in individual foraging routes and sites across repeated trips. Second, we tested for individual differences in searching behaviour in response to environmental covariates using reaction norms, estimated from mixed effect models. Adult gannets tracked over multiple foraging trips showed repeatable between-individual differences in terminal points and departure angles of foraging trips, but low repeatability in trip duration and trip length. Importantly, individual birds showed highly repeatable dive locations, with consistently different environmental conditions (such as copepod abundance), suggesting a high degree of foraging site specialisation. Gannets also showed between-individual differences in searching behaviour along environmental gradients, such that individuals intensified searching under different conditions. Together these results suggest that widespread individual foraging consistency may represent specialisation and be linked with individual responses to environmental conditions. Such divergent searching behaviour could provide a mechanism by which consistent foraging behaviour arises and is maintained among animals that forage across large spatial scales.

Sexual segregation in foraging and migratory behaviour is widespread among sexu- ally dimorphic marine vertebrates. It has also been described for a number of monomorphic spe- cies, yet the underlying mechanisms are poorly understood. We examined variation among years, seasons and age-classes in sex-specific foraging and over-wintering behaviour in the northern gannet Morus bassanus , a species with slight sexual dimorphism. Our results revealed consistent sexual differences in the stable isotope ratios of breeding birds: over 3 different breeding periods, adult females consistently consumed prey with significantly lower δ 13 C and δ 15 N values than adult males. Additionally, GPS tracking data showed that breeding females foraged further offshore than breeding males (a result consistent with the δ 13 C data), and the home ranges of the 2 sexes were distinct. Analyses of stable isotope ratios using a Bayesian mixing model (SIAR) revealed that breeding males consumed a higher proportion of fishery discards than females. Analysis of stable isotope ratios in red blood cells of immature gannets (aged 2 to 4) indicated that sexual seg- regation was not present in this age-class. Although sample sizes were small and statistical power correspondingly low, analysis of geolocator data and of stable isotope ratios in winter-grown flight feathers revealed no clear evidence of sexual segregation during the non-breeding period. Together these results provide detailed insight into sex-specific behaviour in gannets throughout the annual cycle, and although the mechanisms remain unclear they are unlikely to be explained by slight differences in size.

The control of generalist predator populations is increasingly adopted as a managementtool to combat declines in ground-nesting bird populations. However, compensatory pre-dation by uncontrolled species frequently occurs, so determining the relative impacts ofdifferent predatory species, and hence the relative benefits of their control, can be diffi-cult. Islands, with their reduced faunas, provide natural experimental units for investigat-ing specific predator–prey interactions in detail. We studied Northern Lapwing Vanellusvanellus breeding success on an island where feral Ferrets Mustela furo and HoodedCrows Corvus cornix were subjected to trial control regimes over 2 years. In both years,Lapwing hatching success was >80%, with neither Ferret nor Crow control selected asimportant predictors. Fledging rates in both years were 0.7 young per pair, despite highlyeffective predator removal, although Crow control potentially resulted in compensatorypredation by Common Ravens C. corax. Neither mustelid nor corvid control producedsignificant immediate benefits for Lapwings. This suggests that mesopredator release ofmustelids in mainland situations is unlikely to be a consistent threat to Lapwing, andprovides further evidence that declines in this species are unlikely to be tackled success-fully through predator management alone.Keywords: compensatory predation, corvid, ground-nesting birds, island, mesopredator release,mustelid, Vanellus vanellus, wader.Many wild bird populations are in decline (BirdlifeInternational 2008), with habitat loss and fragmen-tation and increased predation from both invasiveand generalist predators frequently identified as theprincipal causes (Newton 1998, 2004, Evans2004). Although birds living in open habitats areknown to suffer naturally high nest predation rates(e.g. Martin 1993), populations can still be verysusceptible to any increases in such pressure(Senner & Howe 1984, Suarez et al. 1993, Dowding& Murphy 2001).Changes in landscape characteristics are thoughtto be one of the key elements resulting in increasedpredation risk. These changes act via a variety ofmechanisms, including the creation of additionalforaging opportunities for predators (Newton2004), the reduction in suitable nesting habitatwhich can make nests more easily detected ordisrupt communal defence (Evans 2004), and anincrease in habitat fragmentation which canenhance the accessibility of nests to predators(Marzluff & Neatherlin 2006). Generalist preda-tors can also be subsidized by anthropogenic foodsources in modified landscapes, such as rubbish tips(Boarman 2003) or carrion (Newton 2004). Suchsubsidies can increase the potential for hyperpreda-tion as populations reach densities above thosethat natural prey would ordinarily sustain (Cour-champ et al. 2000).As a consequence of increases in predation pres-sure, control of predators such as corvids and

Introduced mammalian predators have had significant impacts on many native prey species. Although control of such predators for conservation management is becoming increasingly commonplace, it is often undertaken at a relatively small scale in relation to the overall predator population. Processes such as immigration mean that it remains difficult to determine the effectiveness of control measures. We investigated the impacts of feral ferret Mustela furo removal on the entire feral ferret population on Rathlin Island, UK. Removal of ferrets prior to breeding led to a substantial increase in the post-dispersal population through the enhanced survival of juveniles. Despite increased numbers, overwinter survival remained high, potentially aided by the reduced territoriality shown by this feral species compared to wild carnivores. The response of this ferret population to control is a further illustration of the complex ecological processes and outcomes arising from the anthropogenic disruption of wildlife populations. It highlights how partial or localised management may prove ineffective, and at worst might exacerbate the problems that management was designed to avert.

Oxidative stress is a likely consequence of hard physical exertion and thus a potential mediator of life-history trade-offs in migratory animals. However, little is known about the relative importance of intrinsic and extrinsic stressors on the oxidative state of individuals in wild populations. We quantified the relationships between air temperature, sex, body condition and three markers of oxidative state (malondialdehyde, superoxide dismutase and total antioxidant capacity) across hundreds of individuals of a long-distance migrant (the brent goose Branta bernicla hrota ) during wintering and spring staging. We found that air temperature and migratory stage were the strongest predictors of oxidative state. This emphasizes the importance of extrinsic factors in regulating the oxidative state of migrating birds, with differential effects across the migration. The significance of abiotic effects demonstrates an additional mechanism by which changing climates may affect migratory costs.

Non-native invasive mammal species have caused major ecological change on many islands. To conserve native species diversity, invasive mammals have been eradicated from several islands not inhabited by humans. We reviewed the challenges associated with campaigns to eradicate invasive mammals from islands inhabited by humans and domestic animals. On these islands, detailed analyses of the social, cultural, and economic costs and benefits of eradication are required to increase the probability of local communities supporting the eradication campaign. The ecological benefits of eradication (e.g., improvement of endemic species' probability of survival) are difficult to trade-off against social and economic costs due to the lack of a common currency. Local communities may oppose an eradication campaign because of perceived health hazards, inconvenience, financial burdens, religious beliefs, or other cultural reasons. Besides these social challenges, the presence of humans and domestic animals also complicates eradication and biosecurity procedures (measures taken to reduce the probability of unwanted organisms colonizing an island to near zero). For example, houses, garbage-disposal areas, and livestock-feeding areas can provide refuges for certain mammals and therefore can decrease the probability of a successful eradication. Transport of humans and goods to an island increases the probability of inadvertent reintroduction of invasive mammals, and the establishment of permanent quarantine measures is required to minimize the probability of unwanted recolonization after eradication. We recommend a close collaboration between island communities, managers, and social scientists from the inception of an eradication campaign to increase the probability of achieving and maintaining an island permanently free of invasive mammals.

Summary 1. The detrimental impacts of invasive, non-native species on islands are widely acknowledged and it is often best to act rapidly against such species, even where uncertainty exists over the best way to proceed. If management actions are evaluated and refined, using information learnt from the biology of culled animals, this uncertainty can be gradually reduced, increasing the likelihood of a successful outcome. 2. American mink Neovison vison carcasses were collected as part of an eradication campaign on several islands of the Outer Hebrides, Scotland, and stable isotope analysis was used to describe ecological variation in this invasive non-native predator. 3. Isotope profiles from individual mink whiskers demonstrated how behaviour at a population level changed markedly over time. As the eradication campaign progressed, mink increased their reliance on marine food sources and focused their activity on the coastline. Stable isotope analyses also demonstrated sex-related changes in foraging and ranging behaviour in relation to food resource availability on the two main island complexes. 4. Synthesis and applications. Our findings contribute to the refinement of a campaign to extend the successful eradication of mink from Uist and Harris, to the whole of the Outer Hebrides archipelago, UK. They also highlight the potential for stable isotope approaches to provide more detailed postmortem information that can inform adaptive management of wildlife populations for conservation objectives.

Peer-reviewed. This document is the Accepted Manuscript version of a Published Work that appeared in final form in Current Biology. To access the final edited and published work see doi:10.1016/j.cub.2014.04.041, Exploitation of the seas is currently unsustainable, with increasing demand for marine resources placing intense pressure on the Earth’s largest ecosystem [1]. The scale of anthropogenic effects varies from local to entire ocean basins 1, 2 and 3. For example, discards of commercial capture fisheries can have both positive and negative impacts on scavengers at the population and community-level 2, 3, 4, 5 and 6, although this is driven by individual foraging behaviour 3 and 7. Currently, we have little understanding of the scale at which individual animals initiate such behaviours. We use the known interaction between fisheries and a wide-ranging seabird, the Northern gannet Morus bassanus [3], to investigate how fishing vessels affect individual birds’ behaviours in near real-time. We document the footprint of fishing vessels’ (≥15 m length) influence on foraging decisions (≤11 km), and a potential underlying behavioural mechanism, by revealing how birds respond differently to vessels depending on gear type and activity. Such influences have important implications for fisheries, including the proposed discard ban [8]), and wider marine management.

This Is the Place Bats, bees, seals, and many seabirds practice central-place foraging, leaving a central home site, such as a hive or a rookery, to forage in a specific territory. Such species also share the challenge of competing for local resources with individuals from separate colonies. Using satellite tags, Wakefield et al. (p. 68 , published online 6 June; see the Perspective by Weimerskirch ) followed over 180 northern gannets to determine potential drivers of foraging territory division. Boundaries among colonial territories arose as a result of competition with individuals from other territories. Individuals from the same colony appeared to share information about foraging sites, presumably contributing to the establishment and maintenance of specific, long-term colonial territories.