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Evidence of segregation in foraging habitat has been demonstrated in some top marine predators, including cetaceans, pinnipeds and seabirds. However, most data are not adequate to assess differences relating to body size or seasonal influences. This has implications for quantitative modelling of population-level predator-prey interactions and ecosystem structure. We examined potential influence of body size and 'fatness' on the foraging trip characteristics of a top marine predator, the Atlantic grey seal (Halichoerus grypus), in southwest Ireland within the framework of optimal foraging theory to examine how female grey seals foraging behaviour varied with size, and across the period between moult and breeding. Larger seals undertook trips of greater duration and travelled further from haul-out sites than smaller seals. However, body fat was negatively associated with trip duration and extent. Seals spent more time at sea during the summer, but trips were shorter in extent, suggesting more localized foraging during this season., Introduction Optimal foraging theory is an important tool for increasing our understanding of animal behaviour. According to optimal foraging theory (Pyke 1978), long-distance travel should only be taken if there [...]

Relatively simple foraging radius models have the potential to generate predictive distributions for a large number of species rapidly, thus providing a cost‐effective alternative to large‐scale surveys or complex modelling approaches. Their effectiveness, however, remains largely untested. Here we compare foraging radius distribution models for all breeding seabirds in Ireland, to distributions of empirical data collected from tracking studies and aerial surveys. At the local/colony level, we compared foraging radius distributions to GPS tracking data from seabirds with short (Atlantic puffin Fratercula arctica, and razorbill Alca torda) and long (Manx shearwater Puffinus puffinus, and European storm‐petrel Hydrobates pelagicus) foraging ranges. At the regional/national level, we compared foraging radius distributions to extensive aerial surveys conducted over a two‐year period. Foraging radius distributions were significantly positively correlated with tracking data for all species except Manx shearwater. Correlations between foraging radius distributions and aerial survey data were also significant, but generally weaker than those for tracking data. Correlations between foraging radius distributions and aerial survey data were benchmarked against generalised additive models (GAMs) of the aerial survey data that included a range of environmental covariates. While GAM distributions had slightly higher correlations with aerial survey data, the results highlight that the foraging radius approach can be a useful and pragmatic approach for assessing breeding distributions for many seabird species. The approach is likely to have acceptable utility in complex, temporally variable ecosystems and when logistic and financial resources are limited. [ABSTRACT FROM AUTHOR]

Potentially some of the biggest gaps in our knowledge about the ecology of Antarctic fur seals ( Arctocephalus gazella) relate to juvenile animals. We investigated the at-sea distribution of five male and five female fur seal pups post-weaning. The study was carried out at Bird Island, South Georgia during two successive winters using satellite-linked platform transmitter terminals (PTTs). Our results are analysed in relation to pup sex and the physical environment and productivity of those 2 years, as well as in the context of our present knowledge of where post-breeding females and males forage. The available physical and biological data during both of the winters of this study suggest that both years were not unusual. We report marked differences between the sexes with male pups foraging significantly further away from land and their birth site than do females. The pups foraged in areas to the East of Bird Island seldom reported as foraging areas for the adult population. Also as winter progressed they showed a more oceanic distribution leaving the continental shelf, possibly to exploit a different prey source that was more readily available in the upper water column. [ABSTRACT FROM AUTHOR]

Density-dependent competition for food influences the foraging behaviour and demography of colonial animals, but how this influence varies across a species' latitudinal range is poorly understood. Here we used satellite tracking from 21 Northern Gannet Morus bassanus colonies (39% of colonies worldwide, supporting 73% of the global population) during chick-rearing to test how foraging trip characteristics (distance and duration) covary with colony size (138-60 953 breeding pairs) and latitude across 89% of their latitudinal range (46.81-71.23° N). Tracking data for 1118 individuals showed that foraging trip duration and maximum distance both increased with square-root colony size. Foraging effort also varied between years for the same colony, consistent with a link to environmental variability. Trip duration and maximum distance also decreased with latitude, after controlling for colony size. Our results are consistent with density-dependent reduction in prey availability influencing colony size and reveal reduced competition at the poleward range margin. This provides a mechanism for rapid population growth at northern colonies and, therefore, a poleward shift in response to environmental change. Further work is required to understand when and how colonial animals deplete nearby prey, along with the positive and negative effects of social foraging behaviour., Competing Interests: We declare we have no competing interests., (© 2024 The Author(s).)

The importance of scale when investigating ecological patterns and processes is recognised across many species. In marine ecosystems, the processes that drive species distribution have a hierarchical structure over multiple nested spatial and temporal scales. Hence, multi-scale approaches should be considered when developing accurate distribution models to identify key habitats, particularly for populations of conservation concern. Here, we propose a modelling procedure to identify the best spatial and temporal scale for each modelled and remotely sensed oceanographic variable to model harbour porpoise ( Phocoena phocoena ) distribution within the Irish Exclusive Economic Zone. Harbour porpoise sightings were recorded during dedicated line-transect aerial surveys conducted in the summers of 2016, 2021 and 2022. Binary generalised additive models were used to assess the relationships between porpoise presence and oceanographic variables at different spatial (5-40 km) and temporal (daily, monthly and across survey period) scales. Selected variables included sea surface temperature, thermal fronts, chlorophyll-a, sea surface height, mixed layer depth and salinity. A total of 30,514 km was covered on-effort with 216 harbour porpoise sightings recorded. Overall, the best spatial scale corresponded to the coarsest resolution considered in this study (40 km), while porpoise presence showed stronger association with oceanographic variables summarised at a longer temporal scale. Habitat models including covariates at coarse spatial and temporal scales may better reflect the processes driving availability and abundance of resources at these large scales. These findings support the hypothesis that a multi-scale approach should be applied when investigating species distribution. Identifying suitable spatial and temporal scale would improve the functional interpretation of the underlying relationships, particularly when studying how a small marine predator interacts with its environment and responds to climate and ecosystem changes., Competing Interests: The authors declare no conflict of interest., (© 2024 The Author(s). Ecology and Evolution published by John Wiley & Sons Ltd.)

Many vertebrates show lateralized behaviour, or handedness, where an individual preferentially uses one side of the body more than the other. This is generally thought to be caused by brain lateralization and allows functional specializations such as sight, locomotion, and decision-making among other things. We deployed accelerometers on 51 northern gannets, Morus bassanus , to test for behavioural lateralization during plunge dives. When plunge diving, gannets 'roll' to one side, and standard indices indicated that 51% of individuals were left-sided, 43% right-sided, and 6% 'non-lateralized'. Lateralization indices provide no measure of error and do not account for environmental covariance, so we conducted two repeatability analyses on individuals' dive roll direction and angle. Dive side lateralization was highly repeatable among individuals over time at the population level ( R = 0.878, p < 0.001). Furthermore, roll angle was also highly repeatable in individuals ( R = 0.751, p < 0.001) even after controlling for lateralized state. Gannets show individual specializations in two different parts of the plunge diving process when attempting to catch prey. This is the first demonstration of lateralization during prey capture in a foraging seabird. It is also one of the few demonstrations of behavioural lateralization in a mixed model approach, providing a structure for further exploring behavioural lateralization.

Plastic pollution is distributed patchily around the world's oceans. Likewise, marine organisms that are vulnerable to plastic ingestion or entanglement have uneven distributions. Understanding where wildlife encounters plastic is crucial for targeting research and mitigation. Oceanic seabirds, particularly petrels, frequently ingest plastic, are highly threatened, and cover vast distances during foraging and migration. However, the spatial overlap between petrels and plastics is poorly understood. Here we combine marine plastic density estimates with individual movement data for 7137 birds of 77 petrel species to estimate relative exposure risk. We identify high exposure risk areas in the Mediterranean and Black seas, and the northeast Pacific, northwest Pacific, South Atlantic and southwest Indian oceans. Plastic exposure risk varies greatly among species and populations, and between breeding and non-breeding seasons. Exposure risk is disproportionately high for Threatened species. Outside the Mediterranean and Black seas, exposure risk is highest in the high seas and Exclusive Economic Zones (EEZs) of the USA, Japan, and the UK. Birds generally had higher plastic exposure risk outside the EEZ of the country where they breed. We identify conservation and research priorities, and highlight that international collaboration is key to addressing the impacts of marine plastic on wide-ranging species., (© 2023. The Author(s).)

The 2022 highly pathogenic avian influenza (HPAI) outbreak that occurred in many European countries affected several seabird species. Among them, northern gannets ( Morus bassanus ) were particularly impacted. We conducted aerial surveys in waters around the two largest gannet colonies in southwest Ireland (Little Skellig and Bull Rock, together representing 87% of the national population) in September 2022. During surveys dead and alive northern gannets were counted on survey effort. A total of 184 dead gannets were recorded on survey effort, representing 3.74% of the total number of gannets recorded. We estimated the abundance of dead gannets in the surveyed area at 1526 (95% confidence intervals (CIs) 1450-1605) individuals. The percentage of dead gannets observed was used to estimate a minimum local population mortality of 3126 (95% CIs 2993-3260) individuals across both colonies. Aerial surveys provided key information on gannet mortality from HPAI at sea. The study provides the first estimate of gannet mortality in the two largest gannetries in Ireland.

Body-mounted accelerometers provide a new prospect for estimating power use in flying birds, as the signal varies with the two major kinematic determinants of aerodynamic power: wingbeat frequency and amplitude. Yet wingbeat frequency is sometimes used as a proxy for power output in isolation. There is, therefore, a need to understand which kinematic parameter birds vary and whether this is predicted by flight mode (e.g. accelerating, ascending/descending flight), speed or morphology. We investigate this using high-frequency acceleration data from (i) 14 species flying in the wild, (ii) two species flying in controlled conditions in a wind tunnel and (iii) a review of experimental and field studies. While wingbeat frequency and amplitude were positively correlated, R 2 values were generally low, supporting the idea that parameters can vary independently. Indeed, birds were more likely to modulate wingbeat amplitude for more energy-demanding flight modes, including climbing and take-off. Nonetheless, the striking variability, even within species and flight types, highlights the complexity of describing the kinematic relationships, which appear sensitive to both the biological and physical context. Notwithstanding this, acceleration metrics that incorporate both kinematic parameters should be more robust proxies for power than wingbeat frequency alone.

Understanding the sensory ecology of species is vital if we are to predict how they will function in a changing environment. Visual cues are fundamentally important for many predators when detecting and capturing prey. However, many marine areas have become more turbid through processes influenced by climate change, potentially affecting the ability of marine predators to detect prey. We performed the first study that directly relates a pelagic seabird species's foraging behaviour to oceanic turbidity. We collected biologging data from 79 foraging trips and 5472 dives of a visually dependent, pursuit-diving seabird, the Manx shearwater ( Puffinus puffinus ). Foraging behaviour was modelled against environmental variables affecting underwater visibility, including water turbidity, cloud cover and solar angle. Shearwaters were more likely to initiate area-restricted search and foraging dives in clearer waters. Underwater visibility also strongly predicted dive rate and depth, suggesting that fine-scale prey capture was constrained by the detectability of prey underwater. Our novel use of dynamic descriptors of underwater visibility suggests that visual cues are vital for underwater foraging. Our data indicate that climate change could negatively impact seabird populations by making prey more difficult to detect, compounded by the widely reported effects of reduced prey populations.

Many animals show sexually divergent foraging behaviours reflecting different physiological constraints or energetic needs. We used a bioenergetics approach to examine sex differences in foraging behaviour of the sexually monomorphic northern gannet. We derived a relationship between dynamic body acceleration and energy expenditure to quantify the energetic cost of prey capture attempts (plunge dives). Fourteen gannets were tracked using GPS, time depth recorders (TDR) and accelerometers. All plunge dives in a foraging trip represented less than 4% of total energy expenditure, with no significant sex differences in expenditure. Despite females undertaking significantly more dives than males, this low energetic cost resulted in no sex differences in overall energy expenditure across a foraging trip. Bayesian stable isotope mixing models based on blood samples highlighted sex differences in diet; however, calorific intake from successful prey capture was estimated to be similar between sexes. Females experienced 10.28% higher energy demands, primarily due to unequal chick provisioning. Estimates show a minimum of 19% of dives have to be successful for females to meet their daily energy requirements, and 26% for males. Our analyses suggest northern gannets show sex differences in foraging behaviour primarily related to dive rate and success rather than the energetic cost of foraging or energetic content of prey., (© 2022 The Authors.)

Each winter, the North Atlantic Ocean is the stage for numerous cyclones, the most severe ones leading to seabird mass-mortality events called "winter wrecks." 1-3 During these, thousands of emaciated seabird carcasses are washed ashore along European and North American coasts. Winter cyclones can therefore shape seabird population dynamics 4 , 5 by affecting survival rates as well as the body condition of surviving individuals and thus their future reproduction. However, most often the geographic origins of impacted seabirds and the causes of their deaths remain unclear. 6 We performed the first ocean-basin scale assessment of cyclone exposure in a seabird community by coupling winter tracking data for ∼1,500 individuals of five key North Atlantic seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia, and Rissa tridactyla) and cyclone locations. We then explored the energetic consequences of different cyclonic conditions using a mechanistic bioenergetics model 7 and tested the hypothesis that cyclones dramatically increase seabird energy requirements. We demonstrated that cyclones of high intensity impacted birds from all studied species and breeding colonies during winter but especially those aggregating in the Labrador Sea, the Davis Strait, the surroundings of Iceland, and the Barents Sea. Our broad-scale analyses suggested that cyclonic conditions do not increase seabird energy requirements, implying that they die because of the unavailability of their prey and/or their inability to feed during cyclones. Our study provides essential information on seabird cyclone exposure in a context of marked cyclone regime changes due to global warming. 8 ., Competing Interests: Declaration of interests The authors declare no competing interests., (Crown Copyright © 2021. Published by Elsevier Inc. All rights reserved.)

We explored the implications of reaching the Paris Agreement Objective of limiting global warming to <2°C for the future winter distribution of the North Atlantic seabird community. We predicted and quantified current and future winter habitats of five North Atlantic Ocean seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia and Rissa tridactyla) using tracking data for ~1500 individuals through resource selection functions based on mechanistic modeling of seabird energy requirements, and a dynamic bioclimate envelope model of seabird prey. Future winter distributions were predicted to shift with climate change, especially when global warming exceed 2°C under a "no mitigation" scenario, modifying seabird wintering hotspots in the North Atlantic Ocean. Our findings suggest that meeting Paris agreement objectives will limit changes in seabird selected habitat location and size in the North Atlantic Ocean during the 21st century. We thereby provide key information for the design of adaptive marine-protected areas in a changing ocean., (© 2020 John Wiley & Sons Ltd.)

Understanding how animals forage is a central objective in ecology. Theory suggests that where food is uniformly distributed, Brownian movement ensures the maximum prey encounter rate, but when prey is patchy, the optimal strategy resembles a Lévy walk where area-restricted search (ARS) is interspersed with commuting between prey patches. Such movement appears ubiquitous in high trophic-level marine predators. Here, we report foraging and diving behaviour in a seabird with a high cost of flight, the Atlantic puffin ( Fratercula arctica), and report a clear lack of Brownian or Levy flight and associated ARS. Instead, puffins foraged using tides to transport them through their feeding grounds. Energetic models suggest the cost of foraging trips using the drift strategy is 28-46% less than flying between patches. We suggest such alternative movement strategies are habitat-specific, but likely to be far more widespread than currently thought.

In ecological studies it is often assumed that predator foraging strategies and resource use are geographically and seasonally homogeneous, resulting in relatively static trophic relationships. However, certain centrally placed foragers (e.g. seals) often have terrestrial sites for breeding, resting, and moulting that are geographically distinct, and associated with different habitat types. Therefore, accurate estimations of predator diet at relevant spatial and temporal scales are key to understanding energetic requirements, predator-prey interactions and ecosystem structure. We investigate geographic variation in the diet of grey seals (Halichoerus grypus), a relatively abundant and widely distributed central place forager, to provide insights into geographic variation in resource use. Prey composition was identified using scat samples collected over concurrent timescales and a multivariate approach was used to analyse diet from two contrasting habitats. Regional differences in prey assemblages occurred within all years (2011-2013) and all seasons (ANOSIM, all p<0.05), apart from in winter. Telemetry data were used to identify core foraging areas and habitats most likely associated with scat samples collected at the two haul-out sites. Regional differences in the diet appear to reflect regional differences in the physical habitat features, with seals foraging in deeper waters over sandy substrates showing a higher prevalence of pelagic and bentho-pelagic prey species such as blue whiting and sandeels. Conversely, seals foraging in comparatively shallow waters had a greater contribution of demersal and groundfish species such as cephalopods and flatfish in their diet. We suggest that shallower waters enable seals to spend more time foraging along the benthos while remaining within aerobic dive limits, resulting in more benthic species in the diet. In contrast, the diet of seals hauled-out in areas adjacent to deeper waters indicates that either seals engage in a more pelagic foraging strategy, or that seals can spend less time at the benthos, resulting in comparatively more pelagic prey recovered in the diet. The substantial differences in prey assemblages over a small spatial scale (<300 km) demonstrates the importance of using regionally-specific diet information in ecosystem-based models to better account for different trophic interactions., Competing Interests: The authors have declared that no competing interests exist.

There is worldwide concern about the status of elasmobranchs, primarily as a result of overfishing and bycatch with subsequent ecosystem effects following the removal of top predators. Whilst abundant and wide-ranging, blue sharks (Prionace glauca) are the most heavily exploited shark species having suffered marked declines over the past decades, and there is a call for robust abundance estimates. In this study, we utilized depth data collected from two blue sharks using pop-up satellite archival tags, and modelled the proportion of time the sharks were swimming in the top 1-meter layer and could therefore be detected by observers conducting aerial surveys. The availability models indicated that the tagged sharks preferred surface waters whilst swimming over the continental shelf and during daytime, with a model-predicted average proportion of time spent at the surface of 0.633 (SD = 0.094) for on-shelf, and 0.136 (SD = 0.075) for off-shelf. These predicted values were then used to account for availability bias in abundance estimates for the species over a large area in the Northeast Atlantic, derived through distance sampling using aerial survey data collected in 2015 and 2016 and modelled with density surface models. Further, we compared abundance estimates corrected with model-predicted availability to uncorrected estimates and to estimates that incorporated the average time the sharks were available for detection. The mean abundance (number of individuals) corrected with modelled availability was 15,320 (CV = 0.28) in 2015 and 11,001 (CV = 0.27) in 2016. Depending on the year, these estimates were ~7 times higher compared to estimates without the bias correction, and ~3 times higher compared to the abundances corrected with average availability. When the survey area contains habitat heterogeneity that may affect surfacing patterns of animals, modelling animals' availability provides a robust alternative to correcting for availability bias and highlights the need for caution when applying "average" correction factors., Competing Interests: ALNILAM Research and Conservation Ltd provided support in the form of salaries for author AC. This does not alter our adherence to PLOS ONE policies on sharing data and materials.

Which factors shape animals' migration movements across large geographical scales, how different migratory strategies emerge between populations, and how these may affect population dynamics are central questions in the field of animal migration [1] that only large-scale studies of migration patterns across a species' range can answer [2]. To address these questions, we track the migration of 270 Atlantic puffins Fratercula arctica, a red-listed, declining seabird, across their entire breeding range. We investigate the role of demographic, geographical, and environmental variables in driving spatial and behavioral differences on an ocean-basin scale by measuring puffins' among-colony differences in migratory routes and day-to-day behavior (estimated with individual daily activity budgets and energy expenditure). We show that competition and local winter resource availability are important drivers of migratory movements, with birds from larger colonies or with poorer local winter conditions migrating further and visiting less-productive waters; this in turn led to differences in flight activity and energy expenditure. Other behavioral differences emerge with latitude, with foraging effort and energy expenditure increasing when birds winter further north in colder waters. Importantly, these ocean-wide migration patterns can ultimately be linked with breeding performance: colony productivity is negatively associated with wintering latitude, population size, and migration distance, which demonstrates the cost of competition and migration on future breeding and the link between non-breeding and breeding periods. Our results help us to understand the drivers of animal migration and have important implications for population dynamics and the conservation of migratory species., (Copyright © 2017 Elsevier Ltd. All rights reserved.)