Breeding Thin-Billed Prions Use Marine Habitats Ranging from Inshore to Distant Antarctic Waters.

Pelagic seabirds cover large distances efficiently and thus may reach a variety of marine habitats during breeding. Previous studies using stable isotope data and geolocators suggested that Thin-billed Prions breeding in the Falkland Islands in the Southwest Atlantic may forage in temperate waters over the Patagonian Shelf or cross the Drake Passage to forage in Antarctic waters south of the Polar Front. We deployed miniature GPS dataloggers to track Thin-billed prions in the Falkland Islands during incubation (3 seasons) and chick-rearing (2 seasons). Thin-billed Prions had a wide distribution during incubation, covering latitudes between 43 and 60° S, with trip lengths of ca. 2000 km over seven days, on average. Thin-billed Prions from two nearby sites (60 km apart) were spatially segregated in their incubation trips, with New Island Thin-billed Prions foraging over the Patagonian Shelf, compared to Thin-billed Prions from Bird Island, that foraged in the region of the Polar Front. During chick-rearing, Thin-billed Prions from New Island undertook both long trips to the Patagonian Shelf and south of the Polar Front (30% of trips were 5-11 days), and short trips (70% of trips were 1-4 days) when they foraged more locally, including in inshore waters around the Falkland Islands. Females carried out more trips to distant sites. Thus, Thin-billed showed a high flexibility in foraging areas, habitats and foraging trip durations, which enable them to benefit from both, temperate and Antarctic environments.

Mercury contamination and potential health risks to Arctic seabirds and shorebirds.

Since the last Arctic Monitoring and Assessment Programme (AMAP) effort to review biological effects of mercury (Hg) on Arctic biota in 2011 and 2018, there has been a considerable number of new Arctic bird studies. This review article provides contemporary Hg exposure and potential health risk for 36 Arctic seabird and shorebird species, representing a larger portion of the Arctic than during previous AMAP assessments now also including parts of the Russian Arctic. To assess risk to birds, we used Hg toxicity benchmarks established for blood and converted to egg, liver, and feather tissues. Several Arctic seabird populations showed Hg concentrations that exceeded toxicity benchmarks, with 50 % of individual birds exceeding the "no adverse health effect" level. In particular, 5 % of all studied birds were considered to be at moderate or higher risk to Hg toxicity. However, most seabirds (95 %) were generally at lower risk to Hg toxicity. The highest Hg contamination was observed in seabirds breeding in the western Atlantic and Pacific Oceans. Most Arctic shorebirds exhibited low Hg concentrations, with approximately 45 % of individuals categorized at no risk, 2.5 % at high risk category, and no individual at severe risk. Although the majority Arctic-breeding seabirds and shorebirds appeared at lower risk to Hg toxicity, recent studies have reported deleterious effects of Hg on some pituitary hormones, genotoxicity, and reproductive performance. Adult survival appeared unaffected by Hg exposure, although long-term banding studies incorporating Hg are still limited. Although Hg contamination across the Arctic is considered low for most bird species, Hg in combination with other stressors, including other contaminants, diseases, parasites, and climate change, may still cause adverse effects. Future investigations on the global impact of Hg on Arctic birds should be conducted within a multi-stressor framework. This information helps to address Article 22 (Effectiveness Evaluation) of the Minamata Convention on Mercury as a global pollutant.

Comparing non-breeding distribution and behavior of red-legged kittiwakes from two geographically distant colonies.

Knowledge of non-breeding distributions is a crucial component to seabird conservation, as conditions during the non-breeding period can play an important role in regulating seabird populations. Specifically, if seabirds from different colonies share the same wintering grounds, conditions in that shared region could have a widespread impact on multiple breeding populations. Red-legged kittiwakes (Rissa brevirostris) are endemic to the Bering Sea and may be especially susceptible to effects of climate change due to a restricted breeding range, small population size, and specialized diet. To examine whether red-legged kittiwakes from different breeding colonies overlapped in winter distribution and activity patterns, we used geolocation loggers to simultaneously track individuals from the two largest red-legged kittiwake breeding colonies in Alaska (separated by over 1000 km) during two consecutive non-breeding periods. We found that non-breeding activity patterns were generally similar between birds originating from the two colonies, but birds employed different migratory strategies during the early winter. Kittiwakes from Buldir Island in the western Aleutian Islands left the colony in September and immediately headed west, spending October through December around the Sea of Okhotsk and the Kuril Islands. In contrast, birds from St. George Island in the Pribilof Islands remained in the eastern Bering Sea or around the eastern Aleutian Islands for a couple months before traveling farther west. During late winter however, from January through March, birds from both colonies converged south of Kamchatka and east of the Kuril Islands over the Kuril-Kamchatka Trench and in the Western Subarctic Gyre before returning to their respective colonies in the spring. This late winter overlap in distributions along the Kuril-Kamchatka Trench suggests the region is a winter hotspot for red-legged kittiwakes and highlights the importance of this region for the global kittiwake population.

Overlap between marine predators and proposed Marine Managed Areas on the Patagonian Shelf.

Static (fixed-boundary) protected areas are key ocean conservation strategies, and marine higher predator distribution data can play a leading role toward identifying areas for conservation action. The Falkland Islands are a globally significant site for colonial breeding marine higher predators (i.e., seabirds and pinnipeds). However, overlap between marine predators and Falkland Islands proposed Marine Managed Areas (MMAs) has not been quantified. Hence, to provide information required to make informed decisions regarding the implementation of proposed MMAs, our aims were to objectively assess how the proposed MMA network overlaps with contemporary estimates of marine predator distribution. We collated tracking data (1999-2019) and used a combination of kernel density estimation and model-based predictions of spatial usage to quantify overlap between colonial breeding marine predators and proposed Falkland Islands MMAs. We also identified potential IUCN Key Biodiversity Areas (pKBAs) using (1) kernel density based methods originally designed to identify Important Bird and Biodiversity Areas (IBAs) and (2) habitat preference models. The proposed inshore MMA, which extends three nautical miles from the Falkland Islands, overlapped extensively with areas used by colonial breeding marine predators. This reflects breeding colonies being distributed throughout the Falklands archipelago, and use being high adjacent to colonies due to central-place foraging constraints. Up to 45% of pKBAs identified via kernel density estimation were located within the proposed MMAs. In particular, the proposed Jason Islands Group MMA overlapped with pKBAs for three marine predator species, suggesting it is a KBA hot spot. However, tracking data coverage was incomplete, which biased pKBAs identified using kernel density methods, to colonies tracked. Moreover, delineation of pKBA boundaries were sensitive to the choice of smoothing parameter used in kernel density estimation. Delineation based on habitat model predictions for both sampled and unsampled colonies provided less biased estimates, and revealed 72% of the Falkland Islands Conservation Zone was likely a KBA. However, it may not be practical to consider such a large area for fixed-boundary management. In the context of wide-ranging marine predators, emerging approaches such as dynamic ocean management could complement static management frameworks such as MMAs, and provide protection at relevant spatiotemporal scales.

Foraging in marine habitats increases mercury concentrations in a generalist seabird.

Methylmercury concentrations vary widely across geographic space and among habitat types, with marine and aquatic-feeding organisms typically exhibiting higher mercury concentrations than terrestrial-feeding organisms. However, there are few model organisms to directly compare mercury concentrations as a result of foraging in marine, estuarine, or terrestrial food webs. The ecological impacts of differential foraging may be especially important for generalist species that exhibit high plasticity in foraging habitats, locations, or diet. Here, we investigate whether foraging habitat, sex, or fidelity to a foraging area impact blood mercury concentrations in western gulls (Larus occidentalis) from three colonies on the US west coast. Cluster analyses showed that nearly 70% of western gulls foraged primarily in ocean or coastal habitats, whereas the remaining gulls foraged in terrestrial and freshwater habitats. Gulls that foraged in ocean or coastal habitats for half or more of their foraging locations had 55% higher mercury concentrations than gulls that forage in freshwater and terrestrial habitats. Ocean-foraging gulls also had lower fidelity to a specific foraging area than freshwater and terrestrial-foraging gulls, but fidelity and sex were unrelated to gull blood mercury concentrations in all models. These findings support existing research that has described elevated mercury levels in species using aquatic habitats. Our analyses also demonstrate that gulls can be used to detect differences in contaminant exposure over broad geographic scales and across coarse habitat types, a factor that may influence gull health and persistence of other populations that forage across the land-sea gradient.

Hidden Markov models identify major movement modes in accelerometer and magnetometer data from four albatross species.

BACKGROUND: Inertial measurement units (IMUs) with high-resolution sensors such as accelerometers are now used extensively to study fine-scale behavior in a wide range of marine and terrestrial animals. Robust and practical methods are required for the computationally-demanding analysis of the resulting large datasets, particularly for automating classification routines that construct behavioral time series and time-activity budgets. Magnetometers are used increasingly to study behavior, but it is not clear how these sensors contribute to the accuracy of behavioral classification methods. Development of effective  classification methodology is key to understanding energetic and life-history implications of foraging and other behaviors. METHODS: We deployed accelerometers and magnetometers on four species of free-ranging albatrosses and evaluated the ability of unsupervised hidden Markov models (HMMs) to identify three major modalities in their behavior: 'flapping flight', 'soaring flight', and 'on-water'. The relative contribution of each sensor to classification accuracy was measured by comparing HMM-inferred states with expert classifications identified from stereotypic patterns observed in sensor data. RESULTS: HMMs provided a flexible and easily interpretable means of classifying behavior from sensor data. Model accuracy was high overall (92%), but varied across behavioral states (87.6, 93.1 and 91.7% for 'flapping flight', 'soaring flight' and 'on-water', respectively). Models built on accelerometer data alone were as accurate as those that also included magnetometer data; however, the latter were useful for investigating slow and periodic behaviors such as dynamic soaring at a fine scale. CONCLUSIONS: The use of IMUs in behavioral studies produces large data sets, necessitating the development of computationally-efficient methods to automate behavioral classification in order to synthesize and interpret underlying patterns. HMMs provide an accessible and robust framework for analyzing complex IMU datasets and comparing behavioral variation among taxa across habitats, time and space.

Seasonal variation of mercury contamination in Arctic seabirds: A pan-Arctic assessment.

Mercury (Hg) is a natural trace element found in high concentrations in top predators, including Arctic seabirds. Most current knowledge about Hg concentrations in Arctic seabirds relates to exposure during the summer breeding period when researchers can easily access seabirds at colonies. However, the few studies focused on winter have shown higher Hg concentrations during the non-breeding period than breeding period in several tissues. Hence, improving knowledge about Hg exposure during the non-breeding period is crucial to understanding the threats and risks encountered by these species year-round. We used feathers of nine migratory alcid species occurring at high latitudes to study bird Hg exposure during both the breeding and non-breeding periods. Overall, Hg concentrations during the non-breeding period were ~3 times higher than during the breeding period. In addition, spatial differences were apparent within and between the Atlantic and Pacific regions. While Hg concentrations during the non-breeding period were ~9 times and ~3 times higher than during the breeding period for the West and East Atlantic respectively, Hg concentrations in the Pacific during the non-breeding period were only ~1.7 times higher than during the breeding period. In addition, individual Hg concentrations during the non-breeding period for most of the seabird colonies were above 5 µg g-1 dry weight (dw), which is considered to be the threshold at which deleterious effects are observed, suggesting that some breeding populations might be vulnerable to non-breeding Hg exposure. Since wintering area locations, and migration routes may influence seasonal Hg concentrations, it is crucial to improve our knowledge about spatial ecotoxicology to fully understand the risks associated with Hg contamination in Arctic seabirds.

Similar foraging energetics of two sympatric albatrosses despite contrasting life histories and wind-mediated foraging strategies.

Understanding the environmental and behavioral factors that influence how organisms maintain energy balance can inform us about their potential resiliency to rapid environmental changes. Flexibility in maintaining energy balance is particularly important to long-lived, central-place foraging seabirds that are constrained when locating food for offspring in a dynamic ocean environment. To understand the role of environmental interactions, behavioral flexibility and morphological constraints on energy balance, we used doubly labeled water to measure the at-sea daily energy expenditure (DEE) of two sympatrically breeding seabirds, Campbell (Thalassarche impavida) and grey-headed (Thalassarchechrysostoma) albatrosses. We found that species and sexes had similar foraging costs, but DEE varied between years for both species and sexes during early chick rearing in two consecutive seasons. For both species, greater DEE was positively associated with larger proportional mass gain, lower mean wind speeds during water take-offs, greater proportions of strong tailwinds (>12 m s-1), and younger chick age. Greater proportional mass gains were marginally more costly in male albatrosses that already have higher wing loading. DEE was higher during flights with a greater proportion of strong headwinds for grey-headed albatrosses only. Poleward winds are forecasted to intensify over the next century, which may increase DEE for grey-headed albatrosses that heavily use this region during early chick rearing. Female Campbell albatrosses may be negatively affected by forecasted slackening winds at lower latitudes due to an expected greater reliance on less energy efficient sit-and-wait foraging strategies. Behavioral plasticity associated with environmental variation may influence future population responses to climate change of both species.

Author Correction: Important At-Sea Areas of Colonial Breeding Marine Predators on the Southern Patagonian Shelf.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Wintering in the Western Subarctic Pacific Increases Mercury Contamination of Red-Legged Kittiwakes.

Marine methylmercury concentrations vary geographically and with depth, exposing organisms to different mercury levels in different habitats. Red-legged kittiwakes (Rissa brevirostris), a specialist predator, forage on fish and invertebrates from the mesopelagic zone, a part of the ocean with elevated methylmercury concentrations. We used kittiwakes as bioindicators of MeHg concentrations in remote mesopelagic systems by examining how wintering distribution and habitat affected kittiwakes' mercury exposure. In 2011-2017, we sampled winter-grown feathers on St. George Island, Alaska, from birds equipped with geolocation loggers. We measured total mercury (THg) and nitrogen stable isotopes in nape and head feathers grown during winter, respectively. THg concentration of kittiwake nape feathers averaged 4.61 ± 0.97 µg/g dry weight. Hierarchical cluster analysis was used to classify winter habitats with remotely sensed environmental variables along each bird's track. Five habitat clusters were identified. Birds that spent more time in the Western Subarctic Gyre and those that wintered further south had elevated THg concentrations. In contrast to THg, trophic level varied annually but did not show strong spatial patterns. Our results documented spatial variability in THg exposure based on the oceanic wintering locations of red-legged kittiwakes and highlight their use as a bioindicator of MeHg across ocean basins.

Important At-Sea Areas of Colonial Breeding Marine Predators on the Southern Patagonian Shelf.

The Patagonian Shelf Large Marine Ecosystem supports high levels of biodiversity and endemism and is one of the most productive marine ecosystems in the world. Despite the important role marine predators play in structuring the ecosystems, areas of high diversity where multiple predators congregate remains poorly known on the Patagonian Shelf. Here, we used biotelemetry and biologging tags to track the movements of six seabird species and three pinniped species breeding at the Falkland Islands. Using Generalized Additive Models, we then modelled these animals' use of space as functions of dynamic and static environmental indices that described their habitat. Based on these models, we mapped the predicted distribution of animals from both sampled and unsampled colonies and thereby identified areas where multiple species were likely to overlap at sea. Maximum foraging trip distance ranged from 79 to 1,325 km. However, most of the 1,891 foraging trips by 686 animals were restricted to the Patagonian Shelf and shelf slope, which highlighted a preference for these habitats. Of the seven candidate explanatory covariates used to predict distribution, distance from the colony was retained in models for all species and negatively affected the probability of occurrence. Predicted overlap among species was highest on the Patagonian Shelf around the Falkland Islands and the Burdwood Bank. The predicted area of overlap is consistent with areas that are also important habitat for marine predators migrating from distant breeding locations. Our findings provide comprehensive multi-species predictions for some of the largest marine predator populations on the Patagonian Shelf, which will contribute to future marine spatial planning initiatives. Crucially, our findings highlight that spatially explicit conservation measures are likely to benefit multiple species, while threats are likely to impact multiple species.

Comparing imaging, acoustics, and radar to monitor Leach's storm-petrel colonies.

Seabirds are integral components of marine ecosystems and, with many populations globally threatened, there is a critical need for effective and scalable seabird monitoring strategies. Many seabird species nest in burrows, which can make traditional monitoring methods costly, infeasible, or damaging to nesting habitats. Traditional burrow occupancy surveys, where possible, can occur infrequently and therefore lead to an incomplete understanding of population trends. For example, in Oregon, during the last three decades there have been large changes in the abundance of Leach's storm-petrels (Hydrobates leucorhoa), which included drastic declines at some colonies. Unfortunately, traditional monitoring failed to capture the timing and magnitude of change, limiting managers' ability to determine causes of the decline and curtailing management options. New, easily repeatable methods of quantifying relative abundance are needed. For this study, we tested three methods of remote monitoring: passive acoustic monitoring, time-lapse cameras, and radar. Abundance indices derived from acoustics and imagery: call rates, acoustic energy, and counts were significantly related to traditional estimates of burrow occupancy of Leach's storm-petrels. Due to sampling limitations, we were unable to compare radar to burrow occupancy. Image counts were significantly correlated with all other indices, including radar, while indices derived from acoustics and radar were not correlated. Acoustic data likely reflect different aspects of the population and hold the potential for the further development of indices to disentangle phenology, attendance of breeding birds, and reproductive success. We found that image counts are comparable with standard methods (e.g., radar) in producing annual abundance indices. We recommend that managers consider a sampling scheme that incorporates both acoustics and imaging, but for sites inaccessible to humans, radar remains the sole option. Implementation of acoustic and camera based monitoring programs will provide much needed information for a vulnerable group of seabirds.

Microbial Ecology of the Western Gull (Larus occidentalis).

Avian species host diverse communities of microorganisms which have important roles in the life of birds, including increased metabolism, protection from disease, and immune system development. Along with high human populations and a diversity of human uses of coastal zones, anthropogenic food sources are becoming increasingly available to some species, including gulls. Anthropogenic associations increase the likelihood of encountering foreign or pathogenic bacteria. Diseases in birds caused by bacteria are a substantial source of avian mortality; therefore, it is essential to characterize the microbiome of seabirds. Here, we determined both core and environmentally derived microbial communities of breeding western gulls (Larus occidentalis) from six colonies in California and Oregon. Using DNA extracted from bacterial swabs of the bill, cloaca, and feet of gulls, 16S rRNA gene sequencing was performed targeting the V4 region. We identified a total of 8542 operational taxonomic units (OTUs) from 75 gulls. Sixty-eight OTUs were identified in gulls from all six colonies with the greatest representation from phyla's of Firmicutes, Proteobacteria, Bacteroidetes, and Actinobacteria. Overall, microbial richness based on Chao's Abundance-based Coverage Estimator (ACE) index was similar for all colonies (mean = 2347 OTUs) with the smallest coastal colonies having the highest richness (mean = 2626 OTUs) and the largest colonies, located farther off-shore, having the lowest (mean = 2068 OTUs). This survey represents the most in-depth assessment to date of microbes associated with western gulls, and the first study to identify both species-specific and environmentally derived bacteria across multiple populations.

Habitat use and spatial fidelity of male South American sea lions during the nonbreeding period.

Conditions experienced during the nonbreeding period have profound long-term effects on individual fitness and survival. Therefore, knowledge of habitat use during the nonbreeding period can provide insights into processes that regulate populations. At the Falkland Islands, the habitat use of South American sea lions (Otaria flavescens) during the nonbreeding period is of particular interest because the population is yet to recover from a catastrophic decline between the mid-1930s and 1965, and nonbreeding movements are poorly understood. Here, we assessed the habitat use of adult male (n = 13) and juvenile male (n = 6) South American sea lions at the Falkland Islands using satellite tags and stable isotope analysis of vibrissae. Male South American sea lions behaved like central place foragers. Foraging trips were restricted to the Patagonian Shelf and were typically short in distance and duration (127 ± 66 km and 4.1 ± 2.0 days, respectively). Individual male foraging trips were also typically characterized by a high degree of foraging site fidelity. However, the isotopic niche of adult males was smaller than juvenile males, which suggested that adult males were more consistent in their use of foraging habitats and prey over time. Our findings differ from male South American sea lions in Chile and Argentina, which undertake extended movements during the nonbreeding period. Hence, throughout their breeding range, male South American sea lions have diverse movement patterns during the nonbreeding period that intuitively reflects differences in the predictability or accessibility of preferred prey. Our findings challenge the long-standing notion that South American sea lions undertake a winter migration away from the Falkland Islands. Therefore, impediments to South American sea lion population recovery likely originate locally and conservation measures at a national level are likely to be effective in addressing the decline and the failure of the population to recover.

Classification of Animal Movement Behavior through Residence in Space and Time.

Identification and classification of behavior states in animal movement data can be complex, temporally biased, time-intensive, scale-dependent, and unstandardized across studies and taxa. Large movement datasets are increasingly common and there is a need for efficient methods of data exploration that adjust to the individual variability of each track. We present the Residence in Space and Time (RST) method to classify behavior patterns in movement data based on the concept that behavior states can be partitioned by the amount of space and time occupied in an area of constant scale. Using normalized values of Residence Time and Residence Distance within a constant search radius, RST is able to differentiate behavior patterns that are time-intensive (e.g., rest), time & distance-intensive (e.g., area restricted search), and transit (short time and distance). We use grey-headed albatross (Thalassarche chrysostoma) GPS tracks to demonstrate RST's ability to classify behavior patterns and adjust to the inherent scale and individuality of each track. Next, we evaluate RST's ability to discriminate between behavior states relative to other classical movement metrics. We then temporally sub-sample albatross track data to illustrate RST's response to less resolved data. Finally, we evaluate RST's performance using datasets from four taxa with diverse ecology, functional scales, ecosystems, and data-types. We conclude that RST is a robust, rapid, and flexible method for detailed exploratory analysis and meta-analyses of behavioral states in animal movement data based on its ability to integrate distance and time measurements into one descriptive metric of behavior groupings. Given the increasing amount of animal movement data collected, it is timely and useful to implement a consistent metric of behavior classification to enable efficient and comparative analyses. Overall, the application of RST to objectively explore and compare behavior patterns in movement data can enhance our fine- and broad- scale understanding of animal movement ecology.

Regional variability in diving physiology and behavior in a widely distributed air-breathing marine predator, the South American sea lion (Otaria byronia).

Our understanding of how air-breathing marine predators cope with environmental variability is limited by our inadequate knowledge of their ecological and physiological parameters. Because of their wide distribution along both coasts of the sub-continent, South American sea lions (Otaria byronia) provide a valuable opportunity to study the behavioral and physiological plasticity of a marine predator in different environments. We measured the oxygen stores and diving behavior of South American sea lions throughout most of its range, allowing us to demonstrate that diving ability and behavior vary across its range. We found no significant differences in mass-specific blood volumes of sea lions among field sites and a negative relationship between mass-specific oxygen storage and size, which suggests that exposure to different habitats and geographical locations better explains oxygen storage capacities and diving capability in South American sea lions than body size alone. The largest animals in our study (individuals from Uruguay) were the shallowest and shortest duration divers, and had the lowest mass-specific total body oxygen stores, while the deepest and longest duration divers (individuals from southern Chile) had significantly larger mass-specific oxygen stores, despite being much smaller animals. Our study suggests that the physiology of air-breathing diving predators is not fixed, but that it can be adjusted, to a certain extent, depending on the ecological setting and or habitat. These adjustments can be thought of as a 'training effect': as the animal continues to push its physiological capacity through greater hypoxic exposure, its breath-holding capacity increases.

Pup Vibrissae Stable Isotopes Reveal Geographic Differences in Adult Female Southern Sea Lion Habitat Use during Gestation.

Individuals within populations often differ substantially in habitat use, the ecological consequences of which can be far reaching. Stable isotope analysis provides a convenient and often cost effective means of indirectly assessing the habitat use of individuals that can yield valuable insights into the spatiotemporal distribution of foraging specialisations within a population. Here we use the stable isotope ratios of southern sea lion (Otaria flavescens) pup vibrissae at the Falkland Islands, in the South Atlantic, as a proxy for adult female habitat use during gestation. A previous study found that adult females from one breeding colony (Big Shag Island) foraged in two discrete habitats, inshore (coastal) or offshore (outer Patagonian Shelf). However, as this species breeds at over 70 sites around the Falkland Islands, it is unclear if this pattern is representative of the Falkland Islands as a whole. In order to characterize habitat use, we therefore assayed carbon (δ13C) and nitrogen (δ15N) ratios from 65 southern sea lion pup vibrissae, sampled across 19 breeding colonies at the Falkland Islands. Model-based clustering of pup isotope ratios identified three distinct clusters, representing adult females that foraged inshore, offshore, and a cluster best described as intermediate. A significant difference was found in the use of inshore and offshore habitats between West and East Falkland and between the two colonies with the largest sample sizes, both of which are located in East Falkland. However, habitat use was unrelated to the proximity of breeding colonies to the Patagonian Shelf, a region associated with enhanced biological productivity. Our study thus points towards other factors, such as local oceanography and its influence on resource distribution, playing a prominent role in inshore and offshore habitat use.

Disentangling the cause of a catastrophic population decline in a large marine mammal.

Considerable uncertainties often surround the causes of long-term changes in population abundance. One striking example is the precipitous decline of southern sea lions (SSL; Otariaflavescens) at the Falkland Islands, from 80 555 pups in the mid 1930s to just 5506 pups in 1965. Despite an increase in SSL abundance over the past two decades, the population has not recovered, with the number of pups born in 2014 (minimum 4443 pups) less than 6% of the 1930s estimate. The order-of-magnitude decline is primarily attributed to commercial sealing in Argentina. Here, we test this established paradigm and alternative hypotheses by assessing (1) commercial sealing at the Falkland Islands, (2) winter migration of SSL from the Falkland Islands to Argentina, (3) whether the number of SSL in Argentina could have sustained the reported level of exploitation, and (4) environmental change. The most parsimonious hypothesis explaining the SSL population decline was environmental change. Specifically, analysis of 160 years of winter sea surface temperatures revealed marked changes, including a period of warming between 1930 and 1950 that was consistent with the period of SSL decline. Sea surface temperature changes likely influenced the distribution or availability of SSL prey and impacted its population dynamics. We suggest that historical harvesting may not always be the "smoking gun" as is often purported. Rather, our conclusions support the growing evidence for bottom-up forcing on the abundance of species at lower trophic levels (e.g., plankton and fish) and resulting impacts on higher trophic levels across a broad range of ecosystems.

Wintering North Pacific black-legged kittiwakes balance spatial flexibility and consistency.

BACKGROUND: Marine environments are inherently dynamic, yet marine predators are often long-lived and employ strategies where consistency, individual specialization, routine migrations, and spatial memory are key components to their foraging and life-history strategies. Intrinsic determinates of animal movements are linked to physiological and life-history traits (e.g. sex, colony, experience), while extrinsic influences occur as the result of an animal's interactions with either other animals or the environment (e.g. prey availability, weather, competition). Knowledge of the factors affecting animal movements is critical to understand energetic bottlenecks and population dynamics. Here, we attempt to understand the interaction of some of these factors on the winter distributions of a surface-feeding seabird in the North Pacific. Between 2008 and 2011, we tracked 99 black-legged kittiwakes breeding at St. Paul and St. George in the Pribilof Islands, Alaska using geolocation loggers. We tested for colony and sex differences in winter distributions, and individual spatial fidelity over two consecutive winters of 17 individuals. Then we linked tracking data to associated environmental conditions as proxies of prey availability (e.g. sea surface temperature, mesoscale eddies, chlorophyll a, and wind) to understand their influence on kittiwake space use at an ocean basin scale. RESULTS: Black-legged kittiwakes from both Pribilof Islands primarily wintered in pelagic sub-arctic waters, however, distributions spanned seven ecoregions of the North Pacific. There was a high degree of similarity in area use of birds from the two closely situated colonies and between sexes. Birds tracked for two consecutive years showed higher fidelity to wintering areas than occurred at random. Annual changes were apparent, as distributions were further north in 2009/10 than 2008/09 or 2010/11. This occurred because 70 % of birds remained in the Bering Sea in the fall of 2009, which corresponded with lower October sea surface temperatures than the other two years. CONCLUSIONS: Although individuals returned to wintering areas in consecutive years, our results suggest that under current conditions individual black-legged kittiwakes have a high capacity to alter winter distributions.

Body size affects individual winter foraging strategies of thick-billed murres in the Bering Sea.

Foraging and migration often require different energetic and movement strategies. Though not readily apparent, constraints during one phase might influence the foraging strategies observed in another. For marine birds that fly and dive, body size constraints likely present a trade-off between foraging ability and migration as smaller bodies reduce flight costs, whereas larger bodies are advantageous for diving deeper. This study examines individual wintering strategies of deep diving thick-billed murres (Uria lomvia) breeding at three colonies in the south-eastern Bering Sea: St Paul, St George and Bogoslof. These colonies, arranged north to south, are located such that breeding birds forage in a gradient from shelf to deep-water habitats. We used geolocation time-depth recorders and stable isotopes from feathers to determine differences in foraging behaviour and diet of murres during three non-breeding periods, 2008-2011. Body size was quantified by a principal component analysis (wing, culmen, head+bill and tarsus length). A hierarchical cluster analysis identified winter foraging strategies based on individual movement, diving behaviour and diet (inferred from stable isotopes). Structural body size differed by breeding island. Larger birds from St Paul had higher wing loading than smaller birds from St George. Larger birds, mainly from St Paul, dove to deeper depths, spent more time in the Bering Sea, and likely consumed higher trophic-level prey in late winter. Three winter foraging strategies were identified. The main strategy, employed by small birds from all three breeding colonies in the first 2 years, was characterized by high residency areas in the North Pacific south of the Aleutians and nocturnal diving. In contrast, 31% of birds from St Paul remained in the Bering Sea and foraged mainly during the day, apparently feeding on higher trophic-level prey. Throat feather stable isotopes indicated that individuals exhibited flexibility in the use of this colony-specific foraging strategy. The third strategy only occurred in 2010/2011, when birds dove more and deeper, suggesting limited prey resources. Foraging strategies partitioned with respect to annual differences, presumably in response to shifts in distribution of prey, and were linked to body size. The presence of a colony-specific wintering strategy suggests the potential for overwinter survival differences between these populations.