Predicting top predator habitats in the Southwest Indian Ocean

Top predators need to develop optimal strategies of resources and habitats utilization in order to optimize their foraging success. At the individual scale, a predator has to maximize his intake of food while minimizing his cost of foraging to optimize his energetic gain. At the ecosystem scale, we hypothesized that foraging strategies of predators also respond to their general energetic constraints. Predators with energetically costly lifestyles may be constrained to select high quality habitats whereas more phlegmatic predators may occupy both low and high quality habitats. The objectives of this study were 1) to investigate predator responses to heterogeneity in habitat quality with reference to their energetic strategies and 2) to evaluate their responses to contemporaneous versus averaged habitat quality. We collected cetacean and seabird data from an aerial survey in the Southwest Indian Ocean, a region characterized by heterogeneous oceanographic conditions. We classified cetaceans and seabirds into energetic guilds and described their habitats using remotely sensed covariates at contemporaneous and time‐averaged resolutions and static covariates. We used generalized additive models to predict their habitats at the regional scale. Strategies of habitat utilization appeared in accordance with predators energetic constraints. Cetaceans responded to the heterogeneity in habitat quality, with higher densities predicted in more productive areas. However, the costly Delphininae appeared to be more dependent on habitat quality (showing a 1‐to‐13 ratio between the lowest and highest density sectors) than the more phlegmatic sperm and beaked whales (showing only a 1‐to‐3 ratio). For seabirds, predictions primarily reflected colony locations, although the colony effect was stronger for costly seabirds. Moreover, our results suggest that predators may respond better to persistent oceanographic features. To provide a third dimension to habitat quality, cetacean strategies of utilization of the vertical habitat could be related to the distribution of micronekton in the water column.     

Hitting the buffers: conspecific aggression undermines benefits of colonial breeding under adverse conditions

Colonial breeding in birds is widely considered to benefit individuals through enhanced protection against predators or transfer of information about foraging sites. This view, however, is largely based on studies of seabirds carried out under favourable conditions. Recent breeding failures at many seabird colonies in the UK provide an opportunity to re-examine costs and benefits of coloniality under adverse conditions. Common guillemots Uria aalge are highly colonial cliff-nesting seabirds with very flexible parental care. Although the single chick is normally never left alone, more than 50 per cent of offspring were left unattended at a North Sea colony in 2007, apparently because poor conditions forced both parents to forage simultaneously. Contrary to expectation, unattended chicks were not killed by avian predators. Rather, although non-breeders and failed breeders sometimes provided alloparental care, unattended chicks were frequently attacked by breeding guillemots at neighbouring sites, often with fatal consequences. These results highlight a previously unsuspected trade-off between provisioning chicks and avoiding conspecific attacks, and indicate that understanding how environmental conditions affect social dynamics is crucial to interpreting costs and benefits of colonial breeding.     

Circumpolar dynamics of a marine top‐predator track ocean warming rates

Global warming is a nonlinear process, and temperature may increase in a stepwise manner. Periods of abrupt warming can trigger persistent changes in the state of ecosystems, also called regime shifts. The responses of organisms to abrupt warming and associated regime shifts can be unlike responses to periods of slow or moderate change. Understanding of nonlinearity in the biological responses to climate warming is needed to assess the consequences of ongoing climate change. Here, we demonstrate that the population dynamics of a long‐lived, wide‐ranging marine predator are associated with changes in the rate of ocean warming. Data from 556 colonies of black‐legged kittiwakes Rissa tridactyla distributed throughout its breeding range revealed that an abrupt warming of sea‐surface temperature in the 1990s coincided with steep kittiwake population decline. Periods of moderate warming in sea temperatures did not seem to affect kittiwake dynamics. The rapid warming observed in the 1990s may have driven large‐scale, circumpolar marine ecosystem shifts that strongly affected kittiwakes through bottom‐up effects. Our study sheds light on the nonlinear response of a circumpolar seabird to large‐scale changes in oceanographic conditions and indicates that marine top predators may be more sensitive to the rate of ocean warming rather than to warming itself.     

Combining Methods to Describe Important Marine Habitats for Top Predators: Application to Identify Biological Hotspots in Tropical Waters

In tropical waters resources are usually scarce and patchy, and predatory species generally show specific adaptations for foraging. Tropical seabirds often forage in association with sub-surface predators that create feeding opportunities by bringing prey close to the surface, and the birds often aggregate in large multispecific flocks. Here we hypothesize that frigatebirds, a tropical seabird adapted to foraging with low energetic costs, could be a good predictor of the distribution of their associated predatory species, including other seabirds (e.g. boobies, terns) and subsurface predators (e.g., dolphins, tunas). To test this hypothesis, we compared distribution patterns of marine predators in the Mozambique Channel based on a long-term dataset of both vessel- and aerial surveys, as well as tracking data of frigatebirds. By developing species distribution models (SDMs), we identified key marine areas for tropical predators in relation to contemporaneous oceanographic features to investigate multi-species spatial overlap areas and identify predator hotspots in the Mozambique Channel. SDMs reasonably matched observed patterns and both static (e.g. bathymetry) and dynamic (e.g. Chlorophyll a concentration and sea surface temperature) factors were important explaining predator distribution patterns. We found that the distribution of frigatebirds included the distributions of the associated species. The central part of the channel appeared to be the best habitat for the four groups of species considered in this study (frigatebirds, brown terns, boobies and sub-surface predators).     

Sex-Specific Habitat Utilization and Differential Breeding Investments in Christmas Island Frigatebirds throughout the Breeding Cycle

In seabirds, equal bi-parental care is the rule, as it is considered crucial for raising chicks successfully because seabirds forage in an environment with unpredictable and highly variable food supply. Frigatebirds forage in poor tropical waters, yet males reduce and even stop parental care soon after chick brooding, leaving the female to provision the chick alone for an extended fledging period. Using bird-borne tracking devices, male and female Christmas Island Frigatebirds (Fregata andrewsi) were investigated during the brooding, late chick rearing and post-fledging period to examine whether sexes exhibit foraging strategies that may be linked to differential breeding investments. During brooding, males and females showed similar foraging behaviour under average marine productivity of oceanic waters close to the colony, but males shifted to more distant and more productive habitats when conditions deteriorated to continue with reduced chick provisioning. During the late chick rearing period, females progressively increased their foraging range to the more distant but productive marine areas that only males had visited during brooding. Birds spent the non-breeding period roosting in highly productive waters of the Sunda Shelf. The sex-specific utilisation of three different foraging habitats with different primary productivity (oceanic, coastal, and shelf areas) allowed for temporal and spatial segregation in the exploitation of favourable habitats which seems to enable each sex to optimise its foraging profitability. In addition, post-fledging foraging movements of females suggest a biennial breeding cycle, while limited information on males suggests the possibility of an annual breeding cycle.     

Facilitative interactions among the pelagic community of temperate migratory terns,tunas and dolphins

We studied habitat and behavioral interactions among the marine community of top pelagic predators over the Atlantic continental shelf, observed from shipboard surveys off the northeastern United States. We hypothesized that foraging seabirds, specifically common terns Sterna hirundo and roseate terns S. dougallii, associate positively with the distribution and abundance of large, easily‐detected subsurface marine predators, as a result of facilitative interactions. Few rigorous tests examine the effect of interspecific interactions on seabird distributions, though many papers note the importance of environmental influences. Fewer, still, assess facilitation, defined as positive interactions among multiple taxa (birds, fish, mammals), where individuals use foraging neighbors for improved prey detection or enhanced prey availability. Our use of spatiotemporal‐structured Bayesian hierarchical models allowed us to test for fine‐scale associations of common and roseate terns with aggregations of tunas and cetaceans, and with standard oceanographic parameters. High tern abundance was linked to relatively high tuna densities and low dolphin densities, as well as high sea surface temperatures, shallow water, and proximity to shore. The fact that terns foraged when in the presence of tunas or relatively dense dolphin pods supports our hypothesis that the mechanism behind this spatial association involves positive interspecific interactions (attraction), a phenomenon that has been described but rarely quantified. This study reveals that community structure depends on static and dynamic covariates alike, ranging from bathymetry to the movement of other guild members. To improve the efficacy of predictive modeling, ecosystem approaches to applied conservation management should integrate not only habitat, but also behavior and community factors into analyses, especially in the case of marine spatial planning.     

Prey capture and selection throughout the breeding season in a deep‐diving generalist seabird,the thick‐billed murre

Generalist seabirds forage on a variety of prey items providing the opportunity to monitor diverse aquatic fauna simultaneously. For example, the coupling of prey consumption rates and movement patterns of generalist seabirds might be used to create three‐dimensional prey distribution maps (‘preyscapes’) for multiple prey species in the same region. However, the complex interaction between generalist seabird foraging behaviour and the various prey types clouds the interpretation of such preyscapes, and the mechanisms underlying prey selection need to be understood before such an application can be realized. Central place foraging theory provides a theoretical model for understanding such selectivity by predicting that larger prey items should be 1) selected farther from the colony and 2) for chick‐feeding compared with self‐feeding, but these predictions remain untested on most seabird species. Furthermore, rarely do we know how foraging features such as handling time, capture methods or choice of foraging location varies among prey types. We used three types of animal‐borne biologgers (camera loggers, GPS and depth‐loggers) to examine how a generalist Arctic seabird, the thick‐billed murre Uria lomvia, selects and captures their prey throughout the breeding season. Murres captured small prey at all phases of a dive, including while descending and ascending, but captured large fish mostly while ascending, with considerably longer handling times. Birds captured larger prey and dove deeper during chick‐rearing. As central place foraging theory predicted, birds travelling further also brought bigger prey items for their chick. The location of a dive (distance from colony and distance to shore) best explained which prey type was the most likely to get caught in a dive, and we created a preyscape surrounding our study colony. We discuss how these findings might aid the use of generalist seabirds as bioindicators.     

A review of the occurrence of inter‐colony segregation of seabird foraging areas and the implications for marine environmental impact assessment

Understanding the determinants of species’ distributions is a fundamental aim in ecology and a prerequisite for conservation but is particularly challenging in the marine environment. Advances in bio‐logging technology have resulted in a rapid increase in studies of seabird movement and distribution in recent years. Multi‐colony studies examining the effects of intra‐ and inter‐colony competition on distribution have found that several species exhibit inter‐colony segregation of foraging areas, rather than overlapping distributions. These findings are timely given the increasing rate of human exploitation of marine resources and the need to make robust assessments of likely impacts of proposed marine developments on biodiversity. Here we review the occurrence of foraging area segregation reported by published tracking studies in relation to the density‐dependent hinterland (DDH) model, which predicts that segregation occurs in response to inter‐colony competition, itself a function of colony size, distance from the colony and prey distribution. We found that inter‐colony foraging area segregation occurred in 79% of 39 studies. The frequency of occurrence was similar across the four seabird orders for which data were available, and included species with both smaller (10–100 km) and larger (100–1000 km) foraging ranges. Many predictions of the DDH model were confirmed, with examples of segregation in response to high levels of inter‐colony competition related to colony size and proximity, and enclosed landform restricting the extent of available habitat. Moreover, as predicted by the DDH model, inter‐colony overlap tended to occur where birds aggregated in highly productive areas, often remote from all colonies. The apparent prevalence of inter‐colony foraging segregation has important implications for assessment of impacts of marine development on protected seabird colonies. If a development area is accessible from multiple colonies, it may impact those colonies much more asymmetrically than previously supposed. Current impact assessment approaches that do not consider spatial inter‐colony segregation will therefore be subject to error. We recommend the collection of tracking data from multiple colonies and modelling of inter‐colony interactions to predict colony‐specific distributions.     

Climate oscillation and the invasion of alien species influence the oceanic distribution of seabirds

Spatial and temporal distribution of seabird transiting and foraging at sea is an important consideration for marine conservation planning. Using at‐sea observations of seabirds (n = 317), collected during the breeding season from 2012 to 2016, we built boosted regression tree (BRT) models to identify relationships between numerically dominant seabird species (red‐footed booby, brown noddy, white tern, and wedge‐tailed shearwater), geomorphology, oceanographic variability, and climate oscillation in the Chagos Archipelago. We documented positive relationships between red‐footed booby and wedge‐tailed shearwater abundance with the strength in the Indian Ocean Dipole, as represented by the Dipole Mode Index (6.7% and 23.7% contribution, respectively). The abundance of red‐footed boobies, brown noddies, and white terns declined abruptly with greater distance to island (17.6%, 34.1%, and 41.1% contribution, respectively). We further quantified the effects of proximity to rat‐free and rat‐invaded islands on seabird distribution at sea and identified breaking point distribution thresholds. We detected areas of increased abundance at sea and habitat use‐age under a scenario where rats are eradicated from invaded nearby islands and recolonized by seabirds. Following rat eradication, abundance at sea of red‐footed booby, brown noddy, and white terns increased by 14%, 17%, and 3%, respectively, with no important increase detected for shearwaters. Our results have implication for seabird conservation and island restoration. Climate oscillations may cause shifts in seabird distribution, possibly through changes in regional productivity and prey distribution. Invasive species eradications and subsequent island recolonization can lead to greater access for seabirds to areas at sea, due to increased foraging or transiting through, potentially leading to distribution gains and increased competition. Our approach predicting distribution after successful eradications enables anticipatory threat mitigation in these areas, minimizing competition between colonies and thereby maximizing the risk of success and the conservation impact of eradication programs.     

Colony size and foraging range in seabirds

The reasons for variation in group size among animal species remain poorly understood. Using ‘Ashmole's halo’ hypothesis of food depletion around colonies, we predict that foraging range imposes a ceiling on the maximum colony size of seabird species. We tested this with a phylogenetic comparative study of 43 species of seabirds (28 262 colonies), and investigated the interspecific correlation between colony size and foraging ranges. Foraging range showed weak relationships with the low percentiles of colony size of species, but the strength of the association increased for larger percentiles, peaking at the maximum colony sizes. To model constraints on the functional relationship between the focal traits, we applied a quantile regression based on maximum colony size. This showed that foraging range imposes a constraint to species’ maximum colony sizes with a slope around 2. This second‐order relationship is expected from the equation of the area of a circle. Thus, our large dataset and innovative statistical approach shows that foraging range imposes a ceiling on seabird colony sizes, providing strong support to the hypothesis that food availability is an important regulator of seabird populations.     

The diet of Common Guillemot Uria aalge chicks provides evidence of changing prey communities in the North Sea

Local differences in feeding conditions have been suggested as a cause of regional variation in seabird demography but multi‐colony comparisons of diet are rare. In UK waters the main fish eaten by seabirds during the breeding season belong to three families: Ammodytidae, Clupeidae and Gadidae. Climate change and fishing are affecting these fish stocks and so probably impact on predators such as seabirds. We used standardized observations of prey brought in for chicks to make the first integrated assessment of the diet of Common Guillemot Uria aalge chicks at a UK scale. Chick diet varied markedly among the 23 colonies sampled between 2006 and 2011. Sandeels (Ammodytidae), probably Lesser Sandeels Ammodytes marinus, were the commonest prey. Their contribution to the diet varied both latitudinally and among marine regions, with the proportion significantly higher for a given latitude on the west coast compared to the east. The non‐sandeel component of the diet showed latitudinal changes, with small clupeids, probably Sprats Sprattus sprattus, predominant at southern colonies whereas juvenile gadids were the main alternative to sandeels in the north. Comparison of our Guillemot chick diet with data collected 15–30 years earlier suggests that the proportion of sandeels in the diet has decreased at colonies bordering the North Sea. No significant change was apparent in Atlantic colonies but historical data were limited. The early years of our study coincided with a population explosion of Snake Pipefish Entelurus aequoreus in the Northeast Atlantic and North Sea. Pipefish were recorded in Guillemot chick diet at several northern and northwestern colonies in 2006 and 2007 but have been absent since 2009. Spatial and temporal variation in chick diet accorded broadly with patterns expected as a result of rising sea temperatures and impacts of fishing. Guillemot chick diet could potentially be a useful indicator of changes in the distribution and abundance of forage fish.     

Large‐scale oceanographic fluctuations drive Antarctic petrel survival and reproduction

Polar Regions are experiencing environmental changes at unprecedented rates. These changes can spread throughout entire food webs from lower trophic levels to apex predators. As many top predators forage over large areas, these indirect effects may be associated with large‐scale patterns of climate variability. Using global climate indices that are known to impact the Southern Ocean ecosystem (the El Niño Southern Oscillation and Antarctic Oscillation Indices) we assessed their efficacy to predict variation in the demographic parameters of Antarctic seabirds. First, we used a long‐term dataset on adult survival (estimated from capture–mark–recapture data) and reproduction of Antarctic petrel Thalassoica antarctica, from the largest known breeding colony (Svarthamaren, Dronning Maud Land) and examined whether large‐scale oceanographic fluctuations impact survival and reproduction. Second, we conducted an exhaustive literature review to determine whether the effects of large‐scale environmental variability on Antarctic seabirds have a coherent fingerprint across the Antarctic continent and nearby islands. We found that most of the variation in both reproductive success, timing of hatching, and survival of Antarctic petrels can be accurately modeled using the two modes of large‐scale climate variability in Antarctica. The literature review, combined with the results from our field study, suggests that while the anticipated trends in the global patterns of climatic variability will generally have detrimental effects on populations of top predators in the Southwest Atlantic, these conclusions cannot be extrapolated to all seabird populations in Antarctica without additional data.     

Seabird predation by great skuas Stercorarius skua– intra‐specific competition for food?

Competition for food is widely cited as an important cost of coloniality among birds and much of the evidence in support of this hypothesis comes from studies of colonial piscivorous seabirds. However, for generalist seabirds able to switch between different prey types, the role of food availability in relation to colony size is unclear. Here we investigate patterns of the consumption of seabird prey in relation to colony size in a generalist seabird, the great skua Stercorarius skua, in Shetland, UK. At the population level skuas feed mainly on sandeels Ammodytes marinus and fishery discards, but respond to declines in fish availability to facultatively prey on other seabirds. By comparing the consumption of seabirds among seven different sized colonies, including one colony with artificially reduced numbers of skuas (Fair Isle), we investigate whether consumption of seabird prey is influenced by skua population size, while simultaneously measuring seabird prey availability. Data from five years also enables us to investigate the influence of annual variation in environmental conditions on seabird consumption. Using measures of body condition and reproductive performance we investigate the consequences of living in different sized colonies, which may provide insight into ultimate costs of nesting at high population density. Skua diets varied among colonies and the proportion of seabird prey in the diet was inversely related to skua colony size, despite similar per capita numbers of seabirds across colonies. At the colony where their numbers were artificially suppressed, skuas consumed a greater proportion of seabirds per capita. Highly significant year effects in seabird predation were observed but the pattern among colonies remained consistent over time. Two measures of adult body condition (pectoral muscle index and mean corpuscular volume) revealed that adult great skuas were in poorer condition at the largest colony (Foula), but reproductive performance did not alter significantly among colonies. This study provides evidence that intra‐specific competition among skuas may limit opportunities for obtaining seabird prey, which may be particularly important during periods of poor availability of sandeels and fishery discards, and has implications for assessing the impact of skuas on seabird populations.     

Divergent diving behavior during short and long trips of a bimodal forager,the little auk Alle alle

The purpose of this study was to characterize for the first time seabird diving behavior during bimodal foraging. Little auks Alle alle, small zooplanktivorous Alcids of the High Arctic, have recently been shown to make foraging trips of short and long duration. Because short (ST) and long trips (LT) are thought to occur in different locations and serve different purposes (chick‐ and self‐feeding, respectively) we hypothesized that foraging differences would be apparent, both in terms of water temperature and diving characteristics. Using Time Depth Recorders (TDRs), we tested this hypothesis at three colonies along the Greenland Sea with contrasting oceanographic conditions. We found that diving behavior generally differed between ST and LT. However, the magnitude of the disparity in diving characteristics depended on local foraging conditions. At the study site where conditions were favorable, diving behavior differed only to a small degree between LT and ST. Together with a lack of difference in diving depth and ocean temperature, this indicates that these birds did not increase their foraging effort during ST nor did they travel long distances to seek out more profitable prey. In contrast, where local foraging conditions were poor, birds increased their diving effort substantially to collect a chick meal during ST as indicated by longer, more U‐shaped dives with slower ascent rates and shorter resting times (post‐dive intervals and extended surface pauses). In addition, large differences in diving depth and ocean temperature indicate that birds forage on different prey species and utilize different foraging areas during LT, which may be up to 200 km away from the colony. Continued warming and deteriorating near‐colony foraging conditions may have energetic consequences for little auks breeding in the eastern Greenland Sea.     

Environmental heterogeneity amplifies behavioural response to a temporal cycle

Resource acquisition is integral to maximise fitness, however in many ecosystems this requires adaptation to resource abundance and distributions that seldom stay constant. For predators, prey availability can vary at fine spatial and temporal scales as a result of changes in the physical environment, and therefore selection should favour individuals that can adapt their foraging behaviour accordingly. The tidal cycle is a short, yet predictable, temporal cycle, which can influence prey availability at temporal scales relevant to movement decisions. Here, we ask whether black‐legged kittiwakes Rissa tridactyla can adjust their foraging habitat selection according to the tidal cycle using GPS tracking studies at three sites of differing environmental heterogeneity. We used a hidden Markov model to classify kittiwake behaviour, and analysed habitat selection during foraging. As expected for a central‐place forager, we found that kittiwakes preferred to forage nearer to the breeding colony. However, we also show that habitat selection changed over the 12.4‐h tidal cycle, most likely because of changes in resource availability. Furthermore, we observed that environmental heterogeneity was associated with amplified changes in kittiwake habitat selection over the tidal cycle, potentially because environmental heterogeneity drives greater resource variation. Both predictable cycles and environmental heterogeneity are ubiquitous. Our results therefore suggest that, together, predictable cycles and environmental heterogeneity may shape predator behaviour across ecosystems.     

The jellyfish buffet: jellyfish enhance seabird foraging opportunities by concentrating prey

High levels of jellyfish biomass have been reported in marine ecosystems around the world, but understanding of their ecological role remains in its infancy. Jellyfish are generally thought to have indirect negative impacts on higher trophic-level predators, through changes in lower trophic pathways. However, high densities of jellyfish in the water column may affect the foraging behaviour of marine predators more directly, and the effects may not always be negative. Here, we present novel observations of a diving seabird, the thick-billed murre, feeding on fish aggregating among the long tentacles of large jellyfish, by using small video loggers attached to the birds. We show that the birds encountered large jellyfish, Chrysaora melanaster, during most of their dives, commonly fed on fish associated with jellyfish, and appeared to specifically target jellyfish with a high number of fish aggregating in their tentacles, suggesting the use of jellyfish may provide significant energetic benefits to foraging murres. We conclude that jellyfish provide feeding opportunities for diving seabirds by concentrating forage fish, and that the impacts of jellyfish on marine ecosystems are more complex than previously anticipated and may be beneficial to seabirds.     

An energetic correlate between colony size and foraging effort in seabirds, an example of the Adélie penguin Pygoscelis adeliae

Central-place foraging seabirds alter the availability of their prey around colonies, forming a "halo" of reduced prey access that ultimately constrains population size. This has been indicated indirectly by an inverse correlation between colony size and reproductive success, numbers of conspecifics at other colonies within foraging range, foraging effort (i.e. trip duration), diet quality and colony growth rate. Although ultimately mediated by density dependence relative to food through intraspecific exploitative or interference competition, the proximate mechanism involved has yet to be elucidated. Herein, we show that Adélie penguin Pygoscelis adeliae colony size positively correlates to foraging trip duration and metabolic rate, that the metabolic rate while foraging may be approaching an energetic ceiling for birds at the largest colonies, and that total energy expended increases with trip duration although uncompensated by increased mass gain. We propose that a competition-induced reduction in prey availability results in higher energy expenditure for birds foraging in the halo around large colonies, and that to escape the halo a bird must increase its foraging distance. Ultimately, the total energetic cost of a trip determines the maximum successful trip distance, as on longer trips food acquired is used more for self maintenance than for chick provisioning. When the net cost of foraging trips becomes too high, with chicks receiving insufficient food, chick survival suffers and subsequent colony growth is limited. Though the existence of energetic studies of the same species at multiple colonies is rare, because foraging metabolic rate increases with colony size in at least two other seabird species, we suggest that an energetic constraint to colony size may generally apply to other seabirds.     

Feather corticosterone reveals stress associated with dietary changes in a breeding seabird

Changes in climate and anthropogenic pressures might affect the composition and abundance of forage fish in the world's oceans. The junk‐food hypothesis posits that dietary shifts that affect the quality (e.g., energy content) of food available to marine predators may impact their physiological state and consequently affect their fitness. Previously, we experimentally validated that deposition of the adrenocortical hormone, corticosterone, in feathers is a sensitive measure of nutritional stress in seabirds. Here, we use this method to examine how changes in diet composition and prey quality affect the nutritional status of free‐living rhinoceros auklets (Cerorhinca monocerata). Our study sites included the following: Teuri Is. Japan, Middleton Is. central Gulf of Alaska, and St. Lazaria Is. Southeast Alaska. In 2012 and 2013, we collected “bill loads” delivered by parents to feed their chicks (n = 758) to document dietary changes. We deployed time–depth–temperature recorders on breeding adults (n = 47) to evaluate whether changes in prey coincided with changes in foraging behavior. We measured concentrations of corticosterone in fledgling (n = 71) and adult breeders' (n = 82) feathers to determine how birds were affected by foraging conditions. We found that seasonal changes in diet composition occurred on each colony, adults dove deeper and engaged in longer foraging bouts when capturing larger prey and that chicks had higher concentrations of corticosterone in their feathers when adults brought back smaller and/or lower energy prey. Corticosterone levels in feathers of fledglings (grown during the breeding season) and those in feathers of adult breeders (grown during the postbreeding season) were positively correlated, indicating possible carryover effects. These results suggest that seabirds might experience increased levels of nutritional stress associated with moderate dietary changes and that physiological responses to changes in prey composition should be considered when evaluating the effect of prey quality on marine predators.     

Varying foraging patterns in response to competition? A multicolony approach in a generalist seabird

Reducing resource competition is a crucial requirement for colonial seabirds to ensure adequate self‐ and chick‐provisioning during breeding season. Spatial segregation is a common avoidance strategy among and within species from neighboring breeding colonies. We determined whether the foraging behaviors of incubating lesser black‐backed gulls (Larus fuscus) differed between six colonies varying in size and distance to mainland, and whether any differences could be related to the foraging habitats visited. Seventy‐nine incubating individuals from six study colonies along the German North Sea coast were equipped with GPS data loggers in multiple years. Dietary information was gained by sampling food pellets, and blood samples were taken for stable isotope analyses. Foraging patterns clearly differed among and within colonies. Foraging range increased with increasing colony size and decreased with increasing colony distance from the mainland, although the latter might be due to the inclusion of the only offshore colony. Gulls from larger colonies with consequently greater density‐dependent competition were more likely to forage at land instead of at sea. The diets of the gulls from the colonies furthest from each other differed, while the diets from the other colonies overlapped with each other. The spatial segregation and dietary similarities suggest that lesser black‐backed gulls foraged at different sites and utilized two main habitat types, although these were similar across foraging areas for all colonies except the single offshore island. The avoidance of intraspecific competition results in colony‐specific foraging patterns, potentially causing more intensive utilization of terrestrial foraging sites, which may offer more predictable and easily available foraging compared with the marine environment.     

Variable sea‐ice conditions influence trophic dynamics in an Arctic community of marine top predators

Sea‐ice coverage is a key abiotic driver of annual environmental conditions in Arctic marine ecosystems and could be a major factor affecting seabird trophic dynamics. Using stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) in eggs of thick‐billed murres (Uria lomvia), northern fulmars (Fulmarus glacialis), glaucous gulls (Larus hyperboreus), and black‐legged kittiwakes (Rissa tridactyla), we investigated the trophic ecology of prebreeding seabirds nesting at Prince Leopold Island, Nunavut, and its relationship with sea‐ice conditions. The seabird community of Prince Leopold Island had a broader isotopic niche during lower sea‐ice conditions, thus having a more divergent diet, while the opposite was observed during years with more extensive sea‐ice conditions. Species' trophic position was influenced by sea ice; in years of lower sea‐ice concentration, gulls and kittiwakes foraged at higher trophic levels while the opposite was observed for murres and fulmars. For murres and fulmars over a longer time series, there was no evidence of the effect of sea‐ice concentration on species' isotopic niche. Results suggest a high degree of adaptation in populations of high Arctic species that cope with harsh and unpredictable conditions. Such different responses of the community isotopic niche also show that the effect of variable sea‐ice conditions, despite being subtle at the species level, might have larger implications when considering the trophic ecology of the larger seabird community. Species‐specific responses in foraging patterns, in particular trophic position in relation to sea ice, are critical to understanding effects of ecosystem change predicted for a changing climate.