Differential adult survival at close seabird colonies: The importance of spatial foraging segregation and bycatch risk during the breeding season

Marine megafauna, including seabirds, are critically affected by fisheries bycatch. However, bycatch risk may differ on temporal and spatial scales due to the uneven distribution and effort of fleets operating different fishing gear, and to focal species distribution and foraging behavior. Scopoli's shearwater Calonectris diomedea is a long‐lived seabird that experiences high bycatch rates in longline fisheries and strong population‐level impacts due to this type of anthropogenic mortality. Analyzing a long‐term dataset on individual monitoring, we compared adult survival (by means of multi‐event capture–recapture models) among three close predator‐free Mediterranean colonies of the species. Unexpectedly for a long‐lived organism, adult survival varied among colonies. We explored potential causes of this differential survival by (1) measuring egg volume as a proxy of food availability and parental condition; (2) building a specific longline bycatch risk map for the species; and (3) assessing the distribution patterns of breeding birds from the three study colonies via GPS tracking. Egg volume was very similar between colonies over time, suggesting that environmental variability related to habitat foraging suitability was not the main cause of differential survival. On the other hand, differences in foraging movements among individuals from the three colonies expose them to differential mortality risk, which likely influenced the observed differences in adult survival. The overlap of information obtained by the generation of specific bycatch risk maps, the quantification of population demographic parameters, and the foraging spatial analysis should inform managers about differential sensitivity to the anthropogenic impact at mesoscale level and guide decisions depending on the spatial configuration of local populations. The approach would apply and should be considered in any species where foraging distribution is colony‐specific and mortality risk varies spatially.     

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.     

Predictable anthropogenic food subsidies,density‐dependence and socio‐economic factors influence breeding investment in a generalist seabird

Recent European policies on the ban of fishing discards and the closure of open‐air landfills are expected to reduce predictable and abundant food resources for generalist seabirds. In order to forecast the consequences of this reduction on seabird breeding investment it is important to understand whether diverse anthropogenic foraging resources act synergistically or not and whether their influence is mediated by density‐dependent mechanisms. To assess these effects at large spatio‐temporal scale, we measured mean egg volume as a proxy of breeding investment in ca 5000 three‐egg clutches of the yellow‐legged gull Larus michahellis from 20 colonies of the Western Mediterranean, located both along European and African coasts. In European gull colonies, egg volume increased with the availability of fishing discards and landfills in the vicinity of the colony. However, the landfill effect was weaker than the effect of fishing discards, probably due to the lower quality of waste as food for gulls. In contrast, none of the anthropogenic food subsidies influenced egg volume in African colonies, likely due to socio‐economic differences (i.e. a much lower availability and predictability of both discards and waste food. Finally, results showed that the positive association between fishing discards and open‐air landfills on egg volume was mediated by negative density‐dependent mechanisms probably related to an increase in competition for food.     

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.     

Spatial segregation between immatures and adults in a pelagic seabird suggests age‐related competition

Individual competitiveness conditions access to resources when they are limited. Immature individuals that are less skilled than adults have to adapt their foraging strategies to survive. Among strategies to reduce competition, spatial segregation has been widely demonstrated. However, the use of spatial segregation by immatures to limit intra‐specific competition with adults has rarely been tested. In this study, we investigated and compared habitat preferences and distributions of free‐ranging immature and breeding adult northern gannets Morus bassanus in order to determine whether they compete for similar habitats during the year, and if this results in a spatial segregation between birds of different age groups. Based on > 66 000 km of aerial surveys conducted in the North‐East Atlantic Ocean during winter and summer 2012, habitats selected by immatures and adult birds were modelled independently, linking gannet density to a set of oceanographic and physiographic predictors. Their large‐scale seasonal distribution was then predicted. We found that gannets displayed a strong season‐dependent competition between immatures and adults, as a consequence of immatures and adults using similar habitats in both summer and winter. During summer, when adults are constrained by reproduction, both groups were spatially highly segregated despite similar habitat preferences (thermal fronts), with youngest individuals selecting habitats out of range of central‐place foragers, highlighting intra‐specific competition. Contrastingly during winter, when reproductive constraints disappear, immature and adult distributions largely overlapped. Our study provides new insights into the role played by age, foraging experience and reproductive constraints on the distribution of marine predators. More specifically, these results highlight in seabirds how the youngest fraction mitigates, through spatial segregation, the competition with experienced adults, and suggest a progressive strategy along the maturation process.     

Living on predictability: modelling the density distribution of efficient foraging seabirds

In areas with regular fishing coastal fleets seabirds may benefit from the predictability of discards from fishing vessels, but it is not clear to what extent birds rely on this predictable resource and whether foraging is synchronized with the diel availability of discards. In this paper we investigate if a typical scavenger species, the yellow‐legged gull Larus michahellis, takes advantage of the temporal and spatial predictability of fish discards in the western Mediterranean Sea. The activity and distribution of the trawling fleet in this area is regulated and very predictable in time and space. We gathered aerial survey data across a relatively large area close to the coast to study the spatial distribution and density of L. michahellis, and modelled the density distribution of the species in relation to several oceanographic, ecological and temporal variables, using two different modelling approaches: MARS (multivariate adaptative regression splines) and GLM (generalized linear models). Our models suggest that the spatial density of trawlers at sea and the time of the day are the best explanatory variables of gull distribution, and that gulls concentrate in areas with vessels mainly during fish discarding time, supporting the hypothesis that gulls optimize time foraging to take advantage of fishery waste predictability. Additional surveys from the main gull roosting sites inshore support this hypothesis, as gulls start leaving to the sea just before fishing is completed and vessels begin discarding fish scraps when back to the harbour. This study represents one of the few examples of applying MARS to density distribution modelling, although its application to marine ecosystems should be conducted with caution because of large areas with real absence data. GLMs have shown to be more adaptable to such kind of data. Our data confirm the importance of fishery waste for L. michahellis, not only as a food resource but also as a major driver of their activity and distribution patterns. The ability of seabirds to predict accurately when a food resource will be available implies that modelling their distribution at sea needs to include such variables, both in spatial and temporal dimensions.     

The depth of Sooty Shearwater Ardenna grisea burrows varies with habitat and increases with competition for space

The Sooty Shearwater Ardenna grisea, an abundant but declining petrel, is one of many seabird species that construct breeding burrows, presumably because these confer protection from predators and the elements. Little is known about the causes of variation in Sooty Shearwater burrow architecture, which can differ markedly both within and between breeding sites. We hypothesize that burrow architecture varies in response to habitat type and competition for space. To address these hypotheses, we recorded Sooty Shearwater burrow dimensions on Kidney Island, the largest Sooty Shearwater colony in the Falkland Islands, South Atlantic, and modelled these as functions of burrow density (a proxy for competition) and habitat indices. Our models suggest that Sooty Shearwaters burrow further underground in response to competition for breeding space, and that soil underlying dense tussac grass Poa flabellata is more easily excavated than other substrates, indicating how vegetation restoration could aid the conservation of this species.     

Sex‐specific foraging behavior in response to fishing activities in a threatened seabird

Some seabird species have learnt to efficiently exploit fishing discards from trawling activities. However, a discard ban has been proposed as necessary in Europe to ensure the sustainability of the seas. It is of crucial importance for the management and conservation purposes to study the potential consequences of a discard ban on the foraging ecology of threatened seabirds. We assessed the influence of fishing activities on the feeding habits of 22 male and 15 female Audouin's gulls (Larus audouinii) from the Ebro Delta (Mediterranean Sea) during the breeding period using GPS loggers together with Stable Isotope Analysis (SIA), which provided new insights into their foraging behavior and trophic ecology, respectively. GPS data revealed different sex‐specific foraging patterns between workdays and weekends. Females were highly consistent in that they foraged at sea throughout the week even though discarding stops at weekends. In contrast, males switched from foraging at sea during the week (when discards are produced) to an increased use of rice field habitats at weekends (when fishermen do not work). This sex‐specific foraging behavior could be related to specific nutritional requirements associated with previous egg production, an energetically demanding period for females. However, on a broader time scale integrated by the SIA, both sexes showed a high degree of individual specialization in their trophic ecology. The need to obtain detailed information on the dependence and response of seabirds to fishing activities is crucial in conservation sciences. In this regard, sex‐specific foraging behavior in relation to fisheries has been overlooked, despite the ecological and conservation implications. For instance, this situation may lead to sex differentiation in bycatch mortality in longlines when trawlers do not operate. Moreover, any new fisheries policy will need to be implemented gradually to facilitate the adaptation of a specialized species to a discard ban scenario.     

Sensory‐based conservation of seabirds: a review of management strategies and animal behaviours that facilitate success

Sensory‐based conservation harnesses species' natural communication and signalling behaviours to mitigate threats to wild populations. To evaluate this emerging field, we assess how sensory‐based manipulations, sensory mode, and target taxa affect success. To facilitate broader, cross‐species application of successful techniques, we test which behavioural and life‐history traits correlate with positive conservation outcomes. We focus on seabirds, one of the world's most rapidly declining groups, whose philopatry, activity patterns, foraging, mate choice, and parental care behaviours all involve reliance on, and therefore strong selection for, sophisticated sensory physiology and accurate assessment of intra‐ and inter‐species signals and cues in several sensory modes. We review the use of auditory, olfactory, and visual methods, especially for attracting seabirds to newly restored habitat or deterring birds from fishing boats and equipment. We found that more sensory‐based conservation has been attempted with Procellariiformes (tube‐nosed seabirds) and Charadriiformes (e.g. terns and gulls) than other orders, and that successful outcomes are more likely for Procellariiformes. Evolutionary and behavioural traits are likely to facilitate sensory‐based techniques, such as social attraction to suitable habitat, across seabird species. More broadly, successful application of sensory‐based conservation to other at‐risk animal groups is likely to be associated with these behavioural and life‐history traits: coloniality, philopatry, nocturnal, migratory, long‐distance foraging, parental care, and pair bonds/monogamy.     

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.     

Varying demographic impacts of different fisheries on three Mediterranean seabird species

Fisheries have an enormous economic importance, but reconciling their socio‐economic features with the conservation and sustainability of marine ecosystems presents major challenges. Bycatch mortality from fisheries is clearly among the most serious global threats for marine ecosystems, affecting a wide range of top predators. Recent estimates report ca. 200,000 seabirds killed annually by bycatch in European waters. However, there is an urgent need to rigorously estimate actual mortality rates and quantify effects of bycatch on populations. The Mediterranean Sea is one of the most impacted regions. Here, we estimate for the first time both bycatch mortality rates and their population‐level effects on three endemic and vulnerable Mediterranean taxa: Scopoli's shearwater, Mediterranean shag, and Audouin's gull, that die in different types of fishing gears: longlines, gillnets and sport trolling, respectively. We use multi‐event capture–recapture modelling to estimate crucial demographic parameters, including the probabilities of dying in different fishing gears. We then build stochastic demography models to forecast the viability of the populations under different management scenarios. Longline bycatch was particularly severe for adults of Scopoli's shearwaters and Audouin's gulls (ca. 28% and 23% of total mortality, respectively) and also for immature gulls (ca. 90% of mortality). Gillnets had a lower impact, but were still responsible for ca. 9% of juvenile mortality on shags, whereas sport trolling only slightly influenced total mortality in gulls. Bycatch mortality has high population‐level impacts in all three species, with shearwaters having the highest extinction risk under current mortality rates. Different life‐history traits and compensatory demographic mechanisms between the three species are probably influencing the different bycatch impact: for shearwaters, urgent conservation actions are required to ensure the viability of their populations. Results will be very useful for guiding future seabird conservation policies and moving towards an ecosystem‐based approach to sustainable fisheries management.     

Movement patterns of large juvenile loggerhead turtles in the Mediterranean Sea: Ontogenetic space use in a small ocean basin

Mechanisms that determine how, where, and when ontogenetic habitat shifts occur are mostly unknown in wild populations. Differences in size and environmental characteristics of ontogenetic habitats can lead to differences in movement patterns, behavior, habitat use, and spatial distributions across individuals of the same species. Knowledge of juvenile loggerhead turtles' dispersal, movements, and habitat use is largely unknown, especially in the Mediterranean Sea. Satellite relay data loggers were used to monitor movements, diving behavior, and water temperature of eleven large juvenile loggerhead turtles (Caretta caretta) deliberately caught in an oceanic habitat in the Mediterranean Sea. Hidden Markov models were used over 4,430 spatial locations to quantify the different activities performed by each individual: transit, low‐, and high‐intensity diving. Model results were then analyzed in relation to water temperature, bathymetry, and distance to the coast. The hidden Markov model differentiated between bouts of area‐restricted search as low‐ and high‐intensity diving, and transit movements. The turtles foraged in deep oceanic waters within 60 km from the coast as well as above 140 km from the coast. They used an average area of 194,802 km², where most individuals used the deepest part of the Southern Tyrrhenian Sea with the highest seamounts, while only two switched to neritic foraging showing plasticity in foraging strategies among turtles of similar age classes. The foraging distribution of large juvenile loggerhead turtles, including some which were of the minimum size of adults, in the Tyrrhenian Sea is mainly concentrated in a relatively small oceanic area with predictable mesoscale oceanographic features, despite the proximity of suitable neritic foraging habitats. Our study highlights the importance of collecting high‐resolution data about species distribution and behavior across different spatio‐temporal scales and life stages for implementing conservation and dynamic ocean management actions.     

At‐sea distribution and habitat use in king penguins at sub‐Antarctic Marion Island

King penguins make up the bulk of avian biomass on a number of sub‐Antarctic islands where they have a large functional effect on terrestrial and marine ecosystems. The same applies at Marion Island where a substantial proportion of the world population breeds. In spite of their obvious ecological importance, the at‐sea distribution and behavior of this population has until recently remained entirely unknown. In addressing this information deficiency, we deployed satellite‐linked tracking instruments on 15 adult king penguins over 2 years, April 2008 and 2013, to study their post‐guard foraging distribution and habitat preferences. Uniquely among adult king penguins, individuals by and large headed out against the prevailing Antarctic Circumpolar Current, foraging to the west and southwest of the island. On average, individuals ventured a maximum distance of 1,600 km from the colony, with three individuals foraging close to, or beyond, 3,500 km west of the colony. Birds were mostly foraging south of the Antarctic Polar Front and north of the southern boundary of the Antarctic Circumpolar Current. Habitat preferences were assessed using boosted regression tree models which indicated sea surface temperate, depth, and chorophyll a concentration to be the most important predictors of habitat selection. Interestingly, king penguins rapidly transited the eddy‐rich area to the west of Marion Island, associated with the Southwest Indian Ocean Ridge, which has been shown to be important for foraging in other marine top predators. In accordance with this, the king penguins generally avoided areas with high eddy kinetic energy. The results from this first study into the behavioral ecology and at‐sea distribution of king penguins at Marion Island contribute to our broader understanding of this species.     

Short‐term prey field lability constrains individual specialisation in resource selection and foraging site fidelity in a marine predator

Spatio‐temporally stable prey distributions coupled with individual foraging site fidelity are predicted to favour individual resource specialisation. Conversely, predators coping with dynamic prey distributions should diversify their individual diet and/or shift foraging areas to increase net intake. We studied individual specialisation in Scopoli's shearwaters (Calonectris diomedea) from the highly dynamic Western Mediterranean, using daily prey distributions together with resource selection, site fidelity and trophic‐level analyses. As hypothesised, we found dietary diversification, low foraging site fidelity and almost no individual specialisation in resource selection. Crucially, shearwaters switched daily foraging tactics, selecting areas with contrasting prey of varying trophic levels. Overall, information use and plastic resource selection of individuals with reduced short‐term foraging site fidelity allow predators to overcome prey field lability. Our study is an essential step towards a better understanding of individual responses to enhanced environmental stochasticity driven by global changes, and of pathways favouring population persistence.     

Fine‐scale spatial distribution of murundus structures in the semi‐arid region of Brazil

Murundus are earth mounds widespread in most landscapes in the semi‐arid region of Brazil. Evidence obtained from predictive modelling has suggested a termite origin for these structures, opening up new opportunities for further research. Distribution of densely packed murundus at larger spatial scales is most related to climatic regime and soil nutrient availability. However, factors and processes underlying their distribution and density at smaller spatial scales are not yet fully understood. In this study, we adopted an approach based on mapping point data using high‐resolution satellite imagery, multi‐scale second‐order analysis and general linear models to examine the fine‐scale spatial distribution and density of murundus. Our results suggest that the distribution of those structures within densely packed areas is regulated by more than one process acting or interacting across multiple spatial scales. All densely packed murundus showed a significant regular distribution at the distance scale of up to 50 m radially and a completely random distribution across all other upper distance scales. We interpret the regular pattern as a result of competition for foraging territories between different termite colonies during the process formation of densely packed murundus. The random pattern at larger distance scales (above 50 m radially) can be attributed to habitat selection preferences by termite species builders of murundus mediated by local environmental resources and conditions (i.e. availability of food resources and nesting and open habitat), which would be randomly distributed in space. Thus, at finer spatial scales murundus distributions are associated with biotic interactions acting on an abiotic template. On the basis of significant linear correlations, we suggest that the density of murundus is strongly related to local temperature regime with soil‐type influencing its effect on the murundus densities. Our findings provide novel evidences that mound‐building termites are involved in the formation of murundus in the semi‐arid region of Brazil.     

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.     

Ontogenetic variation in diet and habitat of Irrawaddy dolphins (Orcaella brevirostris) in the Gulf of Thailand and the Andaman Sea

Irrawaddy dolphins (Orcaella brevirostris) are threatened, little‐known cetaceans primarily jeopardized by fishing gear entanglement. Information regarding life history, foraging ecology, and movement are crucial to determine management units and conservation regions. This is the first analysis of the ontogenetic changes in diet and habitat of Irrawaddy dolphins from the Gulf of Thailand and the Andaman Sea using stable isotopes from skin and teeth. Results suggest limited age and sex structure indicated by weak ontogenetic and sexual variability in δ¹⁵N and δ¹³C in teeth. Significant variation in δ¹³C and δ¹⁵N values were observed between three Gulf regions and the Andaman Sea, indicating distinct regional groups. Isotope mixing models run on soft tissues from the eastern Gulf revealed a diet based primarily on fishes and secondarily on crustaceans, suggesting overlap with fishery targets. Conservation strategies that focus on reducing regional competition with humans and bycatch mortality are proposed. Fishing gear improvements and monitoring of fishing operations are a potential immediate response to reduce anthropogenic impact.     

Women from Venus,men from Mars: inter‐sex foraging differences in the imperial cormorant Phalacrocorax atriceps a colonial seabird

Colonial seabirds are central place foragers and likely to be subject to substantial competition for resources. Mechanisms proposed for reducing intra‐specific competition include differential inter‐sex area use mediated by adult choice. We used GPS loggers and dive recorders to study area use and dive depth in a total of 27 male and 26 female imperial cormorants Phalacrocorax atriceps breeding at a colony of some 6500 birds at Punta Leon, Chubut, Argentina during 2004 and 2005. Although time spent travelling and distances between the colony and foraging sites were similar for both sexes, males and females travelled away from their colony using routes virtually perpendicular to each other so that their foraging areas were distinctly different; females hunted close to the coast while males foraged offshore in deeper water. Consideration of foraging efficiency underwater, defined as the duration spent on the bottom divided by the dive cycle duration, showed that females were more efficient at depths < 40 m while males more efficient at depths > 40 m. We suggest that the substantial sexual dimorphism in this species may be responsible for the different depth‐linked foraging efficiencies and that selection for appropriate depths could lead to differential habitat use and putative differences in prey selection.     

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.     

Bio‐energetics underpins the spatial response of North Sea plaice (Pleuronectes platessa L.) and sole (Solea solea L.) to climate change

Climate change is currently one of the main driving forces behind changes in species distributions, and understanding the mechanisms that underpin macroecological patterns is necessary for a more predictive science. Warming sea water temperatures are expected to drive changes in ectothermic marine species ranges due to their thermal tolerance levels. Here, we develop a mechanistic tool to predict size‐ and season‐specific distributions based on the physiology of the species and the temperature and food conditions in the sea. The effects of climate conditions on physiological‐based habitat utilization was then examined for different size‐classes of two commercially important fish species in the North Sea, plaice, Pleuronectes platessa, and sole, Solea solea. The two species provide an attractive comparison as they differ in their physiology (e.g. preferred temperature range). Combining dynamic energy budget (DEB) models with the temperature and food conditions estimated by an ecosystem model (ERSEM), allowed spatial differences in potential growth (as a proxy for habitat quality) to be estimated for 2 years with contrasting temperature and food conditions. The resulting habitat quality maps were in broad agreement with observed ontogenetic and seasonal changes in distribution as well as with the recent changes in distribution which could be attributed to an increase in coastal temperatures. Our physiological‐based model provides a powerful tool to explore the effect of climate change on the spatio‐temporal fish dynamics, predict effects of local or broad‐scale environmental changes and provide a physiological basis for observed changes in species distributions.