Boldness predicts foraging behaviour, habitat use and chick growth in a central place marine predator.

Animal personality can shape individual's fitness. Yet, the mechanistic relationship by which individual's personality traits lead to variations in fitness remains largely underexplored. Here, we used novel object tests to measure boldness of chick-provisioning Cory's shearwaters (Calonectris borealis) from a coastal colony off west Portugal, and deployed GPS loggers to study their at-sea behaviour and distribution. We then tested whether boldness predicts individual differences in adult's trophic ecology and variations in chick growth, to assess potential implications of personality-specific foraging behaviours. Foraging effort was higher for shyer than for bolder individuals, which, during short forays, exhibited larger foraging ranges, and foraged in regions of higher and more variable bathymetry. This suggests that nearby the colony bolder individuals expanded their foraging area to maximize resource acquisition and increase the probability of foraging success. When endeavouring to longer distances, bolder individuals exhibited comparably shorter foraging ranges and targeted low bathymetry regions, likely with enhanced prey availability, while shyer individuals exhibited much larger foraging ranges indicating greater flexibility when foraging in oceanic realms. Despite such differences between bolder and shyer individuals their isotopic niches were similar. Yet, chicks raised by bolder parents grew at a faster rate than those raised by shyer parents. Together, our results suggest that differences in resource acquisition strategies could play a key role through which individual's boldness may influence breeding performance, even when individuals have similar isotopic preferences.

Global assessment of marine plastic exposure risk for oceanic birds.

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

Assessment of contaminants in blue sharks from the Northeast Atlantic: Profiles, accumulation dynamics, and risks for human consumers.

Chemical pollution is a major threat to marine ecosystems, and top predators such as most shark species are extremely vulnerable to being exposed and accumulating contaminants such as metals and persistent organic pollutants (POPs). This work aimed to study the degree, composition, and the sources of contamination in the blue shark (Prionace glauca) inhabiting the Northeast Atlantic, as well as the potential risk faced by human consumers. A total of 60 sharks were sampled in situ aboard fishing vessels, and the concentrations of a set of metals and POPs were analysed in various tissues and complemented with stable isotope analyses. High levels of contaminants were found in most sharks sampled. The concentrations of most metals were higher in the muscle when compared with the liver. Regarding the dangers to consumers posed by the concentrations of arsenic (As), mercury (Hg), and lead (Pb), over 75% of the sharks presented muscle concentrations of at least one contaminant above the legal limits for human consumption, and a risk assessment determined that consumption of meat of these sharks exceeding 0.07 Kg per week could potentially expose human consumers to dangerous amounts of methylmercury (MeHg). Additionally, the assessment of single contaminants may lead to an underestimation of the risk for the human health. Finally, the overall accumulation of contaminants seems to be mostly influenced by the sharks' geographical distribution, rather than sex, size, or trophic level of their prey.

Life History of the Arctic Squid Gonatus fabricii (Cephalopoda: Oegopsida) Reconstructed by Analysis of Individual Ontogenetic Stable Isotopic Trajectories.

Cephalopods are important in Arctic marine ecosystems as predators and prey, but knowledge of their life cycles is poor. Consequently, they are under-represented in the Arctic ecosystems assessment models. One important parameter is the change in ecological role (habitat and diet) associated with individual ontogenies. Here, the life history of Gonatus fabricii, the most abundant Arctic cephalopod, is reconstructed by the analysis of individual ontogenetic trajectories of stable isotopes (δ13C and δ15N) in archival hard body structures. This approach allows the prediction of the exact mantle length (ML) and mass when the species changes its ecological role. Our results show that the life history of G. fabricii is divided into four stages, each having a distinct ecology: (1) epipelagic squid (ML < 20 mm), preying mostly on copepods; (2) epi- and occasionally mesopelagic squid (ML 20−50 mm), preying on larger crustaceans, fish, and cephalopods; (3) meso- and bathypelagic squid (ML > 50 mm), preying mainly on fish and cephalopods; and (4) non-feeding bathypelagic gelatinous females (ML > 200 mm). Existing Arctic ecosystem models do not reflect the different ecological roles of G. fabricii correctly, and the novel data provided here are a necessary baseline for Arctic ecosystem modelling and forecasting.

Mercury biomagnification in an Antarctic food web of the Antarctic Peninsula.

Under the climate change context, warming Southern Ocean waters may allow mercury (Hg) to become more bioavailable to the Antarctic marine food web (i.e., ice-stored Hg release and higher methylation rates by microorganisms), whose biomagnification processes are poorly documented. Biomagnification of Hg in the food web of the Antarctic Peninsula, one of the world's fastest-warming regions, was examined using carbon (δ13C) and nitrogen (δ15N) stable isotope ratios for estimating feeding habitat and trophic levels, respectively. The stable isotope signatures and total Hg (T-Hg) concentrations were measured in Antarctic krill Euphausia superba and several Antarctic predator species, including seabirds (gentoo penguins Pygoscelis papua, chinstrap penguins Pygoscelis antarcticus, brown skuas Stercorarius antarcticus, kelp gulls Larus dominicanus, southern giant petrels Macronectes giganteus) and marine mammals (southern elephant seals Mirounga leonina). Significant differences in δ13C values among species were noted with a great overlap between seabird species and M. leonina. As expected, significant differences in δ15N values among species were found due to interspecific variations in diet-related to their trophic position within the marine food web. The lowest Hg concentrations were registered in E. superba (0.007 ± 0.008 µg g-1) and the highest values in M. giganteus (12.090 ± 14.177 µg g-1). Additionally, a significant positive relationship was found between Hg concentrations and trophic levels (reflected by δ15N values), biomagnifying nearly 2 times its concentrations at each level. Our results support that trophic interaction is the major pathway for Hg biomagnification in Southern Ocean ecosystems and warn about an increase in the effects of Hg on long-lived (and high trophic level) Antarctic predators under climate change in the future.

Foraging costs drive within-colony spatial segregation in shearwaters from two contrasting environments in the North Atlantic Ocean.

Foraging spatial segregation is frequent in central-place foragers during the breeding season, but very few studies have investigated foraging spatial segregation between adjacent sub-colonies. Here, we assessed for within-colony differences in the at-sea distribution, habitat use, trophic ecology and chick growth data of two Calonectris colonies differing in size, and breeding in two different environments in the North Atlantic Ocean. For this, we GPS tracked 52 Cory's shearwaters (Calonectris borealis) breeding in 2 small sub-colonies at Berlenga Island (Portugal) and 59 Cape Verde shearwaters (Calonectris edwardsii) breeding in 2 sub-colonies differing greatly in size at Raso Islet (Cabo Verde), over 2 consecutive breeding seasons (2017-2018), during chick-rearing. Cory's shearwaters from the two sub-colonies at Berlenga Island broadly overlapped in repeatedly used foraging patches close to the colony. In contrast, the foraging distribution of Cape Verde shearwaters was partially segregated in the colony surroundings, but overlapped at distant foraging areas off the west coast of Africa. Despite spatial segregation close to the colony, Cape Verde shearwaters from both sub-colonies departed in similar directions, foraged in similar habitats and exhibited mostly short trips within the archipelago of Cabo Verde. These results, corroborated with similar trophic ecology and chick growth rates between sub-colonies, support the idea that foraging spatial segregation in the colony surroundings was not likely driven by interference competition or directional bias. We suggest that high-quality prey patches are able to shape travel costs and foraging distribution of central-place foragers from neighbouring sub-colonies.

Accumulation of chemical elements and occurrence of microplastics in small pelagic fish from a neritic environment.

The assessment of contaminant exposure in marine organisms often focuses on the most toxic chemical elements from upper trophic level species. Information on mid-trophic level species and particularly on potentially less harmful elements is lacking. Additionally, microplastics have been considered emergent contaminants in aquatic environments which have not been extensively studied in species from mid-trophic levels in food chains. This study aims to contribute to an overall assessment of environmental impacts of such chemicals in a community of small pelagic fish in the North Atlantic. The concentrations of 16 chemical elements, rarely simultaneously quantified (including minerals, trace elements and heavy metals), and the presence of microplastics were analysed in sardines (Sardina pilchardus) and mackerels (Scomber spp. and Trachurus trachurus) sampled along the Portuguese coast. Biochemical stress assessments and stable isotope analyses were also performed. The chemical element concentrations in S. pilchardus, T. trachurus, and Scomber spp. were relatively low and lower than the levels reported for the same species in the North Atlantic and adjacent areas. No clear relationships were found between chemical elements and oxidative damage in fish. However, the concentration of several chemical elements showed differences among species, being related with the species' habitat use, trophic niches, and specific feeding strategies. The presence of plastic pieces in the stomachs of 29% of the sampled fishes is particularly concerning, as these small pelagic fish from mid-trophic levels compose a significant part of the diet of humans and other top predators. This study highlights the importance of multidisciplinary approaches focusing on the individual, including position data, stable isotopes, and oxidative stress biomarkers as complementary tools in contamination assessment of the marine mid-trophic levels in food chains.

Untangling local and remote influences in two major petrel habitats in the oligotrophic Southern Ocean.

Ocean circulation connects geographically distinct ecosystems across a wide range of spatial and temporal scales via exchanges of physical and biogeochemical properties. Remote oceanographic processes can be especially important for ecosystems in the Southern Ocean, where the Antarctic Circumpolar Current transports properties across ocean basins through both advection and mixing. Recent tracking studies have indicated the existence of two large-scale, open ocean habitats in the Southern Ocean used by grey petrels (Procellaria cinerea) from two populations (i.e., Kerguelen and Antipodes islands) during their nonbreeding season for extended periods during austral summer (i.e., October to February). In this work, we use a novel combination of large-scale oceanographic observations, surface drifter data, satellite-derived primary productivity, numerical adjoint sensitivity experiments, and output from a biogeochemical state estimate to examine local and remote influences on these grey petrel habitats. Our aim is to understand the oceanographic features that control these isolated foraging areas and to evaluate their ecological value as oligotrophic open ocean habitats. We estimate the minimum local primary productivity required to support these populations to be much <1% of the estimated local primary productivity. The region in the southeast Indian Ocean used by the birds from Kerguelen is connected by circulation to the productive Kerguelen shelf. In contrast, the region in the south-central Pacific Ocean used by seabirds from the Antipodes is relatively isolated suggesting it is more influenced by local factors or the cumulative effects of many seasonal cycles. This work exemplifies the potential use of predator distributions and oceanographic data to highlight areas of the open ocean that may be more dynamic and productive than previously thought. Our results highlight the need to consider advective connections between ecosystems in the Southern Ocean and to re-evaluate the ecological relevance of oligotrophic Southern Ocean regions from a conservation perspective.

Year-round element quantification of a wide-ranging seabird and their relationships with oxidative stress, trophic ecology, and foraging patterns.

Multidisciplinary approaches are essential to diligently assess environmental health status of ecosystems. In this study, year-round chemical elements' exposure and impacts were assessed on the wide-ranging Cory's shearwater Calonectris borealis breeding in Berlenga Island, offshore Portugal, North Atlantic Ocean. The aim was to identify potential contamination and oxidative stress sources associated with trophic ecology, habitat and spatial use, and foraging patterns. A set of 20 chemical elements were quantified, along with oxidative stress biomarkers, stable isotope analyses, and GPS tracking data. Birds presented higher accumulation to some non-essential elements along the year (i.e. arsenic, As; cadmium, Cd; mercury, Hg; lead, Pb; and strontium, Sr), in which concentrations were similar or surpassed other procellariform seabird populations all over the world. No significant differences were found for any of the elements between different periods within the breeding season, with exception of Hg. However, a Principal Component Analysis taking into consideration a group of elements showed differences between pre-laying and chick-rearing periods, with overall higher concentrations in the former. Individuals spending more time engaging in an intensive search for food, and in more coastal environments, presented overall higher element concentrations, and particularly Hg. Contrary to expectations, no relationships were found between chemical elements and oxidative stress. On the other hand, spatial use and foraging patterns of Cory's shearwaters influenced their oxidative stress responses. Our results highlight the need for multidisciplinary approaches to deepen understanding of the large-scale vulnerability of bioindicators such as seabirds and, by extension, the overall environmental health of ecosystems in which they rely.

Mercury biomagnification in a Southern Ocean food web.

Biomagnification of mercury (Hg) in the Scotia Sea food web of the Southern Ocean was examined using the stable isotope ratios of nitrogen (δ15N) and carbon (δ13C) as proxies for trophic level and feeding habitat, respectively. Total Hg and stable isotopes were measured in samples of particulate organic matter (POM), zooplankton, squid, myctophid fish, notothenioid fish and seabird tissues collected in two years (austral summers 2007/08 and 2016/17). Overall, there was extensive overlap in δ13C values across taxonomic groups suggesting similarities in habitats, with the exception of the seabirds, which showed some differences, possibly due to the type of tissue analysed (feathers instead of muscle). δ15N showed increasing enrichment across groups in the order POM to zooplankton to squid to myctophid fish to notothenioid fish to seabirds. There were significant differences in δ15N and δ13C values among species within taxonomic groups, reflecting inter-specific variation in diet. Hg concentrations increased with trophic level, with the lowest values in POM (0.0005 ± 0.0002 µg g-1 dw) and highest values in seabirds (3.88 ± 2.41 µg g-1 in chicks of brown skuas Stercorarius antarcticus). Hg concentrations tended to be lower in 2016/17 than in 2007/08 for mid-trophic level species (squid and fish), but the opposite was found for top predators (i.e. seabirds), which had higher levels in the 2016/17 samples. This may reflect an interannual shift in the Scotia Sea marine food web, caused by the reduced availability of a key prey species, Antarctic krill Euphausia superba. In 2016/17, seabirds would have been forced to feed on higher trophic-level prey, such as myctophids, that have higher Hg burdens. These results suggest that changes in the food web are likely to affect the pathway of mercury to Southern Ocean top predators.

Diet and life history reduce interspecific and intraspecific competition among three sympatric Arctic cephalopods.

Trophic niche and diet comparisons among closely sympatric marine species are important to understand complex food webs, particularly in regions most affected by climate change. Using stable isotope analyses, all ontogenetic stages of three sympatric species of Arctic cephalopods (genus Rossia) were studied to assess inter- and intraspecific competition with niche and diet overlap and partitioning in West Greenland and the Barents Sea. Seven traits related to resource and habitat utilization were identified in Rossia: no trait was shared by all three species. High boreal R. megaptera and Arctic endemic R. moelleri shared three traits with each other, while both R. megaptera and R. moelleri shared only two unique traits each with widespread boreal-Arctic R. palpebrosa. Thus all traits formed fully uncrossing pattern with each species having unique strategy of resource and habitat utilization. Predicted climate changes in the Arctic would have an impact on competition among Rossia with one potential 'winner' (R. megaptera in the Barents Sea) but no potential 'losers'.

Inter-annual changes in oceanic conditions drives spatial and trophic consistency of a tropical marine predator.

Pelagic seabirds exhibit plasticity in foraging characteristics in relation to oceanographic conditions. This should be particularly relevant in tropical marine environments where food resources are naturally more unpredictable. We studied how inter-annual variations (2013-2018) in tropical oceanographic conditions (driver of oceanic productivity) can influence the spatial and trophic ecology of Cape Verde shearwater (Calonectris edwardsii) during the breeding season. During years of poor oceanographic conditions around the colony, birds engaged in longer trips to West Africa, showed higher spatial and behavioural consistency, and presented a wider isotopic niche. Opposite patterns were generally found for years of good oceanographic conditions, when birds foraged more on their colony surroundings. New foraging areas off West Africa were highlighted as relevant, especially during years of poor environmental conditions. This study highlights the need for long-term studies to assess variation in foraging areas and foraging decisions by seabird populations.

Long-term changes in habitat and trophic level of Southern Ocean squid in relation to environmental conditions.

Long-term studies of pelagic nekton in the Southern Ocean and their responses to ongoing environmental change are rare. Using stable isotope ratios measured in squid beaks recovered from diet samples of wandering albatrosses Diomedea exulans, we assessed decadal variation (from 1976 to 2016) in the habitat (δ13C) and trophic level (δ15N) of five important Southern Ocean squid species in relation to indices of environmental conditions-Southern Oscillation Index (SOI) and Southern Annular Mode (SAM). Based on δ13C values, corrected for the Suess effect, habitat had changed over the last 50 years for Taonius sp. B (Voss), Gonatus antarcticus, Galiteuthis glacialis and Histioteuthis atlantica but not Moroteuthopsis longimana. By comparison, mean δ15N values were similar across decades for all five species, suggesting minimal changes in trophic levels. Both SAM and SOI have increased in strength and frequency over the study period but, of the five species, only in Taonius sp. B (Voss) did these indices correlate with, δ13C and δ15N values, indicating direct relationships between environmental conditions, habitat and trophic level. The five cephalopod species therefore changed their habitats with changing environmental conditions over the last 50 years but maintained similar trophic levels. Hence, cephalopods are likely to remain important prey for top predators in Southern Ocean food webs, despite ongoing climate change.

Assessment of environmental health based on a complementary approach using metal quantification, oxidative stress and trophic ecology of two gull species (Larus michahellis & Larus audouinii) breeding in sympatry.

Metal pollution is currently a major issue in marine ecosystems, as organisms, and particularly seabirds, are exposed and accumulating increased levels from several anthropogenic sources. A set of 13 metals were quantified in two gull species breeding in sympatry, and in two distinct colonies separated by ca. 400 km. Oxidative stress was measured, and stable isotope analyses were used to link metal contamination and oxidative stress with the trophic ecology of each species/population. There was a clear segregation of metal contamination between the two species and to a much lesser extent between colonies. Overall, Audouin's gull was the most contaminated species for most metals, once this species relies mainly on fish and other marine resources. The Yellow-legged gull feeds regularly on terrestrial food sources besides fish, which may dilute contamination levels. Oxidative stress responses were related with birds' trophic ecology and foraging habitat, but apparently not with metal contamination.

Oceans of stimuli: an individual-based model to assess the role of olfactory cues and local enhancement in seabirds' foraging behaviour.

Oceans are extremely dynamic environments, which poses challenges for top-predators such as seabirds to find food resources. Yet, seabirds evolved sensorial abilities (olfactory senses) along with complex behaviours (social information transfer through local enhancement) to improve foraging efficiency. Using the Cory's shearwater (Calonectris borealis) as a model species, we developed an individual-based model to explore the complementary role of different searching mechanisms (olfactory foraging and local enhancement) for the optimal foraging behaviour of pelagic seabirds during 1-day foraging trips around breeding colonies. Model outputs were compared with observed patterns of Cory's shearwaters distribution during local foraging trips. Also, the foraging efficiency of virtual individuals was analysed considering hypothetical scenarios of foraging conditions and densities of foraging individuals around breeding colonies. The results support the use of a combination of searching strategies by Cory's shearwaters, which produced representative patterns of space use from tracked individuals, including spatial foraging segregation of neighbouring sub-colonies. Furthermore, while the mechanisms underpinning local enhancement played a key role in mitigating sub-optimal foraging conditions, the use of olfactory senses conferred great adaptive foraging advantages over a wide range of environmental conditions. Our results also indicate a synergistic effect between the two strategies, which suggests that a multimodal foraging strategy is useful to forage in extremely dynamic environments. The developed model provides a basis for further investigation regarding the role of foraging mechanisms in the population dynamics of colonial animals, including the adaptive foraging behaviour of marine top predators to dynamic environmental conditions.

The first global deep-sea stable isotope assessment reveals the unique trophic ecology of Vampire Squid Vampyroteuthis infernalis (Cephalopoda).

Vampyroteuthis infernalis Chun, 1903, is a widely distributed deepwater cephalopod with unique morphology and phylogenetic position. We assessed its habitat and trophic ecology on a global scale via stable isotope analyses of a unique collection of beaks from 104 specimens from the Atlantic, Pacific and Indian Oceans. Cephalopods typically are active predators occupying a high trophic level (TL) and exhibit an ontogenetic increase in δ15N and TL. Our results, presenting the first global comparison for a deep-sea invertebrate, demonstrate that V. infernalis has an ontogenetic decrease in δ15N and TL, coupled with niche broadening. Juveniles are mobile zooplanktivores, while larger Vampyroteuthis are slow-swimming opportunistic consumers and ingest particulate organic matter. Vampyroteuthis infernalis occupies the same TL (3.0-4.3) over its global range and has a unique niche in deep-sea ecosystems. These traits have enabled the success and abundance of this relict species inhabiting the largest ecological realm on the planet.

Inter-annual isotopic niche segregation of wild humboldt penguins through years of different El Niño intensities.

The Humboldt Current System presents high interannual variability, influenced by the El Niño Southern Oscillation (ENSO), whose implications in wildlife are not fully understood. We studied the isotopic niche of wild Humboldt penguins at Punta San Juan (Peru) during the pre-moult foraging trip in 4 consecutive years (2008-2011) under known oceanographic (ENSO) conditions. Our results show that there is a clear isotopic niche segregation (on both δ13C and δ15N values) of wild Humboldt penguins among all years. Besides isotopic niche segregation, niche width also varied significantly among years. The larger isotopic niche displayed in 2008 reflected the opportunistic feeding behaviour of Humboldt penguins when oceanographic conditions were unfavourable (i.e. El Niño of strong intensity). In contrast, and despite strong segregation, penguins displayed a more specialist behaviour in years of mild environmental conditions (i.e. 2009 "warm-weak", 2010 "neutral" and 2011 "warm-moderate"). No evidence of sexual segregation in wild Humboldt penguins during the pre-moulting foraging trip was found. This study highlights the coping mechanisms of an endangered species to changes in environmental conditions (i.e. overall, from strong to neutral El Niño events), which should have important ramifications in the management of the marine ecosystem in Peru, particularly the one related to the anchovy industry.

Show your beaks and we tell you what you eat: Different ecology in sympatric Antarctic benthic octopods under a climate change context.

Sympatry can lead to higher competition under climate change and other environmental pressures, including in South Georgia, Antarctica, where the two most common octopod species, Adelieledone polymorpha and Pareledone turqueti, occur side by side. Since cephalopods are typically elusive animals, the ecology of both species is poorly known. As beaks of cephalopods are recurrently found in top predator's stomachs, we studied the feeding ecology of both octopods through the evaluation of niche overlapping and specific beak adaptations that both species present. A multidisciplinary approach combining carbon (δ13C) and nitrogen (δ15N) stable isotope signatures, mercury (Hg) analysis and biomaterials' engineering techniques was applied to investigate the beaks. An isotopic niche overlap of 95.6% was recorded for the juvenile stages of both octopod species, dropping to 19.2% for the adult stages. Both A. polymorpha and P. turqueti inhabit benthic ecosystems around South Georgia throughout their lifecycles (δ13C: -19.21 ±â€¯1.87‰, mean ±â€¯SD for both species) but explore trophic niches partially different during adult life stages (δ15N: 7.01 ±â€¯0.40‰, in A. polymorpha, and 7.84 ±â€¯0.65‰, in P. turqueti). The beaks of A. polymorpha are less dense and significantly less stiff than in P. turqueti. Beaks showed lower mercury concentration relative to muscle (A. polymorpha - beaks: 0.052 ±â€¯0.009  µg g-1, muscle: 0.322 ±â€¯0.088  µg g-1; P. turqueti - beaks: 0.038 ±â€¯0.009  µg g-1; muscle: 0.434 ±â€¯0.128  µg g-1). Overall, both octopods exhibit similar habitats but different trophic niches, related to morphology/function of beaks. The high Hg concentrations in both octopods can have negative consequences on their top predators and may increase under the present climate change context.

Environmentally driven sexual segregation in a marine top predator.

Sexual segregation in foraging occurs in many animal species, resulting in the partitioning of resources and reduction of competition between males and females, yet the patterns and drivers of such segregation are still poorly understood. We studied the foraging movements (GPS-tracking), habitat use (habitat modelling) and trophic ecology (stable isotope analysis) of female and male Cory's shearwaters Calonectris borealis during the mid chick-rearing period of six consecutive breeding seasons (2010-2015). We found a clear sexual segregation in foraging in years of greater environmental stochasticity, likely years of lower food availability. When food became scarce, females undertook much longer foraging trips, exploited more homogeneous water masses, had a larger isotopic niche, fed on lower trophic level prey and exhibited a lower body condition, when compared to males. Sexual competition for trophic resources may be stronger when environmental conditions are poor. A greater foraging success of one sex may result in differential body condition of pair mates when enduring parental effort, and ultimately, in an increased probability of breeding failure.

Sexual and individual foraging segregation in Gentoo penguins Pygoscelis papua from the Southern Ocean during an abnormal winter.

Knowledge about sexual segregation and gender-specific, or indeed individual specialization, in marine organisms has improved considerably in the past decade. In this context, we tested the "Intersexual Competition Hypothesis" for penguins by investigating the feeding ecology of Gentoo penguins during their austral winter non-breeding season. We considered this during unusual environmental conditions (i.e. the year 2009 had observations of high sea surface and air temperatures) in comparison with the long term average at Bird Island, South Georgia. Through conventional (i.e. stomach contents) and stable isotopic values from red blood cells, plasma and feathers of both male and female Gentoo penguins, we showed that there were significant differences between sexes, with males feeding mainly on fish (54% by mass) followed by crustaceans (38%) whereas females fed mainly on crustaceans (89% by mass) followed by fish (4%). Themisto gaudichaudii was the most important crustacean prey for males (64% by mass; 82% by number; 53% by frequency of occurrence) and females (63% by mass; 77% by number; 89% by frequency of occurrence), contrasting with all previous studies that found Antarctic krill Euphausia superba were generally the main prey. Stable isotopic data showed that, in terms of habitat use (based on δ 13C), there were significant differences in short-term carbon signatures between males and females (based on plasma and red blood cells), suggesting that both sexes explored different habitats, with females exploring more offshore pelagic waters and males feeding more in coastal benthic waters. Based on δ 15N, males fed on significantly higher trophic level than females (based on plasma and red blood cells), in agreement with our diet results., Thus, Gentoo penguins behave in a similar manner to other non-breeding penguins species (e.g. king, macaroni and rockhopper penguins), albeit at a smaller spatial scale (as they do not disperse as these other penguins do), in that they have a wider habitat and trophic niche during the Antarctic Winter (in comparison to Summer). We also detected individual specialization in feeding/trophic levels for each gender, with certain males feeding mainly on fish and certain females mainly on crustaceans, which may be driven the prevailing environmental conditions that lead individuals to search for alternative prey, and cause sexual diet segregation. Our results provide further information to help improve understanding about sexual segregation and individual specialization of marine organisms, while contributing valuable information on the winter diet for Antarctic monitoring programs and for modelling Antarctic marine food webs.