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Seabirds are increasingly used as bioindicators for assessing the chemical contamination of marine ecosystems, including by mercury (Hg) worldwide. However, some geographical areas are still poorly documented, as metropolitan France that is home to 28 seabird species including the black-legged kittiwake Rissa tridactyla, in the part of the southern limit of the North Atlantic range of the species. Here, we investigated Hg contamination and trophic ecology of black-legged kittiwakes breeding in the harbour of Boulogne-sur-Mer, Northern France. Mean blood Hg concentration was 4.81 ± 1.20 μg g -1 dw (dry weight), 3.66 ± 0.75 μg g -1 dw and 0.43 ± 0.07 μg g -1 dw for adult males, adult females, and chicks, respectively. According to Hg toxicity benchmarks for avian blood, 30% of the sampled adults were considered to be at moderate risk to Hg toxicity. Stable isotope and food analyses showed that highest δ 15 N values (reflecting a higher trophic position) were related to highest blood Hg concentrations in adult birds, and that Atlantic herring (Clupea harengus) and Atlantic mackerel (Scomber scombrus) were the main prey. Adult kittiwakes from Boulogne-sur-Mer showed Hg levels three times higher than those found in Arctic nesting kittiwakes, where sublethal effects have been documented. This study provides a first description of Hg contamination of black-legged kittiwakes breeding in France and calls for future ecotoxicological research to assess the vulnerability of this species in the southern part of its distribution range., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier Ltd.. All rights reserved.)

This study pioneers the reporting of Se isotopes in marine top predators and represents the most extensive Se isotopic characterization in animals to date. A methodology based on hydride generation─multicollector inductively coupled plasma mass spectrometry─was established for such samples. The study was conducted on various internal organs of giant petrels ( Macronectes spp.), encompassing bulk tissues (δ 82/78 Se bulk ), distinct Se-specific fractions such as selenoneine (δ 82/78 Se SEN ), and HgSe nanoparticles (δ 82/78 Se NPs ). The δ 82/78 Se bulk results (2.0-5.6‰) offer preliminary insights into the fate of Se in key internal organs of seabirds, including the liver, the kidneys, the muscle, and the brain. Notably, the liver of all individuals was enriched in heavier Se isotopes compared to other examined tissues. In nanoparticle fraction, δ 82/78 Se varies significantly across individuals (δ 82/78 Se NPs from 0.6 to 5.7‰, n = 8), whereas it exhibits remarkable consistency among tissues and individuals for selenoneine (δ 82/78 Se SEN , 1.7 ± 0.3‰, n = 8). Significantly, there was a positive correlation between the shift from δ 82/78 Se bulk to δ 82/78 Se SEN and the proportion of Se present as selenoneine in the internal organs. This pilot study proves that Se species-specific isotopic composition is a promising tool for a better understanding of Se species fate, sources, and dynamics in animals.

How Antarctic species are facing historical and new stressors remains under-surveyed and risks to wildlife are still largely unknown. Adélie penguins Pygoscelis adeliae are well-known bioindicators and sentinels of Antarctic ecosystem changes, a true canary in the coal mine. Immuno-haematological parameters have been proved to detect stress in wild animals, given their rapid physiological response that allows them tracking environmental changes and thus inferring habitat quality. Here, we investigated variation in Erythrocyte Nuclear Abnormalities (ENAs) and White Blood Cells (WBCs) in penguins from three clustered colonies in the Ross Sea, evaluating immuno-haematological parameters according to geography, breeding stage, and individual penguin characteristics such as sex, body condition and nest quality. Concentrations of mercury (Hg) and stable isotopes of carbon and nitrogen (as proxies of the penguin's trophic ecology) were analysed in feathers to investigate the association between stress biomarkers and Hg contamination in Adélie penguins. Colony and breeding stage were not supported as predictors of immuno-haematological parameters. ENAs and WBCs were respectively ∼30 % and ∼20 % higher in male than in female penguins. Body condition influenced WBCs, with penguins in the best condition having a ∼22 % higher level of WBCs than those in the worst condition. Nest position affected the proportion of micronuclei (MNs), with inner-nesting penguins having more than three times the proportion of MNs than penguins nesting in peripheral positions. Heterophils:Lymphocytes (H:L) ratio was not affected by any of the above predictors. Multiple factors acting as stressors are expected to increase prominently in Antarctic wildlife in the near future, therefore extensive monitoring aimed to assess the health status of penguin populations is mandatory., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 The Authors. Published by Elsevier B.V. All rights reserved.)

In vertebrates, fasting is an intricate physiological process associated with strong metabolic changes, yet its effect on pollutant residue variation is poorly understood. Here, we quantified long-term changes in plasma concentrations of 20 organochlorine and 16 perfluoroalkyl pollutants in king penguins Aptenodytes patagonicus during the breeding and molting fasts, which are marked by low and high levels of protein catabolism, respectively, and by strong lipid use. The profile of measured pollutants in plasma was dominated by perfluorooctanesulfonic acid (PFOS, initial relative contribution of 60%). Initial total pollutant concentrations were similar in molting (3.3-5.7 ng g -1 ww) and breeding penguins (range of 4.2-7.3 ng g -1 wet weight, ww). Long-term fasting (25 days) for molting and breeding led, respectively, to a 1.8- and 2.2-fold increase in total plasma pollutant concentrations, although the rate and direction of change were compound-specific. Hexachlorbenzene (HCB) and PFOS concentrations increased in plasma (net mobilization) during both types of fasting, likely due to lipid use. Plasma perfluoroundecanoate (PFUnDA) and perfluorotridecanoate (PFTrDA) concentrations increased in breeders (net mobilization) but decreased in molting individuals (net excretion), suggesting a significant incorporation of these pollutants into feathers. This study is a key contribution to our understanding of pollutant variation in blood during long-term fasting in wildlife.

Due to its persistence and potential ecological and health impacts, mercury (Hg) is a global pollutant of major concern that may reach high concentrations even in remote polar oceans. In contrast to the Arctic Ocean, studies documenting Hg contamination in the Southern Ocean are spatially restricted and large-scale monitoring is needed. Here, we present the first circumpolar assessment of Hg contamination in Antarctic marine ecosystems. Specifically, the Adélie penguin (Pygoscelis adeliae) was used as a bioindicator species, to examine regional variation across 24 colonies distributed across the entire Antarctic continent. Mercury was measured on body feathers collected from both adults (n = 485) and chicks (n = 48) between 2005 and 2021. Because penguins' diet represents the dominant source of Hg, feather δ 13 C and δ 15 N values were measured as proxies of feeding habitat and trophic position. As expected, chicks had lower Hg concentrations (mean ± SD: 0.22 ± 0.08 μg·g ‒1 ) than adults (0.49 ± 0.23 μg·g ‒1 ), likely because of their shorter bioaccumulation period. In adults, spatial variation in feather Hg concentrations was driven by both trophic ecology and colony location. The highest Hg concentrations were observed in the Ross Sea, possibly because of a higher consumption of fish in the diet compared to other sites (krill-dominated diet). Such large-scale assessments are critical to assess the effectiveness of the Minamata Convention on Mercury. Owing to their circumpolar distribution and their ecological role in Antarctic marine ecosystems, Adélie penguins could be valuable bioindicators for tracking spatial and temporal trends of Hg across Antarctic waters in the future., (© 2023. The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature.)

Mercury (Hg) pollution is a global problem affecting remote areas of the open ocean, but the bioaccumulation of this neurotoxic pollutant in tropical top predators remains poorly documented. The objective of this study was to determine Hg contamination of the seabird community nesting on Clipperton Island using blood and feathers to investigate short and longer-term contamination, respectively. We examined the significance of various factors (species, sex, feeding habitat [δ 13 C] and trophic position [δ 15 N]) on Hg concentrations in six seabird species. Among species, Great Frigatebirds had the highest Hg concentrations in blood and feathers, boobies had intermediate values, and Brown Noddies and Sooty Terns the lowest. At the interspecific level, although δ 13 C values segregated boobies from frigatebirds and noddies/terns, Hg concentrations were explained by neither δ 13 C nor δ 15 N values. At the intraspecific level, both Hg concentrations in blood and feathers show relatively small variations (16-32 and 26-74%, respectively), suggesting that feeding ecology had low seasonal variation among individuals. Despite most species being sexually dimorphic, differences in Hg contamination according to sex was detected only in Brown Boobies during the breeding period. Indeed, female Brown Boobies feed at a higher trophic level and in a different area than males during this period, resulting in higher blood Hg concentrations. The present study also shows that most of the seabirds sampled at Clipperton Island had little or no exposure to Hg toxicity, with 30% in the no risk category and 70% in the low risk category., (© 2023. The Author(s), under exclusive licence to Springer Science+Business Media, LLC, part of Springer Nature.)

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

The oceans become increasingly contaminated as a result of global industrial production and consumer behaviour, and this affects wildlife in areas far removed from sources of pollution. Migratory seabirds such as storm-petrels may forage in areas with different contaminant levels throughout the annual cycle and may show a carry-over of mercury from the winter quarters to the breeding sites. In this study, we compared mercury levels among seven species of storm-petrels breeding on the Antarctic South Shetlands and subantarctic Kerguelen Islands, in temperate waters of the Chatham Islands, New Zealand, and in temperate waters of the Pacific off Mexico. We tested for differences in the level of contamination associated with breeding and inter-breeding distribution and trophic position. We collected inert body feathers and metabolically active blood samples in ten colonies, reflecting long-term (feathers) and short-term (blood) exposures during different periods ranging from early non-breeding (moult) to late breeding. Feathers represent mercury accumulated over the annual cycle between two successive moults. Mercury concentrations in feathers ranged over more than an order of magnitude among species, being lowest in subantarctic Grey-backed Storm-petrels (0.5 μg g -1 dw) and highest in subtropical Leach's Storm-petrels (7.6 μg g -1 dw, i.e. posing a moderate toxicological risk). Among Antarctic Storm-petrels, Black-bellied Storm-petrels had threefold higher values than Wilson's Storm-petrels, and in both species, birds from the South Shetlands (Antarctica) had threefold higher values than birds from Kerguelen (subantarctic Indian Ocean). Blood represents mercury taken up over several weeks, and showed similar trends, being lowest in Grey-backed Storm-petrels from Kerguelen (0.5 μg g -1 dw) and highest in Leach's Storm-petrels (3.6 μg g -1 dw). Among Antarctic storm-petrels, species differences in the blood samples were similar to those in feathers, but site differences were less consistent. Over the breeding season, mercury decreased in blood samples of Antarctic Wilson's Storm-petrels, but did not change in Wilson's Storm-petrels from Kerguelen or in Antarctic Black-bellied Storm-petrels. In summary, we found that mercury concentrations in storm-petrels varied due to the distribution of species and differences in prey choice. Depending on prey choices, Antarctic storm-petrels can have similar mercury concentrations as temperate species. The lowest contamination was observed in subantarctic species and populations. The study shows how seabirds, which accumulate dietary pollutants in their tissues in the breeding and non-breeding seasons, can be used to survey marine pollution. Storm-petrels with their wide distributions and relatively low trophic levels may be especially useful, but more detailed knowledge on their prey choice and distributions is needed., (© 2023. The Author(s).)

Mercury (Hg) is a highly toxic metal that adversely impacts human and wildlife health. The amount of Hg released globally in the environment has increased steadily since the Industrial Revolution, resulting in growing contamination in biota. Seabirds have been extensively studied to monitor Hg contamination in the world's oceans. Multidecadal increases in seabird Hg contamination have been documented in polar, temperate and subtropical regions, whereas in tropical regions they are largely unknown. Since seabirds accumulate Hg mainly from their diet, their trophic ecology is fundamental in understanding their Hg exposure over time. Here, we used the sooty tern (Onychoprion fuscatus), the most abundant tropical seabird, as bioindicator of temporal variations in Hg transfer to marine predators in tropical ecosystems, in response to trophic changes and other potential drivers. Body feathers were sampled from 220 sooty terns, from museum specimens (n = 134) and free-living birds (n = 86) from Ascension Island, in the South Atlantic Ocean, over 145 years (1876-2021). Chemical analyses included (i) total- and methyl-Hg, and (ii) carbon (δ 1 ³C) and nitrogen (δ 15 N) stable isotopes, as proxies of foraging habitat and trophic position, respectively, to investigate the relationship between trophic ecology and Hg contamination over time. Despite current regulations on its global emissions, mean Hg concentrations were 58.9% higher in the 2020s (2.0 μg g -1 , n = 34) than in the 1920s (1.2 μg g -1 , n = 107). Feather Hg concentrations were negatively and positively associated with δ 1 ³C and δ 15 N values, respectively. The sharp decline of 2.9 ‰ in δ 1 ³C values over time indicates ecosystem-wide changes (shifting primary productivity) in the tropical South Atlantic Ocean and can help explain the observed increase in terns' feather Hg concentrations. Overall, this study provides invaluable information on how ecosystem-wide changes can increase Hg contamination of tropical marine predators and reinforces the need for long-term regulations of harmful contaminants at the global scale., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

The present work explores for the first time the potential of formic acid on the extraction of tiemannite (HgSe) nanoparticles from seabird tissues, in particular giant petrels. Mercury (Hg) is considered one of the top ten chemicals of major public health concern. However, the fate and metabolic pathways of Hg in living organisms remain unknown. Methylmercury (MeHg), largely produced by microbial activity in the aquatic ecosystems is biomagnified in the trophic web. HgSe is considered the end-product of MeHg demethylation in biota and an increasing number of studies focuses on the characterization of this solid compound to understand its biomineralization. In this study, a conventional enzymatic treatment is compared with a simpler and environmentally friendly extraction by using formic acid (5 mL of = 50 % formic acid) as exclusive reagent. The analyses by spICP-MS of the resulting extracts from a variety of seabird biological tissues (liver, kidneys, brain, muscle) reveal comparable results by both extraction approaches in terms of nanoparticles stability and extraction efficiency. Therefore, the results included in this work demonstrate the good performance of employing organic acid as simple, cost effective and green procedure to extract HgSe nanoparticles from animal tissues. Moreover, an alternative consisting of a classical enzymatic procedure but with ultrasonic assistance reducing the extraction time from 12 h to 2 min is also described for the first time. The sample processing methodologies developed, combined with spICP-MS, have emerged as powerful tools for the rapid screening and quantification of HgSe nanoparticles in animal tissues. Finally, this combination allowed us to identify the possible occurrence of Cd particles and As particles associated with HgSe NPs in seabirds., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023. Published by Elsevier B.V.)

The use of cephalopod beaks in ecological and population dynamics studies has allowed major advances of our knowledge on the role of cephalopods in marine ecosystems in the last 60 years. Since the 1960's, with the pioneering research by Malcolm Clarke and colleagues, cephalopod beaks (also named jaws or mandibles) have been described to species level and their measurements have been shown to be related to cephalopod body size and mass, which permitted important information to be obtained on numerous biological and ecological aspects of cephalopods in marine ecosystems. In the last decade, a range of new techniques has been applied to cephalopod beaks, permitting new kinds of insight into cephalopod biology and ecology. The workshop on cephalopod beaks of the Cephalopod International Advisory Council Conference (Sesimbra, Portugal) in 2022 aimed to review the most recent scientific developments in this field and to identify future challenges, particularly in relation to taxonomy, age, growth, chemical composition (i.e., DNA, proteomics, stable isotopes, trace elements) and physical (i.e., structural) analyses. In terms of taxonomy, new techniques (e.g., 3D geometric morphometrics) for identifying cephalopods from their beaks are being developed with promising results, although the need for experts and reference collections of cephalopod beaks will continue. The use of beak microstructure for age and growth studies has been validated. Stable isotope analyses on beaks have proven to be an excellent technique to get valuable information on the ecology of cephalopods (namely habitat and trophic position). Trace element analyses is also possible using beaks, where concentrations are significantly lower than in other tissues (e.g., muscle, digestive gland, gills). Extracting DNA from beaks was only possible in one study so far. Protein analyses can also be made using cephalopod beaks. Future challenges in research using cephalopod beaks are also discussed., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Xavier, Golikov, Queirós, Perales-Raya, Rosas-Luis, Abreu, Bello, Bustamante, Capaz, Dimkovikj, González, Guímaro, Guerra-Marrero, Gomes-Pereira, Kubodera, Laptikhovsky, Lefkaditou, Lishchenko, Luna, Liu, Pierce, Pissarra, Reveillac, Romanov, Rosa, Roscian, Rose-Mann, Rouget, Sánchez, Sánchez-Márquez, Seixas, Souquet, Varela, Vidal and Cherel.)

Mercury (Hg) is a toxic trace element widely distributed in the environment, which particularly accumulates in top predators, including seabirds. Among seabirds, large gulls (Larus sp) are generalist feeders, foraging in both terrestrial and marine habitats, making them relevant bioindicators of local coastal Hg contamination. In the present study, we reported blood Hg concentrations in adults and chicks of four different gull species breeding on the French Atlantic coast: the European herring gull (Larus argentatus), the Lesser black-backed gull (L. fuscus), the Great black-backed gull (L. marinus) and the Yellow-legged gull (L. michahellis). We also investigated the potential role of foraging ecology in shaping Hg contamination across species, using the unique combination of three dietary tracers (carbon, nitrogen and sulfur stable isotopes) and biologging (GPS tracking). A high concentration of Hg was associated with high trophic position and a marine diet in gulls, which was corroborated by birds' space use strategy during foraging trips. Adults of all four species reached Hg concentrations above reported toxicity thresholds. Specifically, adults of Great black-backed gulls had a high trophic marine specialized diet and significantly higher Hg concentrations than the three other species. Blood Hg was 4-7 times higher in adults than in chicks, although chicks of all species received mainly marine and high trophic position prey, which is expected to be the cause of blood Hg concentrations of toxic concern. By using both stable isotopes and GPS tracking, the present study provides compelling insights on the main feeding habits driving Hg contamination in a seabird assemblage feeding in complex coastal environments., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Mercury contamination is a major threat to the global environment, and is still increasing in some regions despite international regulations. The methylated form of mercury is hazardous to biota, yet its sublethal effects are difficult to detect in wildlife. Body condition can vary in response to stressors, but previous studies have shown mixed effects of mercury on body condition in wildlife. Using birds as study organisms, we provide the first quantitative synthesis of the effect of mercury on body condition in animals. In addition, we explored the influence of intrinsic, extrinsic and methodological factors potentially explaining cross-study heterogeneity in results. We considered experimental and correlative studies carried out in adult birds and chicks, and mercury exposure inferred from blood and feathers. Most experimental investigations (90%) showed a significant relationship between mercury concentrations and body condition. Experimental exposure to mercury disrupted nutrient (fat) metabolism, metabolic rates, and food intake, resulting in either positive or negative associations with body condition. Correlative studies also showed either positive or negative associations, of which only 14% were statistically significant. Therefore, the overall effect of mercury concentrations on body condition was null in both experimental (estimate ± SE = 0.262 ± 0.309, 20 effect sizes, five species) and correlative studies (-0.011 ± 0.020, 315 effect sizes, 145 species). The single and interactive effects of age class and tissue type were accounted for in meta-analytic models of the correlative data set, since chicks and adults, as well as blood and feathers, are known to behave differently in terms of mercury accumulation and health effects. Of the 15 moderators tested, only wintering status explained cross-study heterogeneity in the correlative data set: free-ranging wintering birds were more likely to show a negative association between mercury and body condition. However, wintering effect sizes were limited to passerines, further studies should thus confirm this trend in other taxa. Collectively, our results suggest that (i) effects of mercury on body condition are weak and mostly detectable under controlled conditions, and (ii) body condition indices are unreliable indicators of mercury sublethal effects in the wild. Food availability, feeding rates and other sources of variation that are challenging to quantify likely confound the association between mercury and body condition in natura. Future studies could explore the metabolic effects of mercury further using designs that allow for the estimation and/or manipulation of food intake in both wild and captive birds, especially in under-represented life-history stages such as migration and overwintering., (© 2022 Cambridge Philosophical Society.)

Penguins lost the ability to fly more than 60 million years ago, subsequently evolving a hyper-specialized marine body plan. Within the framework of a genome-scale, fossil-inclusive phylogeny, we identify key geological events that shaped penguin diversification and genomic signatures consistent with widespread refugia/recolonization during major climate oscillations. We further identify a suite of genes potentially underpinning adaptations related to thermoregulation, oxygenation, diving, vision, diet, immunity and body size, which might have facilitated their remarkable secondary transition to an aquatic ecology. Our analyses indicate that penguins and their sister group (Procellariiformes) have the lowest evolutionary rates yet detected in birds. Together, these findings help improve our understanding of how penguins have transitioned to the marine environment, successfully colonizing some of the most extreme environments on Earth., (© 2022. The Author(s).)

Niche theory predicts that to reduce competition for the same resource, sympatric ecologically similar species should exploit divergent niches and segregate in one or more dimensions. Seasonal variations in environmental conditions and energy requirements can influence the mechanisms and the degree of niche segregation. However, studies have overlooked the multi-dimensional aspect of niche segregation over the whole annual cycle, and key facets of species co-existence still remain ambiguous. The present study provides insights into the niche use and partitioning of two morphologically and ecologically similar seabirds, the common (CDP, Pelecanoides urinatrix) and the South Georgian diving petrel (SGDP, Pelecanoides georgicus). Using phenology, at-sea distribution, diving behavior and isotopic data (during the incubation, chick-rearing and non-breeding periods), we show that the degree of partitioning was highly stage-dependent. During the breeding season, the greater niche segregation during chick-rearing than incubation supported the hypothesis that resource partitioning increases during energetically demanding periods. During the post breeding period, while species-specific latitudinal differences were expected (species specific water mass preference), CDP and SGDP also migrated in divergent directions. This segregation in migration area may not be only a response to the selective pressure arising from competition avoidance between sympatric species, but instead, could reflect past evolutionary divergence. Such stage-dependent and context-dependent niche segregation demonstrates the importance of integrative approaches combining techniques from different fields, throughout the entire annual cycle, to better understand the co-existence of ecologically similar species. This is particularly relevant in order to fully understand the short and long-term effects of ongoing environmental changes on species distributions and communities.This work demonstrates the need of integrative multi-dimensional approaches combining concepts and techniques from different fields to understand the mechanism and causal factors of niche segregation., (© 2022. The Author(s).)

Antarctic marine ecosystems are often considered to be pristine environments, yet wildlife in the polar regions may still be exposed to high levels of environmental contaminants. Here, we measured total mercury (THg) concentrations in blood samples from adult brown skuas Stercorarius antarcticus lonnbergi (n = 82) from three breeding colonies south of the Antarctic Polar Front in the Southern Ocean (southwest Atlantic region): (i) Bahía Esperanza/Hope Bay, Antarctic Peninsula; (ii) Signy Island, South Orkney Islands; and, (iii) Bird Island, South Georgia. Blood THg concentrations increased from the Antarctic Peninsula towards the Antarctic Polar Front, such that Hg contamination was lowest at Bahía Esperanza/Hope Bay (mean ± SD, 0.95 ± 0.45 μg g -1 dw), intermediate at Signy Island (3.42 ± 2.29 μg g -1 dw) and highest at Bird Island (4.47 ± 1.10 μg g -1 dw). Blood THg concentrations also showed a weak positive correlation with δ 15 N values, likely reflecting the biomagnification process. Males had higher Hg burdens than females, which may reflect deposition of Hg into eggs by females or potentially differences in their trophic ecology. These data provide important insights into intraspecific variation in contamination and the geographic transfer of Hg to seabirds in the Southern Ocean., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Tiemannite (HgSe) is considered the end-product of methylmercury (MeHg) demethylation in vertebrates. The biomineralization of HgSe nanoparticles (NPs) is understood to be an efficient MeHg detoxification mechanism; however, the process has not yet been fully elucidated. In order to contribute to the understanding of complex Hg metabolism and HgSe NPs formation, the Hg isotopic signatures of 40 samples of 11 giant petrels were measured. This seabird species is one of the largest avian scavengers in the Southern Ocean, highly exposed to MeHg through their diet, reaching Hg concentrations in the liver up to more than 900 µg g -1 . This work constitutes the first species-specific isotopic measurement (δ 202 Hg, Δ 199 Hg) of HgSe NPs in seabirds and the largest characterization of this compound in biota. Similar δ 202 Hg values specifically associated to HgSe (δ 202 Hg HgSe ) and tissues (δ 202 Hg bulk ) dominated by inorganic Hg species were found, suggesting that no isotopic fractionation is induced during the biomineralization step from the precursor (demethylated) species. In contrast, the largest variations between δ 202 Hg bulk and δ 202 Hg HgSe were observed in muscle and brain tissues. This could be attributed to the higher fraction of Hg present as MeHg in these tissues. Hg-biomolecules screening highlights the importance of the isotopic characterization of these (unknown) complexes., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Birds are principally exposed to selenium (Se) through their diet. In long-lived and top predator seabirds, such as the giant petrel, extremely high concentrations of Se are found. Selenium speciation in biota has aroused great interest in recent years; however, there is a lack of information about the chemical form of Se in (sea)birds. The majority of publications focus on the growth performance and antioxidant status in broilers in relation to Se dietary supplementation. The present work combines elemental and molecular mass spectrometry for the characterization of Se species in wild (sea)birds. A set of eight giant petrels ( Macronectes sp.) with a broad age range from the Southern Ocean were studied. Selenoneine, a Se-analogue of ergothioneine, was identified for the first time in wild avian species. This novel Se-compound, previously reported in fish, constitutes the major Se species in the water-soluble fraction of all of the internal tissues and blood samples analyzed. The levels of selenoneine found in giant petrels are the highest reported in animal tissues until now, supporting the trophic transfer in the marine food web. The characterization of selenoneine in the brain, representing between 78 and 88% of the total Se, suggests a crucial role in the nervous system. The dramatic decrease of selenoneine (from 68 to 3%) with an increase of Hg concentrations in the liver strongly supports the hypothesis of its key role in Hg detoxification.

The New Zealand huia (Heteralocha acutirostris) had the most extreme bill sexual dimorphism among modern birds. Given the quick extinction of the species, the cause of the dimorphism could only be hypothesised to reflect different trophic niches and reduce male/female competition. We tested that hypothesis by combining museum specimens, geometric morphometrics, and isotopic analyses. We used geometric morphometrics to describe bill shape; measured bulk (δ 15 N bulk ) and (δ 13 C bulk ) values from feather as proxies of the birds' foraging habitat and diet; and compared compound-specific stable isotopes analyses (CSIA) of nitrogen in amino acids (δ 15 N AA ) in male-female pairs to estimate their trophic position. Sexes had significantly different, but overlapping feather δ 15 N bulk and δ 13 C bulk values, but δ 15 N AA indicated identical trophic positions and δ 15 N bulk was not related to bill shape. Trophic position was less variable among females, consistent with a specialised foraging behaviour and, thus, supporting a partial male/female foraging segregation., (© 2021. The Author(s).)

Interspecific introgression can occur between species that evolve rapidly within an adaptive radiation. Pachyptila petrels differ in bill size and are characterised by incomplete reproductive isolation, leading to interspecific gene flow. Salvin's prion (Pachyptila salvini), whose bill width is intermediate between broad-billed (P. vittata) and Antarctic (P. desolata) prions, evolved through homoploid hybrid speciation. MacGillivray's prion (P. macgillivrayi), known from a single population on St Paul (Indian Ocean), has a bill width intermediate between salvini and vittata and could also be the product of interspecies introgression or hybrid speciation. Recently, another prion population phenotypically similar to macgillivrayi was discovered on Gough (Atlantic Ocean), where it breeds 3 months later than vittata. The similarity in bill width between the medium-billed birds on Gough and macgillivrayi suggest that they could be closely related. In this study, we used genetic and morphological data to infer the phylogenetic position and evolutionary history of P. macgillivrayi and the Gough medium-billed prion relative other Pachyptila taxa, to determine whether species with medium bill widths evolved through common ancestry or convergence. We found that Gough medium-billed prions belong to the same evolutionary lineage as macgillivrayi, representing a new population of MacGillivray's prion that originated through a colonisation event from St Paul. We show that macgillivrayi's medium bill width evolved through divergence (genetic drift) and independently from that of salvini, which evolved through hybridisation (gene flow). This represents the independent convergence towards a similarly medium-billed phenotype. The newly discovered MacGillivray's prion population on Gough is of utmost conservation relevance, as the relict macgillivrayi population in the Indian Ocean is very small., (© 2021. The Author(s).)

A prerequisite for environmental and toxicological applications of mercury (Hg) stable isotopes in wildlife and humans is quantifying the isotopic fractionation of biological reactions. Here, we measured stable Hg isotope values of relevant tissues of giant petrels ( Macronectes spp.). Isotopic data were interpreted with published HR-XANES spectroscopic data that document a stepwise transformation of methylmercury (MeHg) to Hg-tetraselenolate (Hg(Sec) 4 ) and mercury selenide (HgSe) (Sec = selenocysteine). By mathematical inversion of isotopic and spectroscopic data, identical δ 202 Hg values for MeHg (2.69 ± 0.04‰), Hg(Sec) 4 (-1.37 ± 0.06‰), and HgSe (0.18 ± 0.02‰) were determined in 23 tissues of eight birds from the Kerguelen Islands and Adélie Land (Antarctica). Isotopic differences in δ 202 Hg between MeHg and Hg(Sec) 4 (-4.1 ± 0.1‰) reflect mass-dependent fractionation from a kinetic isotope effect due to the MeHg → Hg(Sec) 4 demethylation reaction. Surprisingly, Hg(Sec) 4 and HgSe differed isotopically in δ 202 Hg (+1.6 ± 0.1‰) and mass-independent anomalies (i.e., changes in Δ 199 Hg of ≤0.3‰), consistent with equilibrium isotope effects of mass-dependent and nuclear volume fractionation from Hg(Sec) 4 → HgSe biomineralization. The invariance of species-specific δ 202 Hg values across tissues and individual birds reflects the kinetic lability of Hg-ligand bonds and tissue-specific redistribution of MeHg and inorganic Hg, likely as Hg(Sec) 4 . These observations provide fundamental information necessary to improve the interpretation of stable Hg isotope data and provoke a revisitation of processes governing isotopic fractionation in biota and toxicological risk assessment in wildlife.

Oceanic frontal zones have been shown to deeply influence the distribution of primary producers and, at the other extreme of the trophic web, top predators. However, the relationship between these structures and intermediate trophic levels is much more obscure. In this paper we address this knowledge gap by comparing acoustic measurements of mesopelagic fish concentrations to satellite-derived fine-scale Lagrangian Coherent Structures in the Indian sector of the Southern Ocean. First, we demonstrate that higher fish concentrations occur more frequently in correspondence with strong Lagrangian Coherent Structures. Secondly, we illustrate that, while increased fish densities are more likely to be observed over these structures, the presence of a fine-scale feature does not imply a concomitant fish accumulation, as other factors affect fish distribution. Thirdly, we show that, when only chlorophyll-rich waters are considered, front intensity modulates significantly more the local fish concentration. Finally, we discuss a model representing fish movement along Lagrangian features, specifically built for mid-trophic levels. Its results, obtained with realistic parameters, are qualitatively consistent with the observations and the spatio-temporal scales analysed. Overall, these findings may help to integrate intermediate trophic levels in trophic models, which can ultimately support management and conservation policies., (© 2021. The Author(s).)

Marine litter has become a global issue with 'garbage patches' documented in all ocean gyres. The Pacific and Atlantic garbage patches have been well described, but there are few empirical data for the Indian Ocean. In the austral summer 2019-2020, we conducted an at-sea survey of macro-litter in the rarely investigated south-west Indian Ocean. Over 24 days, 1623 man-made items were observed including plastic fragments, packaging and fishing-related items during 216 h of observations covering 5464 km. More than 99% of the litter items were plastics of which almost 60% were white. Floating litter was patchily distributed with only five items (0.2%) recorded south of 40°S (0.1 items·km - 2 ). Half of the items were encountered over a two-day period south-east of Madagascar (30°S; 59-67°E; 75.2 items·km - 2 ). Our survey detected an accumulation of litter in the southern Indian Ocean and demonstrated that this area warrants more research., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Capital breeders accumulate nutrients prior to egg development, then use these stores to support offspring development. In contrast, income breeders rely on local nutrients consumed contemporaneously with offspring development. Understanding such nutrient allocations is critical to assessing life-history strategies and habitat use. Despite the contrast between these strategies, it remains challenging to trace nutrients from endogenous stores or exogenous food intake into offspring. Here, we tested a new solution to this problem. Using tissue samples collected opportunistically from wild emperor penguins Aptenodytes forsteri, which exemplify capital breeding, we hypothesized that the stable carbon (δ 13 C) and nitrogen (δ 15 N) isotope values of individual amino acids (AAs) in endogenous stores (e.g. muscle) and in egg yolk and albumen reflect the nutrient sourcing that distinguishes capital versus income breeding. Unlike other methods, this approach does not require untested assumptions or diet sampling. We found that over half of essential AAs had δ 13 C values that did not differ between muscle and yolk or albumen, suggesting that most of these AAs were directly routed from muscle into eggs. In contrast, almost all non-essential AAs differed in δ 13 C values between muscle and yolk or between muscle and albumen, suggesting de novo synthesis. Over half of AAs that have labile nitrogen atoms (i.e. 'trophic' AA) had higher δ 15 N values in yolk and albumen than in muscle, suggesting that they were transaminated during their routing into egg tissue. This effect was smaller for AAs with less labile nitrogen atoms (i.e. 'source' AA). Our results indicate that the δ 15 N offset between trophic-source AAs (Δ 15 N trophic-source ) may provide an index of the extent of capital breeding. The value of emperor penguin Δ 15 N Pro-Phe was higher in yolk and albumen than in muscle, reflecting the mobilization of endogenous stores; in comparison, the value of Δ 15 N Pro-Phe was similar across muscle and egg tissue in previously published data for income-breeding herring gulls Larus argentatus smithsonianus. Our results provide a quantitative basis for using AA δ 13 C and δ 15 N, and isotopic offsets among AAs (e.g. Δ 15 N Pro-Phe ), to explore the allocation of endogenous versus exogenous nutrients across the capital versus income spectrum of avian reproduction., (© 2020 British Ecological Society.)

Technical advances in 3D imaging have contributed to quantifying and understanding biological variability and complexity. However, small, dry-sensitive objects are not easy to reconstruct using common and easily available techniques such as photogrammetry, surface scanning, or micro-CT scanning. Here, we use cephalopod beaks as an example as their size, thickness, transparency, and dry-sensitive nature make them particularly challenging. We developed a new, underwater, photogrammetry protocol in order to add these types of biological structures to the panel of photogrammetric possibilities.We used a camera with a macrophotography mode in a waterproof housing fixed in a tank with clear water. The beak was painted and fixed on a colored rotating support. Three angles of view, two acquisitions, and around 300 pictures per specimen were taken in order to reconstruct a full 3D model. These models were compared with others obtained with micro-CT scanning to verify their accuracy.The models can be obtained quickly and cheaply compared with micro-CT scanning and have sufficient precision for quantitative interspecific morphological analyses. Our work shows that underwater photogrammetry is a fast, noninvasive, efficient, and accurate way to reconstruct 3D models of dry-sensitive objects while conserving their shape. While the reconstruction of the shape is accurate, some internal parts cannot be reconstructed with photogrammetry as they are not visible. In contrast, these structures are visible using reconstructions based on micro-CT scanning. The mean difference between both methods is very small (10 -5 to 10 -4  mm) and is significantly lower than differences between meshes of different individuals.This photogrammetry protocol is portable, easy-to-use, fast, and reproducible. Micro-CT scanning, in contrast, is time-consuming, expensive, and nonportable. This protocol can be applied to reconstruct the 3D shape of many other dry-sensitive objects such as shells of shellfish, cartilage, plants, and other chitinous materials., Competing Interests: None declared., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Adeno-associated virus (AAV) vectors are considered efficient vectors for gene transfer, as illustrated by recent successful clinical trials targeting retinal or neurodegenerative disorders. However, limitations as host immune responses to AAV capsid or transduction of limited regions must still be overcome. Here, we focused on locoregional (LR) intravenous perfusion vector delivery that allows transduction of large muscular areas and is considered to be less immunogenic than intramuscular (IM) injection. To confirm this hypothesis, we injected 6 cynomolgus monkeys with an AAV serotype 8 (AAV8) vector encoding for the highly immunogenic GFP driven by either a muscle-specific promoter (n = 3) or a cytomegalovirus (CMV) promoter (n = 3). We report that LR delivery allows long-term GFP expression in the perfused limb (up to 1 year) despite the initiation of a peripheral transgene-specific immune response. The analysis of the immune status of the perfused limb shows that LR delivery induces persisting inflammation. However, this inflammation is not sufficient to result in transgene clearance and is balanced by resident regulatory T cells. Overall, our results suggest that LR delivery promotes persisting transgene expression by induction of Treg cells in situ and might be a safe alternative to IM route to target large muscle territories for the expression of secreted therapeutic factors., Competing Interests: G.G., M.G., M.D., N.J., V.P., M.J., A.L., J.L.D., J.-Y.D., C.L.G., P.M., Y.C., and O.A. declare no competing interests. P.M. is an employee of Asklepios BioPharmaceutical., (© 2021.)

In vivo and in vitro evidence for detoxification of methylmercury (MeHg) as insoluble mercury selenide (HgSe) underlies the central paradigm that mercury exposure is not or little hazardous when tissue Se is in molar excess (Se:Hg > 1). However, this hypothesis overlooks the binding of Hg to selenoproteins, which lowers the amount of bioavailable Se that acts as a detoxification reservoir for MeHg, thereby underestimating the toxicity of mercury. This question was addressed by determining the chemical forms of Hg in various tissues of giant petrels Macronectes spp. using a combination of high energy-resolution X-ray absorption near edge structure and extended X-ray absorption fine structure spectroscopy, and transmission electron microscopy coupled to elemental mapping. Three main Hg species were identified, a MeHg-cysteinate complex, a four-coordinate selenocysteinate complex (Hg(Sec) 4 ), and a HgSe precipitate, together with a minor dicysteinate complex Hg(Cys) 2 . The amount of HgSe decreases in the order liver > kidneys > brain = muscle, and the amount of Hg(Sec) 4 in the order muscle > kidneys > brain > liver. On the basis of biochemical considerations and structural modeling, we hypothesize that Hg(Sec) 4 is bound to the carboxy-terminus domain of selenoprotein P (SelP) which contains 12 Sec residues. Structural flexibility allows SelP to form multinuclear Hg x (Se,Sec) y complexes, which can be biomineralized to HgSe by protein self-assembly. Because Hg(Sec) 4 has a Se:Hg molar ratio of 4:1, this species severely depletes the stock of bioavailable Se for selenoprotein synthesis and activity to one μg Se/g dry wet in the muscle of several birds. This concentration is still relatively high because selenium is naturally abundant in seawater, therefore it probably does not fall below the metabolic need for essential selenium. However, this study shows that this may not be the case for terrestrial animals, and that muscle may be the first tissue potentially injured by Hg toxicity.

Trace metals such as Cu, Hg, and Zn have been widely investigated in marine ecotoxicological studies considering their bioaccumulation, transfer along trophic webs, and the risks they pose to ecosystems and human health. Comparatively, Li has received little attention, although this element is increasingly used in the high-tech, ceramics/glass, and medication industries. Here, we report Li concentrations in more than 400 samples, including whole organisms and different organs of bivalves, cephalopods, crustaceans, and fish. We investigated species from three contrasting biogeographic areas, i.e. temperate (Bay of Biscay, northeast Atlantic Ocean), tropical (New Caledonia, Pacific Ocean), and subpolar climates (Kerguelen Islands, southern Indian Ocean), among diverse trophic groups (filter-feeders to meso-predators) and habitats (benthic, demersal, and pelagic). Although Li is homogeneously distributed in the ocean (at 0.18 μg/mL), Li concentrations in soft tissues vary greatly, from 0.01 to 1.20 μg/g dry weight. Multiple correspondence analyses reveal two clusters of high and low Li concentrations. Li distributions in marine organisms appear to be mostly geographically independent, though our results highlight a temperature dependency in fish muscles. Li is consistently bio-reduced through the trophic webs, with filter-feeders showing the highest concentrations and predatory fish the lowest. Strong variations are observed among organs, consistent with the biochemical similarity between Na and Li during transport in the brain and in osmoregulatory organs. Fish gills and kidneys show relatively high Li concentrations (0.26 and 0.15 μg/g, respectively) and fish brains show a large range of Li contents (up to 0.34 μg/g), whereas fish liver and muscles are Li depleted (0.07 ± 0.03 and 0.06 ± 0.08 μg/g, respectively). Altogether, these results provide the first exhaustive baseline for future Li ecotoxicology studies in marine coastal environments., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Seabirds accumulate significant amounts of mercury (Hg) due to their long-life span together with their medium to high trophic position in marine food webs. Hg speciation and Hg isotopic analyses of total Hg in different tissues (pectoral muscles, liver, brain, kidneys, blood and feathers) were assessed to investigate their detoxification mechanisms. Three species with contrasted ecological characteristics were studied: the Antarctic prion (zooplankton feeder), the white-chinned petrel (pelagic generalist consumer) and the southern giant petrel (scavenger on seabirds and marine mammals). The difference of mass-dependent fractionation (MDF, δ 202 Hg) values between liver and muscles (up to 0.94 ‰) in all three seabirds strongly suggests hepatic demethylation of the isotopically lighter methylmercury (MeHg) and subsequent redistribution of the isotopically heavier fraction of MeHg towards the muscles. Similarly, higher δ 202 Hg values in feathers (up to 1.88 ‰) relative to muscles and higher proportion of MeHg in feathers (94-97%) than muscles (30-70%) likely indicate potential MeHg demethylation in muscle and preferential excretion of MeHg (isotopically heavier) in the growing feathers during moult. The extents of these key detoxification processes were strongly dependent on the species-specific detoxification strategies and levels of dietary MeHg exposure. We also found higher mass-independent fractionation (MIF, Δ 199 Hg) values in feathers relative to internal tissues, possibly due to different integration times of Hg exposure between permanently active organs and inert tissues as feathers. Hg isotope variations reported in this study show evidence of detoxification processes in seabirds and propose a powerful approach for deep investigation of the Hg metabolic processes in seabirds., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Mercury (Hg) is an environmental contaminant which, at high concentrations, can negatively influence avian physiology and demography. Albatrosses (Diomedeidae) have higher Hg burdens than all other avian families. Here, we measure total Hg (THg) concentrations of body feathers from adult grey-headed albatrosses ( Thalassarche chrysostoma ) at South Georgia. Specifically, we (i) analyse temporal trends at South Georgia (1989-2013) and make comparisons with other breeding populations; (ii) identify factors driving variation in THg concentrations and (iii) examine relationships with breeding success. Mean ± s.d. feather THg concentrations were 13.0 ± 8.0 µg g -1 dw, which represents a threefold increase over the past 25 years at South Georgia and is the highest recorded in the Thalassarche genus. Foraging habitat, inferred from stable isotope ratios of carbon ( δ 13 C), significantly influenced THg concentrations-feathers moulted in Antarctic waters had far lower THg concentrations than those moulted in subantarctic or subtropical waters. THg concentrations also increased with trophic level ( δ 15 N), reflecting the biomagnification process. There was limited support for the influence of sex, age and previous breeding outcome on feather THg concentrations. However, in males, Hg exposure was correlated with breeding outcome-failed birds had significantly higher feather THg concentrations than successful birds. These results provide key insights into the drivers and consequences of Hg exposure in this globally important albatross population.

The non-breeding period plays a major role in seabird survival and population dynamics. However, our understanding of the migratory behaviour, moulting and feeding strategies of non-breeding seabirds is still very limited, especially for small-sized species. The present study investigated the post-breeding behaviour of three distant populations (Kerguelen Archipelago, southeastern Australia, New Zealand) of the common diving petrel (CDP) ( Pelecanoides urinatrix ), an abundant, widely distributed zooplanktivorous seabird breeding throughout the southern Atlantic, Indian and Pacific oceans. The timing, geographical destination and activity pattern of birds were quantified through geolocator deployments during the post-breeding migration, while moult pattern of body feathers was investigated using stable isotope analysis. Despite the high energetic cost of flapping flight, all the individuals quickly travelled long distances (greater than approx. 2500 km) after the end of the breeding season, targeting oceanic frontal systems. The three populations, however, clearly diverged spatially (migration pathways and destinations), and temporally (timing and duration) in their post-breeding movements, as well as in their period of moult. Philopatry to distantly separated breeding grounds, different breeding phenologies and distinct post-breeding destinations suggest that the CDP populations have a high potential for isolation, and hence, speciation. These results contribute to improving knowledge of ecological divergence and evolution between populations, and inform the challenges of conserving migratory species., Competing Interests: We declare we have no competing interests., (© 2020 The Authors.)

Since mercury (Hg) biogeochemistry in the Southern Ocean is minimally documented, we investigated Hg stable isotopes in the blood of seabirds breeding at different latitudes in the Antarctic, Subantarctic and Subtropical zones. Hg isotopic composition was determined in adult penguins (5 species) and skua chicks (2 species) from Adélie Land (66°39'S, Antarctic) to Crozet (46°25'S, Subantarctic) and Amsterdam Island (37°47'S, Subtropical). Mass-dependent (MDF, δ 202 Hg) and mass-independent (MIF, Δ 199 Hg) Hg isotopic values separated populations geographically. Antarctic seabirds exhibited lower δ 202 Hg values (-0.02 to 0.79 ‰, min-max) than Subantarctic (0.88 to 2.12 ‰) and Subtropical (1.44 to 2.37 ‰) seabirds. In contrast, Δ 199 Hg values varied slightly from Antarctic (1.31 to 1.73 ‰) to Subtropical (1.69 to 2.04 ‰) waters. The extent of methylmercury (MeHg) photodemethylation extrapolated from Δ 199 Hg values was not significantly different between locations, implying that most of the bioaccumulated MeHg was of mesopelagic origin. The larger increase of MDF between the three latitudes co-varies with MeHg concentrations. This supports an increasing effect of specific biogenic Hg pathways from Antarctic to Subtropical waters, such as Hg biological transformations and accumulations. This "biogenic effect" among different productive southern oceanic regions can also be related to different mixed layer depth dynamics and biological productivity turnover that specifically influence the vertical transport between the mesopelagic and the photic zones. This study shows the first Hg isotopic data of the Southern Ocean at large scale and reveals how regional Southern Ocean dynamics and productivity control marine MeHg biogeochemistry and the exposure of seabirds to Hg contamination., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

In coastal ecosystems, top predators are exposed to a wide variety of nutrient and contaminant sources due to the diversity of trophic webs within inshore marine habitats. Mercury contamination could represent an additional threat to shark populations that are declining worldwide. Here we measured total mercury, carbon and nitrogen isotopes, as well as mercury isotopes, in two co-occurring shark species (the bull shark Carcharhinus leucas and the tiger shark Galeocerdo cuvier) and their potential prey from a coastal ecosystem of the western Indian Ocean (La Réunion Island). Our primary goals were to (i) determine the main trophic Hg sources for sharks and (ii) better characterize their diet composition and foraging habitat. Hg isotope signatures (Δ 199 Hg and δ 202 Hg) of shark prey suggested that bull sharks were exposed to methylmercury (MeHg) produced in offshore epipelagic waters, while tiger sharks were exposed to offshore mesopelagic MeHg with additional microbial transformation in slope sediments. Δ 199 Hg values efficiently traced the ecology of the two predators, demonstrating that bull sharks targeted coastal prey in shallow waters while tiger sharks were mainly foraging on mesopelagic species in the deeper waters of the island slope. Unexpectedly, we found a positive shift in δ 202 Hg (>1‰) between sharks and their prey, leading to high δ 202 Hg values in the two shark species (e.g. 1.91 ± 0.52‰ in bull sharks). This large shift in δ 202 Hg indicates that sharks may display strong MeHg demethylation abilities, possibly reflecting evolutionary pathways for mitigating their MeHg contamination., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Cephalopods represent an important pathway for mercury transfer through food webs. Due to the general difficulties in capturing oceanic squid, beaks found in the diet of top predators can be used to study their life-cycles and ecological role. Using upper beaks of the giant warty squid Moroteuthopsis longimana (major prey in the Southern Ocean), we describe a method to assess mercury concentrations along the life of cephalopods through the segmentary analysis of beak sections (i.e. tip of the rostrum and subsections along the hood). Distinct total mercury concentrations in the different subsections support that beaks can be used to study mercury levels in different periods of cephalopods' life-cycle. Mercury values in the anterior (1.3-7.9 μg kg -1 dw) and posterior (7.8-12.5 μg kg -1 dw) subsections of the hood reflect juvenile and adult stages, respectively. Furthermore, these results confirm that mercury bioaccumulates continuously throughout the individuals' life, with adults doubling their mercury concentrations to juveniles., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

To mediate competition, similar sympatric species are assumed to use different resources, or the same but geographically separated resources. The two giant petrels ( Macronectes spp.) are intriguing in that they are morphologically similar seabirds with overlapping diets and distributions. To better understand the mechanisms allowing their coexistence, we investigated intra- and interspecific niche segregation at Marion Island (Southern Indian Ocean), one of the few localities where they breed in sympatry. We used GPS tracks from 94 individuals and remote-sensed environmental data to quantify habitat use, combined with blood carbon and nitrogen stable isotope ratios from 90 individuals to characterize their foraging habitat and trophic ecology. Females of both species made distant at-sea foraging trips and fed at a similar trophic level. However, they used distinct pelagic habitats. By contrast, males of both species mainly foraged on or near land, resulting in significant sexual segregation, but high interspecific habitat and diet overlap. However, some males showed flexible behavioural strategies, also making distant, pelagic foraging trips. Using contemporaneous tracking, environmental and stable isotope data we provide a clear example of how sympatric sibling species can be segregated along different foraging behaviour dimensions., Competing Interests: We have no competing interests., (© 2020 The Authors.)

Changes in the foraging environment and at-sea distribution of southern elephant seals from Kerguelen Islands were investigated over a decade (2004-2018) using tracking, weaning mass, and blood δ 13 C values. Females showed either a sub-Antarctic or an Antarctic foraging strategy, and no significant shift in their at-sea distribution was detected between 2004 and 2017. The proportion of females foraging in sub-Antarctic versus Antarctic habitats did not change over the 2006-2018 period. Pup weaning mass varied according to the foraging habitat of their mothers. The weaning mass of sub-Antarctic foraging mothers' pups decreased by 11.7 kg over the study period, but they were on average 5.8 kg heavier than pups from Antarctic foraging mothers. Pup blood δ 13 C values decreased by 1.1‰ over the study period regardless of their sex and the presumed foraging habitat of their mothers. Together, these results suggest an ecological change is occurring within the Indian sector of the Southern Ocean with possible consequences on the foraging performance of southern elephant seals. We hypothesize that this shift in δ 13 C is related to a change in primary production and/or in the composition of phytoplankton communities, but this requires further multidisciplinary investigations.

The Chatham Rise, one of the highest offshore-primary production regions in New Zealand waters, hosts a great abundance and diversity of deep-sea cephalopods including the greater hooked squid, Moroteuthopsis ingens. Stable isotope ratios of carbon (δ 13 C) and nitrogen (δ 15 N) and trace element concentrations (Ag, As, Cd, Co, Cr, Cu, Fe, Hg, Ni, Pb, Se, V, and Zn) were assessed in female and male specimens of different size classes (89-563 mm mantle length). Values of δ 13 C and δ 15 N were overall higher in females and δ 13 C was further influenced by size and sex. Both muscular mantle (the largest fraction of the total body mass) and digestive gland (the known main storage organ for Ag, Cd, Cu and Zn in many cephalopods) tissues were analysed. Higher levels of Cd were observed in males than in females. A positive effect was found between size and Hg concentrations, which could be related to the ontogenetic descent of larger specimens into deeper waters, where they are exposed to higher Hg concentrations, and/or dietary shifts toward Hg-enriched prey with increasing size. This study provides trace element data for this abundant and ecologically important species, and further reveals higher trace element concentrations (especially Hg) in M. ingens from the Chatham Rise, compared to specimens from the sub-Antarctic zone., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Background: Precise and accurate retrospective geolocation of marine predators via their tissues' isotopic composition relies on quality reference maps of relevant isotopic gradients ("isoscapes"). Additionally, a good working knowledge of any discrimination factors that may offset a marine predator's isotopic composition from baseline isotopic values, as well as tissue specific retention rates, are imperative. We provide a critical assessment of inter-specific differences among marine predator-level isoscapes within the Indian Sector of the Southern Ocean., Methods: We combined fine-scale GPS tracking data and concurrent blood plasma δ 13 C and δ 15 N values of eight seabird species (three albatross, two giant petrel and three penguin species) breeding at Marion Island to produce species- and guild-specific isoscapes., Results: Overall, our study revealed latitudinal spatial gradients in both δ 13 C and δ 15 N for far-ranging seabirds (albatrosses and giant petrels) as well as inshore-offshore gradients for near-ranging seabirds (penguins). However, at the species level, latitudinal spatial gradients were not reflected in the δ 13 C and δ 15 N isoscapes of two and three, respectively, of the five far-ranging species studied. It is therefore important when possible to estimate and apply species-specific isoscapes or have a good understanding of any factors and pathways affecting marine predators' isotopic composition when estimating the foraging distribution of marine predators via their tissues' stable isotope compositions., Conclusions: Using a multi-species approach, we provide evidence of large and regional scale systematic spatial variability of δ 13 C and δ 15 N at the base of the marine food web that propagates through trophic levels and is reflected in the isotopic composition of top predators' tissues., Competing Interests: Competing interestsThe authors have no competing interests., (© The Author(s) 2020.)

Background: The timing of events in the early part of the breeding season is crucially important for successful reproduction. Long-lived animals that migrate large distances independently of each other meet at the breeding sites to re-establish their pair bonds and coordinate their breeding duties with their partners., Methods: Using miniature light-geolocation and immersion data together with blood stable isotopes, we studied the early breeding season in Thin-billed prions Pachyptila belcheri, Antarctic prions P. desolata and Blue petrels Halobaena caerulea breeding at Kerguelen Islands in the Indian Ocean. These three species exhibit differences in their winter habitat and timing of migration, moult and breeding. We hypothesised that these differences would influence their behaviour during the early breeding season., Results: In line with our hypothesis, we found clear differences not only in the timing of colony attendance, but also in the time budgets while at sea and in habitat use. Both early breeding Blue petrels and late breeding Antarctic prions spent about 8 h per day in flight and 15 h foraging. In comparison, Thin-billed prions, which breed in mid-summer, spent less time (5 h daily) in flight and more time (18 h daily) foraging, thus maximizing the time spent foraging during the longest daylight days of the year. While the ecological habitat parameters (sea temperature, wind, productivity) of Thin-billed prions and Blue petrels were relatively stable throughout the year, Antarctic prions showed clear niche switching, caused by leapfrogging between the northernmost winter distribution to the southernmost distribution during the early breeding season. Blood stable isotopes confirmed the habitat switch between the inter-breeding and early breeding periods and highlighted trophic segregation with Blue petrels feeding more on fish and Antarctic petrels more on crustaceans during the early breeding period., Conclusion: We found that the three sympatric petrel species segregated in time and space, both in the winter and the early breeding season. The interplay of timing and distribution meant that the three species show the full range of migratory strategies, from niche-tracking Blue petrels to niche-switching Antarctic prions. The latitudinal distribution resembled the leapfrogging of terrestrial avian migrant species or populations., Competing Interests: Competing interestsWe declare that we have no conflict of interest., (© The Author(s) 2020.)

There is a paucity of information on the foraging ecology, especially individual use of sea-ice features and icebergs, over the non-breeding season in many seabird species. Using geolocators and stable isotopes, we defined the movements, distribution and diet of adult Antarctic petrels Thalassoica antarctica from the largest known breeding colony, the inland Svarthamaren, Antarctica. More specifically, we examined how sea-ice concentration and free-drifting icebergs affect the distribution of Antarctic petrels. After breeding, birds moved north to the marginal ice zone (MIZ) in the Weddell sector of the Southern Ocean, following its northward extension during freeze-up in April, and they wintered there in April-August. There, the birds stayed predominantly out of the water (60-80% of the time) suggesting they use icebergs as platforms to stand on and/or to rest. Feather δ 15 N values encompassed one full trophic level, indicating that birds fed on various proportions of crustaceans and fish/squid, most likely Antarctic krill Euphausia superba and the myctophid fish Electrona antarctica and/or the squid Psychroteuthis glacialis . Birds showed strong affinity for the open waters of the northern boundary of the MIZ, an important iceberg transit area, which offers roosting opportunities and rich prey fields. The strong association of Antarctic petrels with sea-ice cycle and icebergs suggests the species can serve, year-round, as a sentinel of environmental changes for this remote region., Competing Interests: We declare we have no competing interests., (© 2020 The Authors.)

The foraging niches of seabirds are driven by a variety of factors, including competition for prey that promotes divergence in trophic niches. Bass Strait, south-eastern Australia, is a key region for seabirds, with little penguins Eudyptula minor , short-tailed shearwaters Ardenna tenuirostris , fairy prions Pachyptila turtur and common diving-petrels Pelecanoides urinatrix being particularly abundant in the region. The trophic niches of these species were investigated using isotopic values in whole blood and by identifying prey remains in stomach contents. The four species occupied different isotopic niches that varied among years, seasons and regions. Little penguins consumed mainly fish whereas the three procellariforms primarily consumed coastal krill Nyctiphanes australis . The dietary similarities between the procellariforms suggest that food resources are segregated in other ways, with interspecific differences in isotope niches possibly reflecting differential consumption of key prey, divergent foraging locations and depth, and differences in breeding phenology. Because oceanographic changes predicted to occur due to climate change may result in reduced coastal krill availability, adversely affecting these seabird predators, further information on foraging zones and feeding behaviour of small procellariform species is needed to elucidate more fully the segregation of foraging niches, the capacity of seabirds to adapt to climate change and the potential for interspecific competition in the region., Competing Interests: The authors declare there are no competing interests and Peter Dann is employed by Phillip Island Nature Parks., (©2020 Fromant et al.)

Seabirds from remote regions are mainly exposed to environmental contaminants from non-point contamination of their food webs. Pre-fledging seabird chicks are fed by their parents with marine prey captured in the vicinity of breeding colonies. Contaminant concentrations in tissues of pre-fledging chicks can thus be mostly related to local dietary sources, and have the potential to unravel spatial patterns of environmental contamination in marine ecosystems. Here, mercury (Hg), 13 other trace elements, and 18 persistent organic pollutants (POPs) were quantified in blood of chicks across four breeding locations that encompass a large latitudinal range in the southern Indian Ocean (from Antarctica, through subantarctic areas, to the subtropics), over a single breeding season. Thirteen species of penguins, albatrosses and petrels were studied, including endangered and near-threatened species, such as Amsterdam albatrosses and emperor penguins. Blood Hg burdens varied widely between species, with a factor of ~50 between the lowest and highest concentrations (mean ± SD, 0.05 ± 0.01 and 2.66 ± 0.81 µg g -1 dry weight, in thin-billed prions and Amsterdam albatrosses, respectively). Species relying on Antarctic waters for feeding had low Hg exposure. Concentrations of POPs were low in chicks, with the exception of hexachlorobenzene. Contaminant concentrations were mainly explained by species differences, but feeding habitat (inferred from δ 13 C values) and chicks' body mass also contributed to explain variation. Collectively, our findings call for further toxicological investigations in Amsterdam albatrosses and small petrel species, because they were exposed to high and diverse sources of contaminants, and in macaroni penguins, which specifically showed very high selenium concentrations. CAPSULE: Seabird chicks from four distant sites in the southern Indian Ocean had contrasted blood metallic and organic contaminant patterns depending on species, feeding habitat and body mass., (Copyright © 2019 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Climate shifts are key drivers of ecosystem change. Despite the critical importance of Antarctica and the Southern Ocean for global climate, the extent of climate-driven ecological change in this region remains controversial. In particular, the biological effects of changing sea ice conditions are poorly understood. We hypothesize that rapid postglacial reductions in sea ice drove biological shifts across multiple widespread Southern Ocean species. We test for demographic shifts driven by climate events over recent millennia by analyzing population genomic datasets spanning 3 penguin genera ( Eudyptes , Pygoscelis , and Aptenodytes ). Demographic analyses for multiple species (macaroni/royal, eastern rockhopper, Adélie, gentoo, king, and emperor) currently inhabiting southern coastlines affected by heavy sea ice conditions during the Last Glacial Maximum (LGM) yielded genetic signatures of near-simultaneous population expansions associated with postglacial warming. Populations of the ice-adapted emperor penguin are inferred to have expanded slightly earlier than those of species requiring ice-free terrain. These concerted high-latitude expansion events contrast with relatively stable or declining demographic histories inferred for 4 penguin species (northern rockhopper, western rockhopper, Fiordland crested, and Snares crested) that apparently persisted throughout the LGM in ice-free habitats. Limited genetic structure detected in all ice-affected species across the vast Southern Ocean may reflect both rapid postglacial colonization of subantarctic and Antarctic shores, in addition to recent genetic exchange among populations. Together, these analyses highlight dramatic, ecosystem-wide responses to past Southern Ocean climate change and suggest potential for further shifts as warming continues.

Rockhopper penguins are delimited as 2 species, the northern rockhopper (Eudyptes moseleyi) and the southern rockhopper (Eudyptes chrysocome), with the latter comprising 2 subspecies, the western rockhopper (Eudyptes chrysocome chrysocome) and the eastern rockhopper (Eudyptes chrysocome filholi). We conducted a phylogeographic study using multilocus data from 114 individuals sampled across 12 colonies from the entire range of the northern/southern rockhopper complex to assess potential population structure, gene flow, and species limits. Bayesian and likelihood methods with nuclear and mitochondrial DNA, including model testing and heuristic approaches, support E. moseleyi and E. chrysocome as distinct species lineages with a divergence time of 0.97 Ma. However, these analyses also indicated the presence of gene flow between these species. Among southern rockhopper subspecies, we found evidence of significant gene flow and heuristic approaches to species delimitation based on the genealogical diversity index failed to delimit them as species. The best-supported population models for the southern rockhoppers were those where E. c. chrysocome and E. c. filholi were combined into a single lineage or 2 lineages with bidirectional gene flow. Additionally, we found that E. c. filholi has the highest effective population size while E. c. chrysocome showed similar effective population size to that of the endangered E. moseleyi. We suggest that the current taxonomic definitions within rockhopper penguins be upheld and that E. chrysocome populations, all found south of the subtropical front, should be treated as a single taxon with distinct management units for E. c. chrysocome and E. c. filholi., (© The American Genetic Association 2019. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

The Southern Ocean represents a continuous stretch of circumpolar marine habitat, but the potential physical and ecological drivers of evolutionary genetic differentiation across this vast ecosystem remain unclear. We tested for genetic structure across the full circumpolar range of the white-chinned petrel (Procellaria aequinoctialis) to unravel the potential drivers of population differentiation and test alternative population differentiation hypotheses. Following range-wide comprehensive sampling, we applied genomic (genotyping-by-sequencing or GBS; 60,709 loci) and standard mitochondrial-marker approaches (cytochrome b and first domain of control region) to quantify genetic diversity within and among island populations, test for isolation by distance, and quantify the number of genetic clusters using neutral and outlier (non-neutral) loci. Our results supported the multi-region hypothesis, with a range of analyses showing clear three-region genetic population structure, split by ocean basin, within two evolutionary units. The most significant differentiation between these regions confirmed previous work distinguishing New Zealand and nominate subspecies. Although there was little evidence of structure within the island groups of the Indian or Atlantic oceans, a small set of highly-discriminatory outlier loci could assign petrels to ocean basin and potentially to island group, though the latter needs further verification. Genomic data hold the key to revealing substantial regional genetic structure within wide-ranging circumpolar species previously assumed to be panmictic., (© 2019 John Wiley & Sons Ltd.)

Background: Penguins (Sphenisciformes) are a remarkable order of flightless wing-propelled diving seabirds distributed widely across the southern hemisphere. They share a volant common ancestor with Procellariiformes close to the Cretaceous-Paleogene boundary (66 million years ago) and subsequently lost the ability to fly but enhanced their diving capabilities. With ∼20 species among 6 genera, penguins range from the tropical Galápagos Islands to the oceanic temperate forests of New Zealand, the rocky coastlines of the sub-Antarctic islands, and the sea ice around Antarctica. To inhabit such diverse and extreme environments, penguins evolved many physiological and morphological adaptations. However, they are also highly sensitive to climate change. Therefore, penguins provide an exciting target system for understanding the evolutionary processes of speciation, adaptation, and demography. Genomic data are an emerging resource for addressing questions about such processes., Results: Here we present a novel dataset of 19 high-coverage genomes that, together with 2 previously published genomes, encompass all extant penguin species. We also present a well-supported phylogeny to clarify the relationships among penguins. In contrast to recent studies, our results demonstrate that the genus Aptenodytes is basal and sister to all other extant penguin genera, providing intriguing new insights into the adaptation of penguins to Antarctica. As such, our dataset provides a novel resource for understanding the evolutionary history of penguins as a clade, as well as the fine-scale relationships of individual penguin lineages. Against this background, we introduce a major consortium of international scientists dedicated to studying these genomes. Moreover, we highlight emerging issues regarding ensuring legal and respectful indigenous consultation, particularly for genomic data originating from New Zealand Taonga species., Conclusions: We believe that our dataset and project will be important for understanding evolution, increasing cultural heritage and guiding the conservation of this iconic southern hemisphere species assemblage., (© The Author(s) 2019. Published by Oxford University Press.)

Speciation through homoploid hybridization (HHS) is considered extremely rare in animals. This is mainly because the establishment of reproductive isolation as a product of hybridization is uncommon. Additionally, many traits are underpinned by polygeny and/or incomplete dominance, where the hybrid phenotype is an additive blend of parental characteristics. Phenotypically intermediate hybrids are usually at a fitness disadvantage compared with parental species and tend to vanish through backcrossing with parental population(s). It is therefore unknown whether the additive nature of hybrid traits in itself could lead successfully to HHS. Using a multi-marker genetic data set and a meta-analysis of diet and morphology, we investigated a potential case of HHS in the prions (Pachyptila spp.), seabirds distinguished by their bills, prey choice, and timing of breeding. Using approximate Bayesian computation, we show that the medium-billed Salvin's prion (Pachyptila salvini) could be a hybrid between the narrow-billed Antarctic prion (Pachyptila desolata) and broad-billed prion (Pachyptila vittata). Remarkably, P. salvini's intermediate bill width has given it a feeding advantage with respect to the other Pachyptila species, allowing it to consume a broader range of prey, potentially increasing its fitness. Available metadata showed that P. salvini is also intermediate in breeding phenology and, with no overlap in breeding times, it is effectively reproductively isolated from either parental species through allochrony. These results provide evidence for a case of HHS in nature, and show for the first time that additivity of divergent parental traits alone can lead directly to increased hybrid fitness and reproductive isolation., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution.)