Blood Kinetics of Lipophilic and Proteinophilic Pollutants during Two Types of Long-Term Fast in King Penguins.

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.

Living in a challenging environment: Monitoring stress ecology by non-destructive methods in an Antarctic seabird.

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.

Circumpolar assessment of mercury contamination: the Adélie penguin as a bioindicator of Antarctic marine ecosystems.

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 δ13C and δ15N 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.

Mercury contamination in the tropical seabird community from Clipperton Island, eastern Pacific Ocean.

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 [δ13C] and trophic position [δ15N]) 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 δ13C values segregated boobies from frigatebirds and noddies/terns, Hg concentrations were explained by neither δ13C nor δ15N 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.

Mercury in Ten Storm-Petrel Populations from the Antarctic to the Subtropics.

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.

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.

A century of mercury: Ecosystem-wide changes drive increasing contamination of a tropical seabird species in the South Atlantic Ocean.

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 (δ1³C) and nitrogen (δ15N) 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 δ1³C and δ15N values, respectively. The sharp decline of 2.9 ‰ in δ1³C 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.

Corrigendum: The significance of cephalopod beaks as a research tool: An update.

[This corrects the article DOI: 10.3389/fphys.2022.1038064.].

The significance of cephalopod beaks as a research tool: An update.

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.

Genomic insights into the secondary aquatic transition of penguins.

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.

Blood mercury concentrations in four sympatric gull species from South Western France: Insights from stable isotopes and biologging.

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.

Stage-dependent niche segregation: insights from a multi-dimensional approach of two sympatric sibling seabirds.

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.

Reply to the comment on "New insights into the biomineralization of mercury selenide nanoparticles through stable isotope analysis in giant petrel tissues" by A. Manceau, J. Hazard. Mater. 425 (2021) 127922. doi: 10.1016/j.jhazmat.2021.127922.

In the comments reported by A. Manceau [1], relating to our recent paper on mercury (Hg) species-specific isotopic characterization in giant petrel tissues [2] two critical questions were raised. Firstly, according to A. Manceau, our method of extraction and isolation of nanoparticles was not able to efficiently isolate mercury selenide nanoparticles (HgSe NPs) and therefore the δ202Hg values measured are not species-specific, but rather δ202Hg of mixtures of complexes such as MeHgCys, Hg(Sec)4, and HgSe. Secondly, he suggests that our main findings showing that no isotopic fractionation is induced during the HgSe NPs biomineralization step from the precursor-demethylated species is erroneous because it contradicts the conclusion of two recent articles by A. Manceau and co-workers [3,4]. In this reply we defend our scientific findings and respectively respond to the questions and comments raised by A. Manceau.

First Time Identification of Selenoneine in Seabirds and Its Potential Role in Mercury Detoxification.

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.

Quantitative meta-analysis reveals no association between mercury contamination and body condition in birds.

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.

Spatial and sex differences in mercury contamination of skuas in the Southern Ocean.

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 δ15N 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.

New insights into the biomineralization of mercury selenide nanoparticles through stable isotope analysis in giant petrel tissues.

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 (δ202Hg, Δ199Hg) of HgSe NPs in seabirds and the largest characterization of this compound in biota. Similar δ202Hg values specifically associated to HgSe (δ202HgHgSe) and tissues (δ202Hgbulk) 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 δ202Hgbulk and δ202HgHgSe 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.

Independent evolution of intermediate bill widths in a seabird clade.

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.

Extreme bill dimorphism leads to different but overlapping isotopic niches and similar trophic positions in sexes of the charismatic extinct huia.

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 (δ15Nbulk) and (δ13Cbulk) 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 (δ15NAA) in male-female pairs to estimate their trophic position. Sexes had significantly different, but overlapping feather δ15Nbulk and δ13Cbulk values, but δ15NAA indicated identical trophic positions and δ15Nbulk 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.

Mercury Isotope Fractionation by Internal Demethylation and Biomineralization Reactions in Seabirds: Implications for Environmental Mercury Science.

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 δ202Hg 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 δ202Hg 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 δ202Hg (+1.6 ± 0.1‰) and mass-independent anomalies (i.e., changes in Δ199Hg 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 δ202Hg 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.