Oceanic seabirds chase tropical cyclones.

In late summer and autumn, the passage of intense tropical cyclones can profoundly perturb oceanic and coastal ecosystems. Direct negative effects on individuals and marine communities can be dramatic, especially in the coastal zone,1,2,3,4 but cyclones can also enhance pelagic primary and secondary production.5,6,7,8,9 However, cyclone impacts on open ocean marine life remain poorly understood. Here, we investigate their effects on the foraging movements of a wide-ranging higher predator, the Desertas petrel (Pterodroma deserta), in the mid-latitude North Atlantic during hurricane season. Contrary to previously studied pelagic seabirds in tropical and mid-latitude regions,10,11 Desertas petrels did not avoid cyclones by altering course, nor did they seek calmer conditions within the cyclone eye. Approximately one-third of petrels tracked from their breeding colony interacted with approaching cyclones. Upon encountering strong winds, the birds reduced ground speed, likely by spending less time in flight. A quarter of birds followed cyclone wakes for days and over thousands of kilometers, a behavior documented here for the first time. Within these wakes, tailwind support was higher than along alternative routes. Furthermore, at the mesoscale (hours-weeks and hundreds of kilometers), sea surface temperature dropped and surface chlorophyll sharply increased, suggesting direct effects on ocean stratification, primary production, and therefore presumably prey abundance and accessibility for surface-feeding petrels. We therefore hypothesize that cyclone wakes provide both predictably favorable wind conditions and foraging opportunities. As such, cyclones may have positive net effects on the demography of many mid-latitude pelagic seabirds and, likely, other marine top-predators.

Allochrony is shaped by foraging niche segregation rather than adaptation to the windscape in long-ranging seabirds.

BACKGROUND: Ecological segregation allows populations to reduce competition and coexist in sympatry. Using as model organisms two closely related gadfly petrels endemic to the Madeira archipelago and breeding with a two month allochrony, we investigated how movement and foraging preferences shape ecological segregation in sympatric species. We tested the hypothesis that the breeding allochrony is underpinned by foraging niche segregation. Additionally, we investigated whether our data supported the hypothesis that allochrony is driven by species-specific adaptations to different windscapes. METHODS: We present contemporaneous tracking and stable isotopes datasets for Zino's (Pterodroma madeira) and Desertas (Pterodroma deserta) petrels. We quantified the year-round distribution of the petrels, characterised their isotopic niches and quantified their habitat preferences using machine learning (boosted regression trees). Hidden-Markov-models were used to investigate the effect of wind on the central-place movement speed, and a simulation framework was developed to investigate whether each species breeds at times when the windscape is most favourable to sustain their trips. RESULTS: Despite substantial spatial overlap throughout the year, the petrels exhibited diverging isotopic niches and habitat preferences during breeding. Both species used a vast pelagic region in the North Atlantic, but targeted two different mesopelagic ecoregions and showed a preference for habitats mostly differing in sea surface temperature values. Based on our simulation framework, we found that both species would perform trips of similar speed during the other species' breeding season. CONCLUSIONS: The different breeding schedules between the species are underpinned by differences in foraging habitat preferences and adaptation to the local environment, rather than to the windscape. Nevertheless, the larger Desertas petrels exploited significantly windier conditions, potentially unsustainable for the smaller Zino's petrels. Furthermore, due to larger mass and likely higher fasting endurance, Desertas petrels engaged in central-place-foraging movements that covered more ground and lasted longer than those of Zino's petrels. Ultimately, patterns of ecological segregation in sympatric seabirds are shaped by a complex interplay between foraging and movement ecology, where morphology, foraging trip regulation and fasting endurance have an important- yet poorly understood- role.

Food web of the oceanic region of the archipelago of Madeira: The role of marine megafauna in the subtropical northeast Atlantic ecosystem.

Many oceanic areas are still in need of baseline information on their structure and functioning. This is particularly important due to the ever-increasing impacts of global changes, which have led to the decline of marine life, and top predators in particular. The study of the structure and functioning of food webs can help understand the consequences of the disappearance of this group in marine ecosystems. Here, we develop a mass-balanced model for the marine Exclusive Economic Zone of the archipelago of Madeira, with emphasis on the role of marine megafauna in this ecosystem. A total of 50 functional groups were defined, representing coastal and open ocean areas, and epipelagic and deep-sea levels. The total biomass of the Madeira system was calculated at 52.68 t km-2, with lower trophic level organisms comprising 89.9 % of its biomass. Marine megafauna, namely pelagic sharks and coastal birds had the highest impacts across other trophic levels and were classified as keystone species, together with monk seals. The food web was characterized by a linear-like food chain, with a large proportion of specialized organisms, including dolphins, shearwaters, and large pelagic fish. The low mean trophic level of the system was 2.03, much lower than that of fisheries (4.3) targeting mainly tunas and Black scabbardfish. Considering the importance of marine megafauna in this food web and the threats they are facing; monitoring studies of key species in the region should be a priority. This study can now be used to build a needed ecosystem-based fisheries management and integrate conservation measures to declining species.

Diet of the critically endangered blackchin guitarfish Glaucostegus cemiculus revealed using DNA metabarcoding.

We present the first assessment of the diet of the blackchin guitarfish Glaucostegus cemiculus (Geoffroy Saint-Hilaire, 1817) for West Africa using DNA metabarcoding on stomach contents of individuals captured in the Bijagós Archipelago, Guinea-Bissau. The diet was dominated by crustaceans, particularly caramote prawn Penaeus kerathurus (frequency of occurrence [FO] = 74%, numerical frequency [NF] = 54%) and fiddler crab Afruca tangeri (FO = 74%, NF = 12%). Bony fishes were present in 30% of the stomachs. We highlight the importance of conservation action for intertidal habitats and their associated benthic invertebrates for the survival of the critically endangered blackchin guitarfish.

Trace elements in relation to trophic ecology of long-distance migratory shorebirds and seabirds in West Africa.

Shorebirds and seabirds are abundant predators in coastal habitats worldwide, relying upon a high diversity of benthic invertebrates and fish, respectively. While occupying different trophic guilds, they are differently exposed to element contamination entering the coastal food web. Therefore, these birds have been used as bioindicators of environmental contamination in marine ecosystems. We analysed the concentration of trace elements in blood samples of 16 shorebird and seabird migratory species in a major non-breeding site, the Bijagós Archipelago, in regard to their trophic ecology. Overall, our study shows low exposure of this bird community to toxic elements, except for Hg. Most species presented Hg burdens within the moderate toxicity threshold, but one species (Dunlin) presented values at a potential high Hg toxicity risk. We found a positive relationship between Se and Ni concentrations and δ15N values (a proxy for trophic level). In addition, a positive relationship was found between δ13C (a proxy for habitat characteristics) and Hg, Fe, Cu and Mn, while the opposite pattern was recorded for As. Differences were also shown for several trace elements between the two studied trophic guilds: concentrations of As, Pb and Se were higher in pelagic fish consumers (seabirds), whereas Cr, Fe and Sr burdens were higher in benthic invertebrate consumers (shorebirds). Although previous studies in the same site revealed very high concentrations of Cd and Pb in some of the prey species of shorebirds and seabirds (bivalves and fishes), values found in birds for these two elements suggest no toxicological risk. Thus, exposure to toxic elements is not currently a threat to coastal bird populations, namely those facing apparent local declines in Guinea-Bissau, one of the most important non-breeding quarters for of West Africa. Still, Hg burdens were high in some species, deserving further monitoring.

Spatial and seasonal variation in macrozoobenthic density, biomass and community composition in a major tropical intertidal area, the Bijagós Archipelago, West-Africa.

The coastal intertidal ecosystem of the Bijagós Archipelago, Guinea-Bissau, one of the largest and most important in West Africa, sustains a considerable proportion of the migratory shorebird populations of the East Atlantic Flyway and operates as a nursery area for benthic fish in the region. The macrozoobenthos in these mudflats constitute the main food source for both groups so that spatial and temporal variation in their abundance and community composition is likely to influence the abundance and distribution of fish and birds. In this study we described the spatial and temporal dynamics in the density, biomass, and community composition of macrozoobenthos across six intertidal flats in three islands of the Bijagós Archipelago. Overall, the Bijagós Archipelago was characterised by a highly species-rich macrozoobenthic community, with ca. 88 taxa identified across all sites, reaching a mean density of 1871 ± 58.3 ind.m-2 (mean ± SE) and mean biomass of 5.65 ± 0.41 g of AFDM.m-2 (ash-free dry mass per m2), values much lower than what was described for nearby intertidal areas, namely the Band d'Arguin, Mauritania. Density and biomass of major macrozoobenthos classes (Bivalvia, Polychaeta, Malacostraca and Gastropoda) differed across sites and months, displaying an overall increase in density towards the final months of the dry season (March and April). Similarly, community composition also differed significantly between sites and throughout the season. The site with most distinct community composition (Adonga) supported low diversity and high abundance of a few bivalve species, whilst other two sites that hosted the most diverse communities, were also the most similar between them (Anrumai and Abu). These spatial and temporal patterns constitute an important baseline to improve knowledge of this intertidal ecosystem and will contribute towards a better understanding of the spatial and temporal distribution patterns of their consumers.

A central place foraging seabird flies at right angles to the wind to jointly optimize locomotor and olfactory search efficiency.

To increase the probability of detecting odour plumes, and so increase prey capture success, when winds are stable central place foraging seabirds should fly crosswind to maximize the round-trip distance covered. At present, however, there is no empirical evidence of this theoretical prediction. Here, using an extensive GPS tracking dataset, we investigate, for the first time, the foraging movements of Bulwer's petrels (Bulweria bulwerii) in the persistent North Atlantic trade winds. To test the hypotheses that, in stable winds, petrels use crosswind to maximize both the distance covered and the probability of detecting olfactory cues, we combine state-space models, generalized additive models and Gaussian plume models. Bulwer's petrels had the highest degree of selectivity for crosswinds documented to date, often leading to systematic zig-zag flights. Crosswinds maximized both the distance travelled and the probability of detecting odour plumes integrated across the round-trip (rather than at any given point along the route, which would result in energetically costly return flight). This evidence suggests that petrels plan round-trip flights at departure, integrating expected costs of homeward journeys. Our findings, which are probably true for other seabirds in similar settings, further highlight the critical role of wind in seabird foraging ecology.

Higher mercury contamination is associated with shorter telomeres in a long-lived seabird - A direct effect or a consequence of among-individual variation in phenotypic quality?

Mercury is a heavy metal, which is pervasive and persistent in the marine environment. It bioaccumulates within organisms and biomagnifies in the marine food chain. Due to its high toxicity, mercury contamination is a major concern for wildlife and human health. Telomere length is a biomarker of aging and health, because it predicts survival, making it a potential tool to investigate sublethal effects of mercury contamination. However, the relationship between telomeres and mercury contamination is unclear. We measured feather mercury concentration in Cory's Shearwaters Calonectris borealis, long-lived seabirds and top predators, between 9 and 35 years of age and related it to telomere length in erythrocytes. Cory's Shearwaters with higher mercury concentrations had shorter telomeres and the effect was sex-dependent, reaching significance in males only. This may be explained by the fact that males have longer telomeres and higher and more variable mercury concentrations than females in this population. The mercury effect on telomere length was stronger on longer telomeres in the genome within individuals. We discuss the hypotheses that the negative correlation could either be a direct effect of mercury on telomere shortening and/or a consequence of variation in phenotypic quality among individuals that results in a covariation between mercury contamination and telomere length.

Contrasting patterns from two invasion fronts suggest a niche shift of an invasive predator of native bees.

Background: The accuracy of predictions of invasive species ranges is dependent on niche similarity between invasive and native populations and on our ability to identify the niche characteristics. With this work we aimed to compare the niche dynamics of two genetically related invasive populations of Vespa velutina (an effective predator of honeybees and wild pollinators), in two distinct climatic regions, one in central Europe and another one in the north-western Iberian Peninsula, and hence to identify uninvaded regions susceptible to invasion. Methods: Niche dynamics and shifts of V. velutina were assessed by comparing the environmental niches of the native and of the two invasive populations, using climatic, topographic and land use variables. We also ran reciprocal distribution models using different algorithms and records from both native and invasive ranges to compare model predictions and estimate which regions are at a greater risk of being invaded. Results: An apparent niche shift was detected in the population of the NW of Iberian Peninsula, where the species is living under environmental conditions different from the native niche. In central Europe, large suitable areas remain unoccupied. The fact that both invasive populations are well established, despite occupying environmentally distinct regions indicates that V. velutina has a high ability to successfully invade different environmental envelopes from those existing in its native range. For example, in north-western Iberian Peninsula the species is now thriving out of its native niche limits. Moreover, the large extent of still unoccupied environmental space with similar conditions to those used by the species in its native range suggests that there is still a large area of central and eastern Europe that can be potentially invaded by the species.

Linking trophic ecology with element concentrations in a coastal fish community of the Bijagós Archipelago, West Africa.

We report the concentration of 13 elements in the muscle and liver of 17 coastal fish species of the Bijagós Archipelago, Guinea-Bissau, and link element concentrations to trophic ecology as assessed by carbon (δ13C) and nitrogen (δ15N) stable isotopes. We found higher concentrations of Hg, Fe, Mn, Cu, Zn, Se and As in liver as compared to muscle tissue, and the opposite pattern for Sr and Ca in all fish species. The concentration of Hg and Se in muscle samples increased significantly with δ15N, suggesting a biomagnification of these elements in this food chain. The concentrations of Ca and Sr, Fe and Cr, Ca and Mn, and Fe and Mn were positively correlated to each other in more than 50% of the studied species. Fish constitute the most important animal protein source for people in Guinea-Bissau, and thus assessing the concentrations of potentially toxic elements is relevant for human health.

Stopover use of a large estuarine wetland by dunlins during spring and autumn migrations: Linking local refuelling conditions to migratory strategies.

Migratory strategies dictate stopover ecology, particularly concerning decisions of when, where and how long to stop, and what to do at stationary periods. In birds, individuals stop primarily to replenish energy stores, although the functions of stopover events vary among and within species, particularly between pre- and post-breeding seasons. Here, we combined plasma metabolite levels and haematological parameters to compare refuelling rates and physiological state within (early, mid, late) and between (spring, autumn) migratory periods, aiming to identify potentially different migratory strategies in a shorebird, the dunlin Calidris alpina, using a key stopover site in Iberia. Plasma triglycerides and ß-hydroxybutyrate concentrations did not differ between seasons, and small differences were found in haematological profiles (higher haemoglobin and hematocrit levels in spring). Similar refuelling rates and physiological status suggests a single migratory strategy in spring and autumn. During both seasons, dunlins arrive at the Tagus estuary with medium-to-high fuel loads, indicating they do not engage in prolonged fuelling. This agrees with a skipping migratory strategy, where birds fly short-to-medium distances while fuelling at moderate rates along a network of sites. Although we may expect late spring migrants to experience stronger pressures to optimally schedule migratory events, we found no significant differences in physiological profiles among early, mid and late migrants. Unexpectedly, such differences were found in autumn: early birds showed the highest triglycerides and haemoglobin levels and lowest ß-hydroxybutyrate concentrations. These results denote enhanced refuelling rates and blood oxygen-carrying capacity in early autumn migrants, which is typical of jumpers, i.e., birds travelling with larger fuel loads and performing fewer stops. Our study adds substantially to previous knowledge of stopover ecology in migratory shorebirds in the East Atlantic Flyway. Importantly, it indicates that the Tagus estuary is a high-quality stopover site for intermediate fuelling. Yet, understanding non-fuelling stopping functions is needed to ultimately inform conservation planning.

Environmental variability directly affects the prevalence of divorce in monogamous albatrosses.

In many socially monogamous species, divorce is a strategy used to correct for sub-optimal partnerships and is informed by measures of previous breeding performance. The environment affects the productivity and survival of populations, thus indirectly affecting divorce via changes in demographic rates. However, whether environmental fluctuations directly modulate the prevalence of divorce in a population remains poorly understood. Here, using a longitudinal dataset on the long-lived black-browed albatross (Thalassarche melanophris) as a model organism, we test the hypothesis that environmental variability directly affects divorce. We found that divorce rate varied across years (1% to 8%). Individuals were more likely to divorce after breeding failures. However, regardless of previous breeding performance, the probability of divorce was directly affected by the environment, increasing in years with warm sea surface temperature anomalies (SSTA). Furthermore, our state-space models show that warm SSTA increased the probability of switching mates in females in successful relationships. For the first time, to our knowledge, we document the disruptive effects of challenging environmental conditions on the breeding processes of a monogamous population, potentially mediated by higher reproductive costs, changes in phenology and physiological stress. Environmentally driven divorce may therefore represent an overlooked consequence of global change.

Diet and trophic position of two mackerel species in the archipelago of Madeira, Portugal.

The Atlantic chub mackerel Scomber colias and the blue jack mackerel Trachurus picturatus are two abundant species in the Macaronesia region which includes the archipelago of Madeira, Portugal. Both are key species in the trophic web, being important prey for several local top predators, such as seabirds and marine mammals. Nonetheless, little is known about their feeding ecology in oceanic environments. In this study, the authors describe the seasonal variation in the diet of S. colias and T. picturatus in the oceanic region of Madeira throughout a year. Visual inspection of stomach contents revealed that S. colias fed on a broader range of prey groups than T. picturatus, but for both species, zooplankton (particularly calanoid copepods) and fish were the most important food items. The diet of S. colias included a higher proportion of fish, namely Atlantic saury Scomberesox saurus and S. colias, than that of T. picturatus, that included mostly the longspine snipefish Macroramphosus scolopax. T. picturatus consumed a higher proportion of decapods and other copepods. Seasonal variation was found in the diet of both species, with zooplanktonic species being more important in colder months (February to April) for S. colias and during warm months (May to October) for T. picturatus. Their diet in other seasons was dominated by fish. Although they consume similar prey, carbon and nitrogen stable isotope analysis of muscle of S. colias and T. picturatus showed little overlap in their diets, and T. picturatus showed higher δ15 N and a narrower isotopic niche.

Assessing the contribution of mangrove carbon and of other basal sources to intertidal flats adjacent to one of the largest West African mangrove forests.

Mangrove forests are productive habitats and major potential exporters of organic matter and nutrients to adjacent habitats. Here we examine the extent to which mangrove carbon is transferred to adjacent intertidal food webs in the second largest mangrove-covered area in Africa, in Guinea-Bissau. Applying stable isotope analysis and mixing models, we made comparisons at two spatial scales: (1) a large scale, comparing intertidal flats with (mangrove sites) and without (control sites) adjacent mangrove forests regarding the carbon isotopic signature of macrozoobenthos and sediment organic matter (SOM), and the relative importance of potential primary food sources in sustaining macrozoobenthos, and (2) a fine scale, performing stable carbon isotope measurements along 200 m transects from the coastline out to open intertidal flats, to trace mangrove carbon in macrozoobenthos and in the SOM. We found no evidence that mangrove carbon sustains intertidal food webs, despite SOM being significantly more depleted in 13C in mangrove sites. Mangrove leaves had the lowest relative contribution to the diet of macrozoobenthos, while macroalgae, benthic microalgae and POM showed variable but overall relevant contributions. Yet, at a smaller scale, mangrove carbon was detectable in SOM and in most macrozoobenthos, being strongest within 50 m of the mangrove edge and quickly fading with increasing distance. Our results suggest that there is only a marginal input of mangrove carbon into the food webs of unvegetated intertidal flats. Still, this leaves open the possibility of mangrove forests acting as sources of dissolved inorganic carbon and processed nitrogen, which can be assimilated by the algae and subsequently fuel adjacent food webs.

Global political responsibility for the conservation of albatrosses and large petrels.

Migratory marine species cross political borders and enter the high seas, where the lack of an effective global management framework for biodiversity leaves them vulnerable to threats. Here, we combine 10,108 tracks from 5775 individual birds at 87 sites with data on breeding population sizes to estimate the relative year-round importance of national jurisdictions and high seas areas for 39 species of albatrosses and large petrels. Populations from every country made extensive use of the high seas, indicating the stake each country has in the management of biodiversity in international waters. We quantified the links among national populations of these threatened seabirds and the regional fisheries management organizations (RFMOs) which regulate fishing in the high seas. This work makes explicit the relative responsibilities that each country and RFMO has for the management of shared biodiversity, providing invaluable information for the conservation and management of migratory species in the marine realm.

Monitoring of mercury in the mesopelagic domain of the Pacific and Atlantic oceans using body feathers of Bulwer's petrel as a bioindicator.

Global mercury pollution has markedly and consistently grown over the past 70 years (although with regional variations in trends) and is a source of major concern. Mercury contamination is particularly prevalent in biota of the mesopelagic layers of the open ocean, but these realms are little studied, and we lack a large scale picture of contamination in living organisms of this region. The Bulwer's petrel Bulweria bulwerii, a species of migratory seabird, is a highly specialised predator of mesopelagic fish and squid, and therefore can be used as a bioindicator for the mesopelagic domain. Mercury accumulated by the birds through diet is excreted into feathers during the moulting process in adults and feather growth in chicks, reflecting contamination in the non-breeding and breeding periods, respectively, and hence the influence of different, largely non-overlapping breeding and non-breeding ranges. We studied mercury in feathers and the trophic position in two colonies from the Atlantic Ocean (Portugal and Cape Verde) and two colonies from the Pacific Ocean (Japan and Hawaii). We found significantly lower levels of mercury in adult and chick samples from the Pacific Ocean compared with samples from the Atlantic Ocean. However, we did not detect differences in trophic position of chicks among colonies and oceans, suggesting that differences in mercury measured in feathers reflect levels of environmental contamination, rather than differences in the structure of the trophic chain in different oceans. We conclude that despite a reduction in mercury levels in the Atlantic in recent decades, mesopelagic organisms in this ocean remain more heavily contaminated than in the Pacific at tropical and subtropical latitudes. We suggest that Bulwer's petrel is a highly suitable species to monitor the global contamination of mercury in the mesopelagic domain.

Costs of reproduction and migration are paid in later return to the colony, not in physical condition, in a long-lived seabird.

Life history theory suggests a trade-off between costly activities such as breeding and migration and somatic self-maintenance. However, how the short-term cost of parental effort is expressed in species with a slow pace-of-life is not well understood. Also, investigating carry-over effects of migration is most meaningful when comparing migratory strategies within the same population, but this has rarely been done. We explore this hypothesis in a long-lived, pelagic seabird, the Cory's Shearwater, Calonectris borealis, where males display partial migration. By manipulating reproductive effort and taking advantage of the natural variation in migratory strategy, we investigate whether early reproductive failure and migratory strategy had implications on the physical condition of males on return to the colony the following year. We experimentally induced breeding failure from mid-incubation, tracked the over-winter movements of these males and of males that invested in parental effort, and assessed innate immunity, stress, and residual body mass the following year. Early breeding failure resulted in earlier return to the colony among all males, associated with greater probability of reproductive success. Residents had a lower tail feather fault bar intensity, an indicator of stress during the non-breeding period, compared to migrants. Reproductive effort and migratory strategy had no impact on physiological condition otherwise. Our results provide evidence that in species with a slow-pace of life, such as the Cory's Shearwater, somatic maintenance is prioritised, with the costs of reproduction and migration paid in delayed arrival date.

Seabird diet analysis suggests sudden shift in the pelagic communities of the subtropical Northeast Atlantic.

The dynamics of the subtropical pelagic ecosystems of the Northeast Atlantic are still poorly known due to the high costs associated with sampling large oceanic areas. Top predators can be used as alternative low-cost samplers and indicators of the temporal variability of such systems. To study the variation in the composition of pelagic species through time in the broad Canary current region, we analysed foraging trips and regurgitations of Cory's shearwaters Calonectris borealis nesting on Selvagens islands, in 2008-2011 and 2016-2018. Fisheries data, oceanographic variables and the North Atlantic Oscillation were explored as possible explanatory variables for trends in behaviour and diet. Cory's shearwaters' diet, complemented by fisheries data, revealed marked changes in the composition of the pelagic fish communities. In 2016 there was a peak in the abundance of the Atlantic chub mackerel Scomber colias, followed by an explosive increase in the abundance of the Longspine snipefish Macroramphosus scolopax in 2017 and 2018, as deduced from the diet composition of the Cory's shearwater, and supported by fisheries data, in the broad oceanic area surrounding the Selvagens islands. Oceanographic variables did not show fluctuations correlated with these marked shifts in pelagic fish availability, the causes of which remain largely unknown. This study highlights the importance of the Atlantic chub mackerel and of the Longspine snipefish in the Madeira/Canary region and exemplifies the efficiency of avian predators in revealing rapid changes in pelagic communities of oceanic domains. Such trends and variations need to be better monitored and understood to measure the impact of ongoing global changes and to sustainably manage the marine environment and resources.

Untangling causes of variation in mercury concentration between flight feathers.

Bird feathers are one of the most widely used animal tissue in mercury biomonitoring, owing to the ease of collection and storage. They are also the principal excretory pathway of mercury in birds. However, limitations in our understanding of the physiology of mercury deposition in feathers has placed doubt on the interpretation of feather mercury concentratoins. Throughout the literature, moult sequence and the depletion of the body mercury pool have been taken to explain patterns such as the decrease in feather mercury from the innermost (P1) to the outermost primary feather (P10) of the wing. However, it has been suggested that this pattern is rather a measurement artefact as a result of the increased feather mass to length ratio along the primaries, resulting in a dilution effect in heavier feathers. Here, we attempt to untangle the causes of variation in feather mercury concentrations by quantifying the mercury concentration as µg of mercury (i) per gram of feather, (ii) per millimetre of feather, and (iii) per day of feather growth in the primary feathers of Bulwer's Petrel Bulweria bulwerii chicks, effectively controlling for some of the axes of variation that may be acting in adults, and monitoring the growth rate of primary feathers in chicks. The mercury concentration in Bulwer's Petrel chicks' primaries increased from the innermost to the outermost primary for all three concentration measures, following the order of feather emergence. These observations confirm that the pattern of mercury concentration across primary feathers is not an artefact of the measure of concentration, but is likely an effect of the order of feather growth, whereby the earlier grown feathers are exposed to higher blood mercury concentrations than are later moulted feathers as a result of blood mercury depletion.