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Este artículo contiene 20 páginas, 6 figuras, 4 tablas., Aim: Over the last decades, the study of movement through tracking data has grown exceeding the expectations of movement ecologists. This has posed new challenges, specifically when using individual tracking data to infer higher- level distributions (e.g. population and species). Sources of variability such as individual site fidelity (ISF), en-vironmental stochasticity over time, and space-use variability across species ranges must be considered, and their effects identified and corrected, to produce accurate estimates of spatial distribution using tracking data. Innovation: We developed R functions to detect the effect of these sources of vari-ability in the distribution of animal groups when inferred from individual tracking data. These procedures can be adapted for their use in most tracking datasets and tracking techniques. We demonstrated our procedures with simulated datasets and showed their applicability on a real-world dataset containing 1346 year- round migratory trips from 805 individuals of three closely related seabird species breeding in 34 colonies in the Mediterranean Sea and the Atlantic Ocean, spanning 10 years. We detected an effect of ISF in one of the colonies, but no effect of the environmental stochasticity on the distribution of birds for any of the species. We also identified among-colony variability in nonbreeding space use for one species, with significant effects of popu-lation size and longitude. Main conclusions: This work provides a useful, much- needed tool for researchers using animal tracking data to model species distributions or establish conservation measures. This methodology may be applied in studies using individual tracking data to accurately infer the distribution of a population or species and support the deline-ation of important areas for conservation based on tracking data. This step, designed to precede any analysis, has become increasingly relevant with the proliferation of studies using large tracking datasets that has accompanied the globalization process in science driving collaborations and tracking data sharing initiatives., We thank the following institutions for funding: EU H2020 pro-gramme through grant 634495; Seventh Framework Programme (Research Executive Agency) through Marie Curie Career Integration Grant 618841 (FP7-PEOPLE-2013- CIG); ESFRI LifeWatch Project; LIFE programme of the European Commission through projects LIFE10 NAT/MT090 and LIFE11 NAT/IT/000093; Ministerio de Ciencia e Innovación/Ministerio de Economia y Competitividad (Spain) through projects CGL2009- 11278/BOS, CGL2013-42585-P, C G L 2 0 1 3 - 4 2 2 0 3 - R , C G L 2 0 16 - 7 8 5 3 0 - R a n d C G L 2 0 17- 8 52 10 - P ; Organismo Autónomo de Parques Nacionales (Spain) through pro-ject 1248/2014; Fundação para a Ciência e a Tecnologia (MCTES, Portugal) through projects MARE-UID/MAR/04292/2019; IF/00502/2013/CP1186/CT0003, PTDC/BIA-ANM/3743/2014, PTDC/MAR-PRO/0929/2014, UID/AMB/50017/2019 and UIDP/50017/2020 + UIDB/50017/2020 (to CESAM); Office Français de la Biodiversité (France), through the Programme PACOMM, Natura2000 en mer; Hellenic Bird Ringing Centre; MSDEC (Malta). VMP was supported by pre-doctoral contract BES-2014- 068025 of the Spanish Ministerio de Industria, Economía y Competitividad; MM by grant SFRH/BPD/47047/2008 from the Portuguese Foundation for Science and Technology; JMRG by Ph.D. grant AP2009-2163 from the Spanish Ministerio de Educación; GDO and MMü by Ornis italica and by the Regione Siciliana and Assessorato Risorse Agricole e Alimentari thoriugh a grant to the Ringing Unit of Palermo; VHP by grant SFRH/BPD/85024/2012 from the Portuguese Foundation for Science and Technology; VN by grant SFRH/BPD/88914/2012 from the Portuguese Foundation for Science and Technology; and JN by the Spanish National Programme Ramón y Cajal (RYC-2015- 17809); GK and SX were partially funded by the Operational Program “Environment and Sustainable Development” (EPPERAA) of the National Strategic Reference Framework (NSRF) 2007-2013, co- financed by the ERDF and Greek EDP; FdF by a Ph.D. grant from the Coordination for the Improvement of Higher Education Personnel (CAPES—Brazilian government agency; Bex Process 1307/13-4); ZZ by a PhD grant from the University of Barcelona (APIF/2012); MCF by a PhD grant from the University of Barcelona; and RR by post-doctoral contracts of the PLEAMAR programme from MINECO and Fundación Biodiversidad (2017/2349), and Ministerio de Ciencia, in-novación y Universidades (RYC-2017- 22055). This publication is part of the project I+D+i/PID2020-117155GB-I00, funded by MCIN/ AEI/10.13039/501100011033.

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 δ

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., This work was funded by the Fundação para a Ciência e a Tecnologia (FCT, Portugal) and the European Regional Development Fund through the projects: UIDB/04292/2020 and UIDP/04292/2020, granted to MARE; UIDP/50017/2020, UIDB/50017/2020 and LA/P/0094/2020, granted to CESAM; and PD/BD/135537/2018 awarded to F.V.

In long-lived marine top-predators with delayed sexual maturity such as seabirds, adult survival is predicted to drive population dynamics. Major knowledge gaps exist for the cryptic sub-adult stages of the population. Yet as the oceans undergo dramatic change, investigating the trends of top-predator populations and their responses to environmental variability is key for a process-based understanding of climate change. Using integrated population models based on a longitudinal data set spanning nearly 2 decades, we investigated the demographic rates and environmental drivers shaping the dynamics of a longevous marine top-predator population, the black-browed albatross Thalassarche melanophris in the Falklands. We quantified the dynamics of the whole population, including its cryptic sub-adult stages, and examined the effects of candidate environmental covariates on adult survival and productivity parameters. We found that high survival rates of both adults and juveniles are the main contributors to the growth of this albatross population, the only one that is increasing in the South Atlantic. The breeding parameters were affected by environmental fluctuations through bottom-up processes, with a negative effect of higher sea surface temperatures and a positive effect of wind intensity. The relatively constant adult survival was influenced by deeper ecosystem changes, captured by large-scale indices (Southern Annular Mode). Approximately half of the total population is composed of sub-adults, and the population dynamics are highly sensitive to changes in juvenile survival; addressing current knowledge gaps in these cryptic life-history stages is therefore a conservation goal of primary importance.

The burrowing activity of fiddler crabs inhabiting intertidal flats creates visually distinct patches within these habitats. However, differences in the composition and abundance of shorebirds and their macroinvertebrate prey between areas inhabited or not by crabs are yet to be studied. Here, we compare the macroinvertebrate and shorebird assemblages in low and high crab density areas in the intertidal flats of the Bijagos archipelago, Guinea-Bissau. High crab density areas are associated with lower richness and densities of macroinvertebrates. Shorebird assemblages were also less rich at high crab density areas and the differences in species composition occurred according to prey type preferences. Fiddler crab density was the most important variable explaining macroinvertebrate abundance, after accounting for the effects of fine fraction of sediment and distance to coast. Nonetheless, a controlled experimental setup would be required to attribute differences found to the engineering activity of fiddler crabs rather than other unaccounted habitat features. Our findings suggest that crab patches should be taken into account when assessing the distribution and abundance of macroinvertebrates and shorebirds in intertidal areas. Since low and high crab density areas differ markedly in terms of shorebird carrying capacity, monitoring variations in their extent will be important to interpret past and present population trends.

SummaryThe increasing pressure of anthropogenic development in areas with high natural value poses a huge challenge for wildlife conservation worldwide. The Tagus estuary in southern Portugal is among the most important wetlands for migratory shorebirds in the East Atlantic Flyway (EAF). However, in 2020 the Portuguese government approved the construction of the new Lisbon international airport at the heart of the Tagus estuary. Intense aircraft traffic, flying at low altitudes during both approach and take-off, is expected to cross the estuary, overlapping to a great extent the important intertidal foraging areas. Here, we aim to quantify the potential loss of conservation value of the intertidal areas of the Tagus estuary for shorebirds resulting from the disturbance (noise) caused by overflying aircraft. Using a comprehensive dataset of wintering shorebird abundance and distribution in the whole intertidal estuarine area we first identified priority areas for conservation using a spatial prioritization approach. We then performed a replacement cost analysis by excluding the areas likely to become unsuitable or severely underused by birds due to intense air traffic noise. Our results suggest that the implementation of the new Lisbon airport may lead to a loss of up to 30% of the conservation value of the Tagus estuary in terms of intertidal feeding areas of wintering birds alone. The global impact will likely be greater when effects on supratidal roosts, as well as on passage birds, are also considered. The Tagus estuary, which is internationally important for six of the 10 species included in our analysis, is just one of a network of already depleted sites along the EAF. Thus, negative impacts on bird populations on the Tagus estuary will have repercussions and undermine conservation efforts elsewhere. The plight of shorebirds at the Tagus estuary is thus a matter of international conservation concern.

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. FCT - Fundação para a Ciência e Tecnologia info:eu-repo/semantics/publishedVersion

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 (δ

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 β-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 β-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.

The conservation of migratory marine species, including pelagic seabirds, is challenging because their movements span vast distances frequently beyond national jurisdictions. Here, we aim to identify important aggregations of seabirds in the North Atlantic to inform ongoing regional conservation efforts. Using tracking, phenology, and population data, we mapped the abundance and diversity of 21 seabird species. This revealed a major hotspot associated with a discrete area of the subpolar frontal zone, used annually by 2.9–5 million seabirds from ≥56 colonies in the Atlantic: the first time this magnitude of seabird concentrations has been documented in the high seas. The hotspot is temporally stable and amenable to site-based conservation and is under consideration as a marine protected area by the OSPAR Commission. Protection could help mitigate current and future threats facing species in the area. Overall, our approach provides an exemplar data-driven pathway for future conservation efforts on the high seas. info:eu-repo/semantics/publishedVersion

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. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

The processes that drive the ontogeny of migratory strategies in long-lived animals with slow maturation remain enigmatic. While some short-lived migrants are known or believed to repeat the same migratory patterns throughout their lives, little is known on the time required for immature long-lived migrants to progressively acquire adult-like migratory behaviours, or which aspects take longer to refine during the maturation process. Here, we studied the ontogeny of long-distance migratory strategies and related patterns of spatial distribution in a long-lived seabird species during the annual cycle. To do so, we deployed light-level geolocators on 4- to 9-year-old immature Cory's shearwaters (Calonectris borealis) and on breeding adults. We revealed that migratory timings and destinations of young shearwaters progressively changed with age. The effect of ageing was remarkably evident on spring migratory performance and phenology. Birds gradually shortened the duration of the non-breeding period by advancing departure date and reducing travelling time, which resulted in a sequential arrival at the colony of the various age contingents. Ageing immatures gradually changed from a more exploratory strategy to a more conservative way of exploiting resources, reducing both their year-round spatial spread across oceanic domains and the total distance travelled. Immatures always performed a trans-equatorial migration to the Southern Hemisphere, contrasting with 17% of the adults which remained in the North Atlantic year-round. Finally, during the breeding season immatures were widely dispersed through the North Atlantic reducing their overlap with breeding adults. Our long-term study provides empirical support to the hypothesis that in long-lived species, the refinement of migratory behaviour and year-round spatial distribution is a progressive process mediated by age and experience, where life stage constraints and competition for resources may also play a role. The emerging pattern suggests that for some avian taxa, the ontogeny of migratory strategy is a prolonged, complex and dynamic process. FEDER; FCT-Fundação para a Ciência e Tecnologia info:eu-repo/semantics/publishedVersion

Marine bivalves are bioindicators of coastal environmental pollution, integrating monitoring programs worldwide. Nonetheless, the choice of particular species as an indicator requires validation, achieved by understanding the differences in element concentrations among and within species. The present study compares the chemical composition of whole body soft tissues of four common bivalve species from the Bijagos Archipelago, a pristine region of West Africa. Significant differences were recorded in the concentrations of various elements among studied species, which likely arise from species-specific uptake and bioaccumulation processes. Overall, there was a segregation between a group including the bloody cockle Senilia senilis and oyster Crassostrea tulipa (with high Cd and Zn concentrations) versus the two other species, Austromacoma nymphalis (with low Cu and high Mn, Se, Hg, Pb concentrations) and Diplodonta spp. (with high values of Cu, Al, Fe, V, Cr, Hg, Pb). C. tulipa was sampled in two different substrates (rock beds and mangrove roots), and the two groups revealed different chemical profiles, with significantly higher concentrations of P, Si, Zn and Cr and lower Cu in specimens fixed in mangrove roots. These results strongly suggest the influence of small-scale environmental variability on the accumulation of particular elements. We found extremely high Cd concentrations in S. senilis (27.1 ± 7.53 μg g−1 DW) and identified C. tulipa as another high Cd-accumulating species (ca. 10 μg g−1 DW). Our results suggest a detoxifying mechanism linked with the presence of Se to reduce the potential toxic effects of Cd in these two species. Cadmium concentrations reported for some bivalve species in this area largely exceed the maximum values proposed by the European Union, emphasizing the need for a regular contamination assessment.

Inter-individual variation in behaviour has been recognised as a major driver of population ecology, but its relationship to migratory strategy has been ill-explored. Here, we investigated whether male migrant and resident Cory’s shearwaters Calonectris borealis, a long-lived partially migratory seabird, are distinguishable by their temperament at the colony. We tracked a large number of individuals over two winters using GLS devices and assessed whether exploratory behaviour and reaction to extraction from the nest corresponded to migratory strategy over this period. While exploratory behaviour was unrelated to migratory strategy, birds that were resident were more likely to be unreactive towards extraction from the nest. This difference in behavioural stress response, together with previous findings that migrants display higher physiological stress over winter, suggests that migrants and residents may be distinguishable by their stress threshold. Fundação para a Ciência e a Tecnologia - FCT info:eu-repo/semantics/publishedVersion

Static (fixed-boundary) protected areas are key ocean conservation strategies, and marine higher predator distribution data can play a leading role toward identifying areas for conservation action. The Falkland Islands are a globally significant site for colonial breeding marine higher predators (i.e., seabirds and pinnipeds). However, overlap between marine predators and Falkland Islands proposed Marine Managed Areas (MMAs) has not been quantified. Hence, to provide information required to make informed decisions regarding the implementation of proposed MMAs, our aims were to objectively assess how the proposed MMA network overlaps with contemporary estimates of marine predator distribution. We collated tracking data (1999–2019) and used a combination of kernel density estimation and model-based predictions of spatial usage to quantify overlap between colonial breeding marine predators and proposed Falkland Islands MMAs. We also identified potential IUCN Key Biodiversity Areas (pKBAs) using (1) kernel density based methods originally designed to identify Important Bird and Biodiversity Areas (IBAs) and (2) habitat preference models. The proposed inshore MMA, which extends three nautical miles from the Falkland Islands, overlapped extensively with areas used by colonial breeding marine predators. This reflects breeding colonies being distributed throughout the Falklands archipelago, and use being high adjacent to colonies due to central-place foraging constraints. Up to 45% of pKBAs identified via kernel density estimation were located within the proposed MMAs. In particular, the proposed Jason Islands Group MMA overlapped with pKBAs for three marine predator species, suggesting it is a KBA hot spot. However, tracking data coverage was incomplete, which biased pKBAs identified using kernel density methods, to colonies tracked. Moreover, delineation of pKBA boundaries were sensitive to the choice of smoothing parameter used in kernel density estimation. Delineation based on habitat model predictions for both sampled and unsampled colonies provided less biased estimates, and revealed 72% of the Falkland Islands Conservation Zone was likely a KBA. However, it may not be practical to consider such a large area for fixed-boundary management. In the context of wide-ranging marine predators, emerging approaches such as dynamic ocean management could complement static management frameworks such as MMAs, and provide protection at relevant spatiotemporal scales. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

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., This project has received funding from the European Union’s Horizon 2020 Research and Innovation Programme under the Marie Skłodowska-Curie grant agreement no. 766417. Research made possible with the support, permissions, and funding granted by the following organizations, institutions, and grant agreements: Marine Conservation Program of DPIPWE Tasmania; American Bird Conservancy; Bureau of Ocean Energy Management; communities of Isla Mocha and Islas Juan Fernández; Corporación Nacional Forestal and Servicio Agrícola y Ganadería (Chile); Environment Canada; the National Fish and Wildlife Foundation; the National Geographic Society; ProDelphinus; and the Wallis Foundation; NSF grants DEB 9304579, DEB 9629539, DEB9806606, DEB0235818, and DEB 0842199 to D.J.A.; the National Geographic Society; the U.S. Fish and Wildlife Service; U.S. Geological Survey Ecosystems Mission Area, Wake Forest University; Max-Planck Society and State of Baden-Wuerttemberg Innovation funding; Colorado State University International Programs; Swiss Friends of Galapagos; the International Center for Tropical Ecology at University of Missouri-St. Louis; the Instituto Antartico Chileno (INACH) and the Australian Antarctic Division (AAD); LIFE “Marine IBAs IN Spain” (LIFE04NAT/ES/000049, 2004-2009) and LIFE+ INDEMARES (2009-2014); Sea World Research and Rescue Foundation Inc.; Holsworth Wildlife Research Endowment; and Winifred Violet Scott Trust; FCT-Portugal through projects (UIDB/04292/2020 and UIDP/04292/2020 and UIDP/50017/2020 and UIDB/50017/2020, granted to MARE and CESAM, respectively); the Falklands Islands Government; Natural Environmental Research Council (NERC) core funding to British Antarctic Survey Ecosystems Programme and Official Development Assistance Atlantic Islands project (NE/ R000 107/1); the New Zealand Department of Conservation; Ministry for Primary Industries; Ngāti Rehua Ngāti Wai ki Aotea; Falklands Island Conservation; University of Barcelona (APIF/2015, to M.C.-F.); the French Polar Institute (program IPEV n°109 to H.W.); Réserve Naturelle Nationale des Terres Australes Françaises; and the Zones Atelier Antarctique (LTSER France, CNRS-INEE); European funds through the European Commission Training and Mobility of Researchers Programme (ERBFMBICT983030); Spanish funds through the Ministerio de Ciencia y Tecnología (REN2002-01164/GLO), Ministerio de Educación y Ciencia (CGL2006-01315/BOS), Ministerio de Ciencia e Innovación (CGL2009-11278/BOS), and Ministerio de Economía y Competitividad (CGL2013-42585-P); Catalan funds through the Generalitat de Catalunya (2001SGR00091); and additional funding from SEO/BirdLife (programa Migra & proyecto LIFE+ Indemares), Fundación Banco Bilbao Vizcaya Argentaria (BIOCON04/099) and Fundación Biodiversidad (18PCA4328, 2012-2013); NSERC Discovery Grant and Government of Canada’s Program for International Polar Year to W.A.M.; and an ACAP AC Grant in 2013-14, predoctoral contract BES-2017-079874 of the Spanish Ministerio de Industria, Economía y Competitividad (to L.N.-H.); Spanish Foundation for Biodiversity and Spanish Ministry of Science grant ref. CGL2013-42203-R; the Pew Environment Group via the Pew Fellowship Award in Marine Conservation (to M.L.C.); National Research Foundation; South Africa and Oceans and Coasts; Department of Environment, Agriculture and Fisheries; Malta Seabird Project (LIFE10NAT/MT/090) co-funded by the LIFE program of the European Commission and the Maltese Ministry for the Environment, Sustainable Development and Climate Change, in partnership with the Royal Society for the Protection of Bird and the Portuguese Society for the Study of Birds; predoctoral contract BES-2014-068025 of the Spanish Ministerio de Industria, Economía y Competitividad (to V.M.-P.); Scientific Expert PIM initiative (Petites Iles de Méditerranée); the PIM initiative (Petites Iles de Méditerranée); the Tunisian Coastal Protection and Planning Agency (APAL); Ministry of the Environment, Japan; Funding by Fundación Ecocentro, Argentina; Wildlife Conservation Society, USA; and Ministerio de Ciencia, Tecnología e Innovación, Argentina; Centro Nacional Patagónico (CONICET), postdoctoral contracts by Beatriu de Pinós (2010-BP_A-00173), Juan de la Cierva (JCI-2009-05426), PLEAMAR (2017/2349), and Ramón y Cajal (RYC-2017-22055) programme (to R.R.); Seventh Framework Programme (Research Executive Agency of the European Commission, 618841, FP7-PEOPLE-2013-CIG); Fondation Total pour la Biodiversité (project: Trophic ecology and impacts of bycatch on the avifauna communities of Zembra archipelago); Agence de Protection et d’Aménagement du Littoral (APAL-Tunisia); Killam Postdoctoral fellowship from Dalhousie University; South African National Antarctic Programme; ACAP; Papahānaumokuākea Marine National Monument; NOAA; Japan Society for the Promotion of Science Kakenhi grant 19651100 and 15H02857; National Parks and Conservation Service (Mauritius) (to M.L.C.); IPEV Prog 109; and NASA. The use of trade, product, or firm names in this publication is for descriptive purposes only and does not imply endorsement by the U.S. government. The scientific results and conclusions, as well as any views or opinions expressed herein, are those of the authors and do not necessarily reflect those of the NOAA or the Department of Commerce. This communication reflects only the authors’ view, and the Research Executive Agency of the European Union is not responsible for any use that may be made of the information it contains. Z.Z. acknowledges funding from a predoctoral grant (APIF/2012) from the University of Barcelona. J.Ad. acknowledges funding from the U.S. Geological Survey Ecosystems Mission Area, U.S. Bureau of Ocean Energy Management, Pacific OCS Region.

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. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

In the context of environmental change, determining the causes underpinning unusual mortality events of vertebrate species is a crucial conservation goal. This is particularly true for polar and sub-polar colonial seabirds, often immunologically naïve to new and emerging diseases. Here, we investigate the patterns of black-browed albatross (Thalassarche melanophris) chick mortality events unrelated to predation recorded between the 2004/05 and 2019/2020 breeding seasons in four colonies across the species range in the Falklands. The prevalence of these mortality events was highly variable across years, causing the death of between 3 and 40% of all chicks in the studied plots. With few exceptions, mortality was patchily distributed. Using clustering methodologies, we identified the spatio-temporal mortality clusters based on the nest locations and chick death date. Using generalised linear models and generalised additive mixed-effects models we found that chicks nearer the first mortality event were predicted to die before those in more distant nests. The probability of death increased with age and was highest for chicks close to nests where a chick had died previously. Our findings, along with the symptoms consistently exhibited by most deceased chicks in the study, strongly suggest the prevalence of a widespread infectious disease, potentially with a common aetiology, both in areas with regular and with very rare human presence. Understanding the causes driving these disease-related mortality events, which seem different from the outbreaks documented in the literature, is a conservation priority for the Falklands black-browed albatross population, which comprises over 70% of the species global population. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

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. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

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. info:eu-repo/semantics/publishedVersion

Islands worldwide have sufered seabird extinctions after the arrival of humans and the alien species they introduced. On Santa Luzia (Cabo Verde), an uninhabited island of 35 km2 ,the presence of an impressive quantity of petrel bones in coastal dunes suggested the previous existence of an important seabird colony. Yet, these remains had not been identifed and no seabird extinctions have been reported for the island. This paper characterizes the extinct seabird colony of Santa Luzia and discusses the chronology and possible causes of its demise. A total of 130 grid points in a 5 km2 area and 38 supplementary points within and outside the main study area were excavated to collect bone remains. A total of 1318 anatomical elements, identifed as White-faced Storm Petrel Pelagodroma marina eadesorum (85.3% of the minimum number of individuals, MNI), Boyd’s Shearwater Pufnus lherminieri boydi (11.8% MNI) and Cape Verde Storm Petrel Hydrobates jabejabe (2.9% MNI), were found in 18% of the sampling points, within 1.25 km2 . Neither of the two former species currently breeds on Santa Luzia. In addition, two bones of the Cape Verde Shearwater Calonectris edwardsii and Pterodroma sp. were detected in the supplementary points. Radiocarbon dating of White-faced Storm Petrel (n=10) and Boyd’s Shearwater bones (n=13) suggests that the colony probably went extinct during the frst half of the twentieth century. The recent extinction of these species on Santa Luzia might be consequent on the arrival of humans and their domestic animals on the island. We estimate that the extinct seabird populations must have been at least hundreds of thousand strong, far larger than current populations of the same species in Cabo Verde. We suggest that, following feral cat eradication, Santa Luzia has a signifcant potential for seabird restoration Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

Tunas are among the most exploited top predators worldwide, with negative impacts on some of their stocks. Changes in their population abundance can impact marine food-webs and have the potential to alter entire ecosystems. To better understand the impacts of the exploitation of tuna stocks in the most critical habitats, basic knowledge on the diet of these species in each region is required. Here, we describe the diet of the two most fished tuna species in the archipelago of Madeira, the Bigeye tuna Thunnus obesus and the Skipjack tuna Katsuwonus pelamis, based on stomach contents analysis. To gain further insights into the diet, and also better assess the possible bias caused by the occurrence of live bait in stomachs, we compared tuna mercury values with those of two other predators with similar diets that are not directly targeted by fisheries, and with Bigeye and Skipjack tunas from other ocean basins. Bigeye tunas fed mostly on Atlantic chub mackerel Scomber colias and mackerel Trachurus sp., which together contributed with 85% of total prey weight. Only 7.5% of prey weight was constituted by mesopelagic prey, including myctophids and cephalopods. Skipjack tunas had an epipelagic diet (NF = 97%), with the Atlantic chub mackerel representing half of the total prey weight, despite Longspine snipefish Macroramphosus scolopax and Sand smelt Atherina sp. accounting for 62.9% of the total number of prey. There were interannual variations in diet likely linked to interannual pelagic community shifts. Bait did not bias the results of the stomach analysis of these tunas and bait species were observed to be part of the natural diet of both tuna species. Baseline data provided by this study should allow for more informed decisions for an efficient ecosystem-based fisheries management. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

The Atlantic chub mackerel Scomber colias and the Blue jack mackerel Trachurus picturatus are two abundant species in the Macaronesia region which include 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. However, little is known about their feeding ecology in oceanic environments. In this study, we describe the seasonal variation of 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. Trachurus 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 were dominated by fish. Despite consuming 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. info:eu-repo/semantics/acceptedVersion

Seabirds are a highly threatened group, yet the foraging ecology of several species remains poorly understood. Brown boobies breed in all oceans in the tropical region and are common across their range. In Tinhosa Grande (São Tomé and Príncipe), this species breeds in one of the largest colonies of seabirds in the east tropical Atlantic. We studied the foraging ecology of Brown boobies from this colony during the chick-rearing period. Thirty-three birds were tracked with GPS loggers and their diet was analysed from 11 regurgitations, using traditional and DNA barcoding techniques for prey identifcation. A total of 127 completed foraging trips were identifed, 89% of which lasted less than 24 h. Females performed signifcantly longer trips and both sexes foraged preferentially over deep oceanic waters. The diet of Brown boobies included juvenile fsh and squid (Sthenoteuthis pteropus), comprising mostly fsh species whose juvenile phases live in the pelagic environment, and only migrate to coastal waters when adults. The most frequent of those prey found was Flying gurnard (Dactylopterus volitans). The relevance of such prey shows that Brown booby conservation depends not only on the management of their foraging areas and breeding sites but also on the correct management of the coastal adjacent areas that support the adult individuals of some of their prey. Our results suggest that the areas closest to the colony do not have available resources for these birds to feed on and that Brown boobies may be associate with subsurface marine predators or with sargassum patches to forage. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

Monitoring mercury concentration in the marine environment is pivotal due to the risks that mercury intake poses to the ecosystem and human health. It is therefore of interest to make reliable, comparative measurements over large geographic areas. Here, the utility of wide-ranging generalist seabirds as mercury biomonitors at an ocean basin scale was assessed, using the Cory's Shearwater as a model species. The mercury concentration in flight feathers moulted at distant non-breeding areas of geolocator-tracked birds was quantified, reflecting contamination in various geographic areas. Compound-specific isotope analysis of amino acids was used to obtain comparable trophic positionestimates controlled for baseline isoscape. Birds that remained resident in the Canary Current integrated less mercury into their feathers than those that migrated to either the Benguela or Agulhas currents. Residents also occupied a significantly lower trophic position during the non-breeding season than migrants, largely explaining the difference in mercury exposure. Both mercury concentration and trophic position were similar in individuals spending the non-breeding period in the Benguela and Agulhas currents. This paper highlights the importance of accurate trophic position calculation in order to understand mercury exposure in wide-ranging predators and for meaningful spatial comparisons. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

Seabirds must often travel vast distances to exploit heterogeneously distributed oceanic resources, but how routes and destinations of foraging trips are optimized remains poorly understood. Among the seabirds, gadfly petrels ( Pterodroma spp.) are supremely adapted for making efficient use of wind energy in dynamic soaring flight. We used GPS tracking data to investigate the role of wind in the flight behaviour and foraging strategy of the Desertas petrel, Pterodroma deserta . We found that rather than visiting foraging hotspots, Desertas petrels maximize prey encounter by covering some of the longest distances known in any animal in a single foraging trip (up to 12 000 km) over deep, pelagic waters. Petrels flew with consistent crosswind (relative wind angle 60°), close to that which maximizes their groundspeed. By combining state–space modelling with a series of comparisons to simulated foraging trips (reshuffled-random, rotated, time-shifted, reversed), we show that this resulted in trajectories that were close to the fastest possible, given the location and time. This wind use is thus consistent both with birds using current winds to fine-tune their routes and, impressively, with an a priori knowledge of predictable regional-scale wind regimes, facilitating efficient flight over great distances before returning to the home colony.

Trace elements' concentration in the ocean is fast growing and is a source of major concern. Being charismatic and at the top of food chains, seabirds are often used as biological monitors of contaminants. We studied the concentration of trace elements in blood of black-browed albatross from the Falklands Islands, which we here show, by tracking with geolocators, forage over most of the Patagonian Shelf. Levels of trace elements were measured in males and females from two different islands. Blood concentrations of trace elements were not significantly different between islands, which is consistent with observations from foraging behavior revealing that birds from both islands foraged in broadly the same areas in the months before sampling. Arsenic and selenium concentrations in females were higher than in males. Sex-related differences in the concentration of these elements may be related to unknown slight differences in diet or to differences in assimilation between sexes. These results provide reference values for monitoring elemental contamination in the Patagonian Shelf Large Marine Ecosystem using black-browed albatrosses, one of the most abundant top predators and a suitable sentinel for the region's environmental health. Fundação para a Ciência e Tecnologia - FCT info:eu-repo/semantics/publishedVersion

1. The identification of geographic areas where the densities of animals are highest across their annual cycles is a crucial step in conservation planning. In marine environments, however, it can be particularly difficult to map the distribution of species, and the methods used are usually biased towards adults, neglecting the distribution of other life-history stages even though they can represent a substantial proportion of the total population. 2. Here we develop a methodological framework for estimating populationlevel density distributions of seabirds, integrating tracking data across the main life-history stages (adult breeders and non-breeders, juveniles and immatures). We incorporate demographic information (adult and juvenile/immature survival, breeding frequency and success, age at first breeding) and phenological data (average timing of breeding and migration) to weight distribution maps according to the proportion of the population represented by each life-history stage. 3. We demonstrate the utility of this framework by applying it to 22 species of albatrosses and petrels that are of conservation concern due to interactions with fisheries. Because juveniles, immatures and non-breeding adults account for 47%–81% of all individuals of the populations analysed, ignoring the distributions of birds in these stages leads to biased estimates of overlap with threats, and may misdirect management and conservation efforts. Population-level distribution maps using only adult distributions underestimated exposure to longline fishing effort by 18%–42%, compared with overlap scores based on data from all lifehistory stages. 4. Synthesis and applications. Our framework synthesizes and improves on previous approaches to estimate seabird densities at sea, is applicable for data-poor situations, and provides a standard and repeatable method that can be easily updated as new tracking and demographic data become available. We provide scripts in the R language and a Shiny app to facilitate future applications of our approach. We recommend that where sufficient tracking data are available, this framework be used to assess overlap of seabirds with at-sea threats such as overharvesting, fisheries bycatch, shipping, offshore industry and pollutants. Based on such an analysis, conservation interventions could be directed towards areas where they have the greatest impact on populations. Fundação para a Ciência e a Tecnologia - FCT; French Polar Institute IPEV; Falkland Islands Government info:eu-repo/semantics/publishedVersion

Biological invasions can have serious consequences for native communities. The Asian date mussel Arcuatula senhousia is an Asian species that has been accidentally introduced into coastal areas of ...

This study was funded by the Department for Environment, Food and Rural Affairs UK as part of the Darwin Plus project DPLUS027 (Marine Spatial Planning for the Falkland Islands). JFM received financial support during this study from the German Research Foundation (DFG SPP 1158, MA2574/6–1). JPG received financial support during this study from FCT - Portugal through the strategic project UID/AMB/ 50017/2013 and from FEDER funds granted to CESAM, within the PT2020 Partnership Agreement and Compete 2020. Marine Spatial Planning (MSP) is becoming a key management approach throughout the world. The process includes the mapping of how humans and wildlife use the marine environment to inform the development of management measures. An integrated multi-species approach to identifying key areas is important for MSP because it allows managers a global representation of an area, enabling them to see where management can have the most impact for biodiversity protection. However, multi-species analysis remains challenging. This paper presents a methodological framework for mapping key areas for marine megafauna (seabirds, pinnipeds, cetaceans) by incorporating different data types across multiple species. The framework includes analyses of tracking data and observation survey data, applying analytical steps according to the type of data available during each year quarter for each species. It produces core-use area layers at the species level, then combines these layers to create megafauna core-use area layers. The framework was applied in the Falkland Islands. The study gathered over 750,000 tracking and at-sea observation locations covering an equivalent of 5495 data days between 1998 and 2015 for 36 species. The framework provides a step-by-step implementation protocol, replicable across geographic scales and transferable to multiple taxa. R scripts are provided. Common repositories, such as the Birdlife International Tracking Database, are invaluable tools, providing a secure platform for storing and accessing spatial data to apply the methodological framework. This provides managers with data necessary to enhance MSP efforts and marine conservation worldwide. Postprint

Sixty sediment samples from four sites in the Bijagos archipelago were characterized for fine fraction, loss on ignition, major, minor and trace elemental composition (Al, Fe, Ca, Mg, Ti, P, Zr, Mn, Cr, Sr, Ba, B, V, Li, Zn, Ni, Pb, As, Co, U, Cu, Cs and Cd), and the elements of the La–Lu series. Element concentrations were largely explained by the Al content and the proportion of fine fraction content, with the exception of Ca and Sr. Sediments showed enhanced Ti, U, Cr, As and Cd concentrations with respect to estimated upper crust values, most likely mirroring a regional signature. Rare earth elements were in deficit relatively to the North American Shale Composite (NASC), mainly in coarser material. No pronounced Ce-anomaly was observed, while Eu-anomalies were positive in most analyzed sediments.

Knowledge of the macrozoobenthic community and its interactions with predators is critical for understanding intertidal ecosystems but this information is still lacking for many tropical sites. We core-sampled three intertidal areas in the Bijagos archipelago, Guinea-Bissau, one of the largest intertidal wetlands in West Africa, to evaluate macroinvertebrate abundance and diversity, investigate ecological variables influencing macroinvertebrate distribution, and estimate food availability for several shorebird species. We found a low macroinvertebrate biomass, which mostly consisted of bivalves and polychaetes. The fiddler crab Afruca tangeri was also among the more abundant species. Both Shannon diversity and rarefied species richness indicate macrozoobenthic communities in the Bijagos have low diversity but there is large variation in species composition among sites. Sediment fine fraction, organic content and distance to the coast were correlated with the abundance of several macroinvertebrate taxa. Harvestable biomass for the shorebird species studied was very low, below the values described for intertidal sites in temperate areas and some other sites in the tropics. Although this fits with low shorebird densities in the Bijagos, it also suggests that wintering shorebird communities must depend on secondary production by macrobenthic invertebrates during the winter, in the same way as has been proposed for other tropical intertidal systems.