Quantifying variation in δ13C and δ15N isotopes within and between feathers and individuals : is one sample enough?

This study represents a contribution to the ecosystems component of the British Antarctic Survey Polar Science for Planet Earth Programme, funded by The Natural Environment Research Council and through a NERC standard grant NE/I02237X/1.Studies of avian migration increasingly use stable isotope analysis to provide vital trophic and spatial markers. However, when interpreting differences in stable isotope values of feathers, many studies are forced to make assumptions about the timing of moult. A fundamental question remains about the consistency of these values within and between feathers from the same individual. In this study, we examine variation in carbon and nitrogen isotopes by sub-sampling feathers collected from the wings of adults of two small congeneric petrel species, the broad-billed Pachyptila vittata and Antarctic prion P. desolata. Broad-billed prion feather vane material was enriched in 15N compared to feather rachis material, but there was no detectable difference in δ13C. Comparison of multiple samples taken from Antarctic prion feathers indicated subtle difference in isotopes; rachis material was enriched in 13C compared to vane material, and there were differences along the length of the feather, with samples from the middle and tip of the feather depleted in 15N compared to those from the base. While the greatest proportion of model variance was explained by differences between feathers and individuals, the magnitude of these within-feather differences was up to 0.5 ‰ in δ15N and 0.8 ‰ in δ13C. We discuss the potential drivers of these differences, linking isotopic variation to individual-level dietary differences, movement patterns and temporal dietary shifts. A novel result is that within-feather differences in δ13C may be attributed to differences in keratin structure within feathers, suggesting further work is required to understand the role of different amino acids. Our results highlight the importance of multiple sampling regimes that consider both within- and between-feather variation in studies using stable isotopes.Publisher PDFPeer reviewe

Marine protected areas show low overlap with projected distributions of seabird populations in Britain and Ireland

EJC was funded by the Irish Petroleum Infrastructure Programme (PIP) IS013/08. WJG was funded by INSITE http://www.insitenorthsea.org/. AK was funded by the Irish Research CouncilGOIPD/2015/81. MJ was funded by the SFI Centre for Marine and Renewable Energy (12/RC/2302).Marine Protected Areas (MPAs) are an important tool for the conservation of seabirds. However, mapping seabird distributions using at-sea surveys or tracking data to inform the designation of MPAs is costly and time-consuming, particularly for far-ranging pelagic species. Here we explore the potential for using predictive distribution models to examine the effectiveness of current MPAs for the conservation of seabirds, using Britain and Ireland as a case study. A distance-weighted foraging radius approach was used to project distributions at sea for an entire seabird community during the breeding season, identifying hotspots of highest density and species richness. The percentage overlap between distributions at sea and MPAs was calculated at the level of individual species, family group, foraging range group (coastal or pelagic foragers), and conservation status. On average, 32.5% of coastal populations and 13.2% of pelagic populations overlapped with MPAs indicating that pelagic species, many of which are threatened, are likely to have significantly less coverage from protected areas. We suggest that a foraging radius approach provides a pragmatic and rapid method of assessing overlap with MPA networks for central place foragers. It can also act as an initial tool to identify important areas for potential designation. This would be particularly useful for regions throughout the world with limited data on seabird distributions at sea and limited resources to collect this data. Future assessment for marine conservation management should account for the disparity between coastal and pelagic foraging species to ensure that wider-ranging seabirds are afforded adequate levels of protection.PostprintPeer reviewe

Environmental drivers of population-level variation in the migratory and diving ontogeny of an Arctic top predator

This work is an output of the ARISE project (NE/P006035/1 and NE/P00623X/1), part of the Changing Arctic Ocean programme jointly funded by the UKRI Natural Environment Research Council (NERC) and the German Federal Ministry of Education and Research (BMBF). Fieldwork in Canada was carried out under a Canadian Council on Animal Care permit no. NAFC2017–2 and funded by Fisheries and Oceans Canada and a bursary from Department for Business, Energy and Industrial Strategy (BEIS) administered by the NERC Arctic Office. Fieldwork in the Greenland Sea was approved by the Greenland Ministry of Fisheries, Hunting and Agriculture and the Norwegian Food Safety Authority (permit no. 11546) as part of the Northeast Greenland Environmental Study Program 2017–2018 (by the Danish Centre for Environment and Energy at Aarhus University, The Greenland Institute of Natural Resources and the Environmental Agency for Mineral Resource Activities of the Government of Greenland) and financed by oil licence holders in the area.The development of migratory strategies that enable juveniles to survive to sexual maturity is critical for species that exploit seasonal niches. For animals that forage via breath-hold diving, this requires a combination of both physiological and foraging skill development. Here, we assess how migratory and dive behaviour develop over the first year of life for a migratory Arctic top predator, the harp seal Pagophilus groenlandicus, tracked using animal-borne satellite relay data loggers. We reveal similarities in migratory movements and differences in diving behaviour between 38 juveniles tracked from the Greenland Sea and Northwest Atlantic breeding populations. In both regions, periods of resident and transitory behaviour during migration were associated with proxies for food availability: sea ice concentration and bathymetric depth. However, while ontogenetic development of dive behaviour was similar for both populations of juveniles over the first 25 days, after this time Greenland Sea animals performed shorter and shallower dives and were more closely associated with sea ice than Northwest Atlantic animals. Together, these results highlight the role of both intrinsic and extrinsic factors in shaping early life behaviour. Variation in the environmental conditions experienced during early life may shape how different populations respond to the rapid changes occurring in the Arctic ocean ecosystem.Publisher PDFPeer reviewe

Sympatric seals, satellite tracking and protected areas : habitat-based distribution estimates for conservation and management

Analysis was funded by the UK Government Department for Business, Energy and Industrial Strategy (BEIS; OESEA-16-76/OESEA-17-78) with support from the Natural Environment Research Council (NERC; INSITE Phase II NE/T010614/1 EcoSTAR), EU INTERREG (MarPAMM), and the Scottish Government (MMSS/002/15). DJFR’s contribution was funded by NERC National Capability Funding (NE/R015007/1). WJG was supported by INSITE Phase I (MAPS). Telemetry tags and their deployment were funded in the UK by BEIS (and previous incarnations), NERC, Marine Scotland, Scottish Government, NatureScot, SMRU, SMRU Instrumentation Group, Marine Current Turbines, Ørsted, the Met Office, the Zoological Society of London (ZSL), the Crown Estate, Highlands & Islands Enterprise, Moray Firth Renewables Limited (MORL), Beatrice Offshore Windfarm Limited (BOWL), SITA Trust, BBC Wildlife Fund and the Hampshire & Isle of Wight Wildlife Trust. Tags and their deployment in Ireland were funded by Inland Fisheries Ireland, the Department of Communications, Marine and Natural Resources, the Higher Education Authority of Ireland, the National Geographic Society, the Department of Agriculture, Food and the Marine, and the National Parks and Wildlife Service. UK aerial surveys conducted by SMRU were funded by NERC (NE/R015007/1), NatureScot, the Department for Agriculture, Environment and Rural Affairs (Northern Ireland), Marine Current Turbines, Marine Scotland, Natural England, and Scottish Power. Aerial surveys in Ireland were funded by the Department for Tourism, Culture, Arts, Gaeltacht, Sport and Media.Marine predator populations are crucial to the structure and functioning of ecosystems. Like many predator taxa, pinnipeds face an increasingly complex array of natural and anthropogenic threats. Understanding the relationship between at-sea processes and trends in abundance at land-based monitoring sites requires robust estimates of at-sea distribution, often on multi-region scales. Such an understanding is critical for effective conservation management, but estimates are often limited in spatial extent by spatial coverage of animal-borne tracking data. Grey (Halichoerus grypus) and harbour seals (Phoca vitulina) are sympatric predators in North Atlantic shelf seas. The United Kingdom (UK) and Ireland represents an important population centre for both species, and Special Areas of Conservation (SACs) are designated for their monitoring and protection. Here we use an extensive high-resolution GPS tracking dataset, unprecedented in both size (114 grey and 239 harbour seals) and spatial coverage, to model habitat preference and generate at-sea distribution estimates for the entire UK and Ireland populations of both species. We found regional differences in environmental drivers of distribution for both species which likely relate to regional variation in diet and population trends. Moreover, we provide SAC-specific estimates of at-sea distribution for use in marine spatial planning, demonstrating that hotspots of at-sea density in UK and Ireland-wide maps cannot always be apportioned to the nearest SAC. We show that for grey seals, colonial capital breeders, there is a mismatch between SACs (where impacts are likely to be detected) and areas where impacts are most likely to occur (at sea). We highlight an urgent need for further research to elucidate the links between at-sea distribution during the foraging season and population trends observed in SACs. More generally, we highlight that the potential for such a disconnect needs to be considered when designating and managing protected sites, particularly for species that aggregate to breed and exhibit partial migration (e.g. grey seals), or spatial variation in migration strategies. We demonstrate the use of strategic tracking efforts to predict distribution across multiple regions, but caution that such efforts should be mindful of the potential for differences in species-environment relationships despite similar accessible habitats.Publisher PDFPeer reviewe

Resolving issues with environmental impact assessment of marine renewable energy installations

Growing concerns about climate change and energy security have fueled a rapid increase in the development of marine renewable energy installations (MREIs). The potential ecological consequences of increased use of these devices emphasizes the need for high quality environmental impact assessment (EIA). We demonstrate that these processes are hampered severely, primarily because ambiguities in the legislation and lack of clear implementation guidance are such that they do not ensure robust assessment of the significance of impacts and cumulative effects. We highlight why the regulatory framework leads to conceptual ambiguities and propose changes which, for the most part, do not require major adjustments to standard practice. We emphasize the importance of determining the degree of confidence in impacts to permit the likelihood as well as magnitude of impacts to be quantified and propose ways in which assessment of population-level impacts could be incorporated into the EIA process. Overall, however, we argue that, instead of trying to ascertain which particular developments are responsible for tipping an already heavily degraded marine environment into an undesirable state, emphasis should be placed on better strategic assessment.Publisher PDFPeer reviewe

Global assessment of marine plastic exposure risk for oceanic birds

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

Global assessment of marine plastic exposure risk for oceanic birds

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

Inter-colony movements, at-sea behaviour and foraging in an immature seabird: results from GPS-PPT tracking, radio-tracking and stable isotope analysis

Seabird populations contain large numbers of immatures––in some instances comprising >50% of the fully grown adults in the population. These birds are significant components of marine food webs and may contribute to compensatory recruitment and dispersal, but remain severely understudied. Here, we use GPS-PTTs, radio-tracking and analysis of stable carbon (δ13C) and nitrogen (δ15N) isotopes to investigate the movements and foraging ecology of immature seabirds. Our study focussed on immature northern gannets Morus bassanus aged 2–4 attending non-breeding aggregations alongside a large breeding colony. GPS-PTT tracking of five birds revealed that immatures have the ability to disperse widely during the breeding season, with some individuals potentially prospecting at other colonies. Overall, however, immatures were faithful to the colony of capture. During returns to the focal colony, immatures acted as central place foragers, conducted looping and commuting flights, and analysis of the variance in first-passage time revealed evidence of area-restricted search (ARS) behaviour. In addition, stable carbon (δ13C) and nitrogen (δ15N) isotope analyses indicate that immatures were isotopically segregated from breeders. Our findings provide insights into the foraging, prospecting and dispersal behaviour of immature seabirds, which may have important implications for understanding seabird ecology and co

Individual seabirds show consistent foraging strategies in response to predictable fisheries discards

Current fishing extraction methods often generate huge quantities of dead or dying biomass that is returned to the sea in the form of discards. This practice produces a readily available clumped resource for many scavengers such as seabirds, but in the face of declining stocks and via policy change, the amount of discards produced is set to decline in the future. To understand how discards have influenced seabird foraging in the past and how birds may respond to future change requires studies examining consistent individual foraging choices. There is increasing evidence that populations may be made up of generalist or specialist foragers and this is key to the population's ability to adapt to change. Here we test for consistent individual foraging behaviour of northern gannets Morus bassanus in relation to fishing vessels and examine consequences of scavenging behaviour in terms of foraging effort and body condition. Using a combination of bird-borne bio-logging devices (GPS and Time Depth Recorders) with high resolution GPS data acquired through vessel monitoring systems on fishing boats, we examined the overlap between birds and fisheries. We found that during repeat foraging trips in the same breeding season, gannets regularly foraged at fishing boats but there were also clear among individual differences in the extent of fisheries overlap. Furthermore, we show for the first time that these differences represent consistent strategies – individual differences in scavenging were highly repeatable across multiple trips within a period of several weeks. However, despite this finding, we found no differences in foraging effort or body condition between scavengers and non-scavengers. Moreover, scavenging strategy did not appear to influence diving behaviour or vary by sex. Scavenging on discards appears to be a strategy employed consistently by a subsection of the population and future work should examine whether these specialisations persist throughout and between years and what causes these individual differences, exploring possible demographic and fitness consequences in light of global changes to fish stocks and fisheries management

Ecological segregation of two superabundant, morphologically similar, sister seabird taxa breeding in sympatry

Prions Pachyptila are the most abundant seabirds in the Southern Ocean and comprise two main groups: those with and without bill lamellae to filter zooplankton. With few exceptions, each breeding location supports at most one species from each of these groups. However, Gough Island supports two morphologically very similar, filter-feeding species: broad-billed P. vittata and MacGillivray’s prions P. macgillivrayi. To understand how these two species co-occur in sympatry, we compared the foraging ranges, habitat selectivity, trophic segregation and moult schedules of these species using combined geolocation-immersion loggers. After breeding, both species showed a well-defined westward migration prior to moulting. Moult lasted 11–19 weeks and was significantly longer in MacGillivray’s than broad-billed prions. Moulting birds occurred in specific areas within the Argentine Basin, with little overlap between the two species. Habitat analysis revealed species-specific preferences, in particular sea surface temperature. Activity patterns also differed; MacGillivray’s prions spent more time in flight, which indicates a more active foraging strategy, relying less on filter feeding. Stable isotope ratios (δ15N) in flight feathers were greater in MacGillivray’s prion, which is consistent with its less specialized bill morphology resulting in feeding at a higher trophic level. Inter-specific spatial segregation was observed for most of the tracking period, in large part because broad-billed prions breed roughly 3 months earlier than MacGillivray’s prions. At Tristan da Cunha, 250 km farther north, where only broad-billed prions breed, they departed, moulted and returned significantly later (15–17 days) than conspecifics from Gough Island, providing evidence for character displacement in sympatry with MacGillivray’s prion