Proof of concept evidence that stable isotopes of carbon, nitrogen and sulphur may identify individual kittiwakes breeding in different colonies.

RATIONALE: Carbon, nitrogen and sulphur stable isotopes in feathers grown by seabirds while breeding reflect the local isoscape and diet in the vicinity of the colony, so may make it possible to discriminate individual birds from different colonies. METHODS: Black-legged kittiwake Rissa tridactyla inner primary feathers from two colonies about 350 km apart in the North Sea were used to test whether δ13C, δ15N and δ34S differed between individuals from the two colonies. Feather tips cut from breeding birds caught at nests were compared with tips of moulted feathers (grown 1 year earlier) found on the ground. RESULTS: Isotopic compositions showed no overlap between the two colonies in δ13C, δ15N or δ34S in tips of newly-grown feathers sampled from breeding adult kittiwakes. There was some overlap in δ13C, δ15N and δ34S from moulted feathers, but discriminant analysis allowed >90% of individuals to be assigned to their colony. In five of six comparisons, mean isotopic compositions were the same in new and moulted feathers but not for δ34S at one of the two colonies. CONCLUSIONS: This study has demonstrated for the first time that stable isotopes in inner primary feathers of kittiwakes can allow accurate identification of the breeding colony of individual birds from two different colonies within the North Sea. Further research is required to determine if this method can be applied with greater spatial resolution and to a larger number of colonies.

HPAIV outbreak triggers short-term colony connectivity in a seabird metapopulation.

Disease outbreaks can drastically disturb the environment of surviving animals, but the behavioural, ecological, and epidemiological consequences of disease-driven disturbance are poorly understood. Here, we show that an outbreak of High Pathogenicity Avian Influenza Virus (HPAIV) coincided with unprecedented short-term behavioural changes in Northern gannets (Morus bassanus). Breeding gannets show characteristically strong fidelity to their nest sites and foraging areas (2015-2019; n = 120), but during the 2022 HPAIV outbreak, GPS-tagged gannets instigated long-distance movements beyond well-documented previous ranges and the first ever recorded visits of GPS-tagged adults to other gannet breeding colonies. Our findings suggest that the HPAIV outbreak triggered changes in space use patterns of exposed individuals that amplified the epidemiological connectivity among colonies and may generate super-spreader events that accelerate disease transmission across the metapopulation. Such self-propagating transmission from and towards high density animal aggregations may explain the unexpectedly rapid pan-European spread of HPAIV in the gannet.

Environmental Samples Test Negative for Avian Influenza Virus H5N1 Four Months after Mass Mortality at A Seabird Colony.

High pathogenicity avian influenza (HPAI) profoundly impacted several seabird populations during the summers of 2021 and 2022. Infection spread rapidly across colonies, causing unprecedented mortality. At Foula, Shetland, 1500 breeding adult great skuas Stercorarius skua, totalling about two tonnes of decomposing virus-laden material, died at the colony in May-July 2022. Carcasses were left where they died as Government policy was not to remove dead birds. The factors influencing risk of further spread of infection are uncertain, but evidence suggests that HPAI can persist in water for many months in cool conditions and may be a major transmission factor for birds living in wetlands. We investigated risk of further spread of infection from water samples collected from under 45 decomposing carcasses and in three freshwater lochs/streams by sampling water in October 2022, by which time the great skua carcasses had rotted to bones, skin, and feathers. No viral genetic material was detected four months after the mortality, suggesting a low risk of seabird infection from the local environment when the seabirds would return the next breeding season. These findings, although based on a relatively small number of water samples, suggest that the high rainfall typical at Shetland probably washed away the virus from the decomposing carcasses. However, limitations to our study need to be taken on board in the design of environmental monitoring at seabird colonies during and immediately after future outbreaks of HPAI.

Risks to different populations and age classes of gannets from impacts of offshore wind farms in the southern North Sea.

The southern North Sea holds the world's highest concentration of offshore wind farms (OWFs). Northern gannets (Morus bassanus), a species considered at high risk from OWF impacts, show a strong seasonal peak there in November, but it is unclear which populations and age classes are most at risk of collision with wind turbines. We tagged adult and juvenile gannets at the world's largest colony (Bass Rock) and reviewed two sources of survey data for different age classes to study their movements through southern North Sea waters. Tracked birds showed peak numbers in the southern North Sea in mid-October, with much smaller numbers there during November. Adults were distributed throughout the area, including waters close to OWFs, whereas juveniles were confined to the coast. Survey data indicated high proportions of immature gannets in southern North Sea waters, suggesting higher collision risk than for adults. Gannets present in November may be predominantly from colonies further north than Bass Rock.

Untangling local and remote influences in two major petrel habitats in the oligotrophic Southern Ocean.

Ocean circulation connects geographically distinct ecosystems across a wide range of spatial and temporal scales via exchanges of physical and biogeochemical properties. Remote oceanographic processes can be especially important for ecosystems in the Southern Ocean, where the Antarctic Circumpolar Current transports properties across ocean basins through both advection and mixing. Recent tracking studies have indicated the existence of two large-scale, open ocean habitats in the Southern Ocean used by grey petrels (Procellaria cinerea) from two populations (i.e., Kerguelen and Antipodes islands) during their nonbreeding season for extended periods during austral summer (i.e., October to February). In this work, we use a novel combination of large-scale oceanographic observations, surface drifter data, satellite-derived primary productivity, numerical adjoint sensitivity experiments, and output from a biogeochemical state estimate to examine local and remote influences on these grey petrel habitats. Our aim is to understand the oceanographic features that control these isolated foraging areas and to evaluate their ecological value as oligotrophic open ocean habitats. We estimate the minimum local primary productivity required to support these populations to be much <1% of the estimated local primary productivity. The region in the southeast Indian Ocean used by the birds from Kerguelen is connected by circulation to the productive Kerguelen shelf. In contrast, the region in the south-central Pacific Ocean used by seabirds from the Antipodes is relatively isolated suggesting it is more influenced by local factors or the cumulative effects of many seasonal cycles. This work exemplifies the potential use of predator distributions and oceanographic data to highlight areas of the open ocean that may be more dynamic and productive than previously thought. Our results highlight the need to consider advective connections between ecosystems in the Southern Ocean and to re-evaluate the ecological relevance of oligotrophic Southern Ocean regions from a conservation perspective.

Effects of Sitka spruce masting on phenology and demography of siskins Spinus spinus.

Masting behaviour of Sitka spruce Picea sitchensis may influence Eurasian siskin Spinus spinus breeding ecology as breeding siskins specialize on spruce seeds. We caught siskins and other small passerines over 16 years using mist nets adjacent to large plantations of mature Sitka spruce. We sexed, aged, measured and weighed the birds and collected feather samples from fledglings to measure nitrogen and carbon stable isotope ratios. Siskins departed in late summer, and returned, and bred earlier in years of higher cone abundance. Nitrogen and carbon isotopes indicated that siskins fed their chicks on Sitka spruce seeds in most years, and more so in years of high cone production. More siskins were caught following heavy rainfall, when the cones had closed, encouraging the birds to seek alternative food sources. Fledglings were not heavier or larger in years with higher cone crops but were more numerous. However, the age ratio of siskins caught the following year was unaffected by cone crop. Given their reliance on Sitka spruce seeds, climate change may have a major impact on siskin numbers by altering the availability of Sitka spruce seeds, either through changes in masting patterns or cone opening, or due to climate-related changes in forestry practices. Siskins represent a valuable study system to conservation ecology, where a native species is ecologically reliant on introduced taxa.

Behavioural plasticity in the early breeding season of pelagic seabirds - a case study of thin-billed prions from two oceans.

BACKGROUND: In long-lived seabirds that migrate large distances independently of each other, the early part of the breeding season is crucially important for a successful reproductive attempt. During this phase, pair bonds are re-established and partners coordinate their breeding duties. We studied the early breeding season in Thin-billed prions Pachyptila belcheri breeding in the Atlantic Ocean (Falkland/Malvinas Islands) and Indian Ocean (Kerguelen). Despite overlap in the wintering areas, these two populations exhibit differences in their timing and direction of migration. We hypothesised that these differences would influence behaviour during the early breeding season. RESULTS: In line with our hypothesis, we found very strong differences in colony attendance patterns. Thin-billed prions of the Falkland population spent the late winter period over shelf waters close to the colony, first arrived back at the colony in September, and attended the nests interruptedly for one month, before departing on a pre-laying exodus. In contrast, Kerguelen birds remained in the non-breeding areas until mid-October and spent much less time attending the burrow before their pre-laying exodus. Despite this asynchronous arrival to the two colonies, the subsequent patterns resulted in remarkably synchronous incubation in both populations, with males taking on the first long incubation shift in late November, whereas females returned to sea soon after egg laying. During the pre-laying exodus and incubation, Thin-billed prions from the Falklands spread north over the Patagonian Shelf, while prions from Kerguelen travelled much further, reaching southern oceanic waters and moved at faster speeds (> 400 km per day). Although prions from Kerguelen moved much further, their isotopic niches were considerably narrower, suggesting a stronger dependence on Antarctic waters. CONCLUSIONS: The study thus suggests that Thin-billed prions show a high intraspecific plasticity in their use of either neritic or oceanic waters during the early breeding season. Breeding birds from the Falkland Islands can exploit an extensive shelf area, while Kerguelen birds have adapted to the need to forage in distant southern open waters. This difference in foraging ecology may thus have shaped the phenology of the early breeding phase.

The effect of long-range transport, trophic position and diet specialization on legacy contaminant occurrence in great skuas, Stercorarius skua, breeding across the Northeast Atlantic.

High levels of halogenated organic contaminants (HOCs) have been found in the marine predatory seabird great skua (Stercorarius skua) from breeding colonies in the Northeastern Atlantic, with large unexplained inter-colony variation. The present study aimed at analyzing if the HOCs occurrence in breeding great skuas in remote colonies was explained by local baseline food web exposure determined by long-range transport, or by ecological factors such as diet specialization and relative trophic position in the breeding area. The occurrence of organochlorine pesticides (OCPs), polychlorinated biphenyls (PCBs), and polybrominated diphenyl ethers (PBDEs) was analyzed in plasma of 204 adult great skuas collected over two years (2008 and 2009) and 5 colonies across the North-Atlantic from Shetland to Svalbard. The ΣHOCs levels in plasma ranged across two orders of magnitude, from 40 to 7600 ng/g (wet weight) and differed significantly across the great skua colonies. The variation in contaminant occurrence among colonies did not reflect long-range transport through a latitudinal or remoteness gradient, as the second northernmost colony (Bjørnøya), had the highest contaminant concentrations. No latitudinal or remoteness gradient was evident in the contaminant pattern among the colonies. The contaminant levels increased significantly with increasing δ15N values, and regurgitated pellets of undigested prey suggested that great skuas with higher δ15N values had a higher proportion of bird prey in their diet, mostly seabirds. In contrast, great skuas from colonies with lower δ15N and lower contaminant level fed mostly on fish. The enrichment of δ13C increased with decreasing δ15N and lower contaminant levels. Therefore, individual behavior of great skuas, such as migration strategies and diet specialization, rather than long-range transport and thus baseline food web exposure, explain among and within colony variance in contaminant occurrence.

Ixodes ricinus parasitism of birds increases at higher winter temperatures.

Increasing winter temperatures are expected to cause seasonal activity of Ixodes ricinus ticks to extend further into the winter. We caught birds during winter months (November to February) at a site in the west of Scotland over a period of 24 years (1993-1994 to 2016-2017) to quantify numbers of attached I. ricinus and to relate these to monthly mean temperature. No adult ticks were found on any of the 21,731 bird captures, but 946 larvae and nymphs were found, with ticks present in all winter months, on 16 different species of bird hosts. All ticks identified to species were I. ricinus. I. ricinus are now active throughout the year in this area providing temperature permits. No I. ricinus were present in seven out of eight months when the mean temperature was below 3.5º C. Numbers of I. ricinus attached to birds increased rapidly with mean monthly temperatures above 7º C. Winter temperatures in Scotland have been above the long-term average in most years in the last two decades, and this is likely to increase risk of tick-borne disease.

Black guillemot ecology in relation to tidal stream energy generation: An evaluation of current knowledge and information gaps.

The black guillemot Cepphus grylle has been identified as a species likely to interact with marine renewable energy devices, specifically tidal turbines, with the potential to experience negative impacts. This likelihood is primarily based on the species being a diving seabird, and an inshore, benthic forager often associating with tidal streams. These behavioural properties may bring them into contact with turbine blades, or make them susceptible to alterations to tidal current speed, and/or changes in benthic habitat structure. We examine the knowledge currently available to assess the potential impacts of tidal stream turbines on black guillemot ecology, highlight knowledge gaps and make recommendations for future research. The key ecological aspects investigated include: foraging movements, diving behaviour, seasonal distribution, other sources of disturbance and colony recovery. Relating to foraging behaviour, between studies there is heterogeneity in black guillemot habitat use in relation to season, tide, diurnal cycles, and bathymetry. Currently, there is also little knowledge regarding the benthic habitats associated with foraging. With respect to diving behaviour, there is currently no available research regarding how black guillemots orientate and manoeuvre within the water column. Black guillemots are considered to be a non-migratory species, however little is known about their winter foraging range and habitat. The effect of human disturbance on breeding habitat and the metapopulation responses to potential mortalities are unknown. It is clear further understanding of black guillemot foraging habitat and behaviour is needed to provide renewable energy developers with the knowledge to sustainably locate tidal turbines and mitigate their impacts.

Seabird diversity hotspot linked to ocean productivity in the Canary Current Large Marine Ecosystem.

Upwelling regions are highly productive habitats targeted by wide-ranging marine predators and industrial fisheries. In this study, we track the migratory movements of eight seabird species from across the Atlantic; quantify overlap with the Canary Current Large Marine Ecosystem (CCLME) and determine the habitat characteristics that drive this association. Our results indicate the CCLME is a biodiversity hotspot for migratory seabirds; all tracked species and more than 70% of individuals used this upwelling region. Relative species richness peaked in areas where sea surface temperature averaged between 15 and 20°C, and correlated positively with chlorophyll a, revealing the optimum conditions driving bottom-up trophic effects for seabirds. Marine vertebrates are not confined by international boundaries, making conservation challenging. However, by linking diversity to ocean productivity, our research reveals the significance of the CCLME for seabird populations from across the Atlantic, making it a priority for conservation action.

Legacy Pollutants are Declining in Great Skuas (Stercorarius skua) but Remain Higher in Faroe Islands than in Scotland.

To monitor environmental pollutants in Faroese biota, samples from a top predator were analysed and put into a spatial and temporal context. Analysis of 20 Great Skua eggs sampled in 2012 from the Faroe Islands showed >70 % lower concentrations of legacy persistent organic pollutants (POPs) than in samples analysed in 1977. The 2012 Faroese eggs showed higher concentrations than for eggs in Shetland from about the same period (2008). Eggshells were analysed for sub-lethal effects but there were no detectable effects of legacy POP levels on eggshell colour or thickness. A temporal decline in legacy POPs would indicate a reduction in the general pollutant levels present in the environment as has been shown in other areas of the North Atlantic, but there are significant geographic differences in POPs levels likely due to differences in diet resulting in significantly different exposures on a relatively limited spatial scale.

Archiving Primary Data: Solutions for Long-Term Studies.

The recent trend for journals to require open access to primary data included in publications has been embraced by many biologists, but has caused apprehension amongst researchers engaged in long-term ecological and evolutionary studies. A worldwide survey of 73 principal investigators (Pls) with long-term studies revealed positive attitudes towards sharing data with the agreement or involvement of the PI, and 93% of PIs have historically shared data. Only 8% were in favor of uncontrolled, open access to primary data while 63% expressed serious concern. We present here their viewpoint on an issue that can have non-trivial scientific consequences. We discuss potential costs of public data archiving and provide possible solutions to meet the needs of journals and researchers.

Multiple Stressors in a Top Predator Seabird: Potential Ecological Consequences of Environmental Contaminants, Population Health and Breeding Conditions.

Environmental contaminants may have impacts on reproduction and survival in wildlife populations suffering from multiple stressors. This study examined whether adverse effects of persistent organic pollutants (POPs) increased with poor population health and breeding conditions in three colonies (60-74°N) of great skua (Stercorarius skua) in the north-eastern Atlantic (Shetland, Iceland and Bjørnøya [Bear Island]). POPs (organochlorines [OCs] and polybrominated diphenyl ethers [BDEs]) were measured in plasma of incubating birds (n = 222), concentrations differing nearly tenfold among colonies: Bjørnøya (2009) > Bjørnøya (2010) > Iceland (2009) > Shetland (2009). Reproductive success (hatching success and chick survival) showed that breeding conditions were favourable in Shetland and at Bjørnøya (2010), but were very poor in Iceland and at Bjørnøya (2009). Biomarkers indicated that health was poor in the Shetland population compared to the other populations. Females whose chicks hatched late had high POP concentrations in all colonies except at Bjørnøya (2010), and females losing their eggs at Bjørnøya (2009) tended to have higher concentrations than those hatching. Moreover, there was a negative relationship between female POP concentrations and chick body condition at hatching in Iceland and at Bjørnøya (2010). Supplementary feeding experiments were conducted, and in Iceland where feeding conditions were poor, significant negative relationships were found between female POP concentrations and daily growth-rate in first-hatched chicks of control nests, but not in food supplemented nests. This suggests that negative impacts of POPs were mitigated by improved feeding conditions. For second-chicks, there was a strong negative relationship between the female POP concentrations and growth-rate, but no effects of supplementary feeding. Lowered adult return-rate between breeding seasons with increasing POP loads were found both at Bjørnøya (2009) and in Shetland, especially related to BDEs. This indicates stronger fitness consequences of POPs following seasons with very poor breeding conditions and/or high reproductive effort. This study suggests that the impacts of POPs may differ depending on population health and breeding conditions, and that even low concentrations of POPs could have ecological consequences during adverse circumstances. This is important with regard to risk assessment of biomagnifying contaminants in marine ecosystems.

Half a world apart? Overlap in nonbreeding distributions of Atlantic and Indian Ocean thin-billed prions.

Distant populations of animals may share their non-breeding grounds or migrate to distinct areas, and this may have important consequences for population differentiation and dynamics. Small burrow-nesting seabirds provide a suitable case study, as they are often restricted to safe breeding sites on islands, resulting in a patchy breeding distribution. For example, Thin-billed prions Pachyptila belcheri have two major breeding colonies more than 8,000 km apart, on the Falkland Islands in the south-western Atlantic and in the Kerguelen Archipelago in the Indian Ocean. We used geolocators and stable isotopes to compare at-sea movements and trophic levels of these two populations during their non-breeding season, and applied ecological niche models to compare environmental conditions in the habitat. Over three winters, birds breeding in the Atlantic showed a high consistency in their migration routes. Most individuals migrated more than 3000 km eastwards, while very few remained over the Patagonian Shelf. In contrast, all Indian Ocean birds migrated westwards, resulting in an overlapping nonbreeding area in the eastern Atlantic sector of the Southern Ocean. Geolocators and isotopic signature of feathers indicated that prions from the Falklands moulted at slightly higher latitudes than those from Kerguelen Islands. All birds fed on low trophic level prey, most probably crustaceans. The phenology differed notably between the two populations. Falkland birds returned to the Patagonian Shelf after 2-3 months, while Kerguelen birds remained in the nonbreeding area for seven months, before returning to nesting grounds highly synchronously and at high speed. Habitat models identified sea surface temperature and chlorophyll a concentration as important environmental parameters. In summary, we show that even though the two very distant populations migrate to roughly the same area to moult, they have distinct wintering strategies: They had significantly different realized niches and timing which may contribute to spatial niche partitioning.

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

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.

Feather corticosterone levels on wintering grounds have no carry-over effects on breeding among three populations of great skuas (Stercorarius skua).

Environmental conditions encountered by migratory seabirds in their wintering areas can shape their fitness. However, the underlying physiological mechanisms remain largely unknown as birds are relatively inaccessible during winter. To assess physiological condition during this period, we measured corticosterone concentrations in winter-grown primary feathers of female great skuas (Stercorarius skua) from three breeding colonies (Bjørnøya, Iceland, Shetland) with wintering areas identified from characteristic stable isotope signatures. We subsequently compared winter feather corticosterone levels between three wintering areas (Africa, Europe and America). Among females breeding in 2009, we found significant differences in feather corticosterone levels between wintering areas. Surprisingly, levels were significantly higher in Africa despite seemingly better local ecological factors (based on lower foraging effort). Moreover, contrary to our predictions, females sharing the same wintering grounds showed significant differences in feather corticosterone levels depending on their colony of origin suggesting that some skuas could be using suboptimal wintering areas. Among females wintering in Africa, Shetland females showed feather corticosterone levels on average 22% lower than Bjørnøya and Iceland females. Finally, the lack of significant relationships between winter feather corticosterone levels and any of the breeding phenology traits does not support the hypothesis of potential carry-over effects of winter feather corticosterone. Yet, the fitness consequences of elevated feather corticosterone levels remain to be determined.

Isotopic analysis of island House Martins Delichon urbica indicates marine provenance of nutrients.

The presence of one of the largest colonies of House Martins in Europe on the small island of Stora Karlsö, Sweden, led us to investigate the source of their food by analysis of stable isotopes of carbon and nitrogen. Carbon isotopic values of House Martin nestlings were the same as those of Common Guillemot Uria aalge nestlings fed on marine fish, but differed from local Collared Flycatcher Ficedula albicollis nestlings fed on woodland insects. We infer that these House Martins fed their chicks almost exclusively on insects that had used nutrients derived from seabirds, indicating a dependence on the presence of a large seabird colony. We suggest by extension that some populations of island passerines of high conservation importance may also be dependent on nutrient subsidies from seabird colonies.

Dietary versus maternal sources of organochlorines in top predator seabird chicks: an experimental approach.

We examined the relative importance of dietary sources and maternal transfer on organochlorine concentrations (∑OCs) in Great skua chicks (Stercorarius skua) in Shetland by food supplementing parents with known wintering area. We predicted that experimental chicks (whose parents were supplemented) should have (i) higher growth rates and, (ii) lower ∑OCs due to growth dilution effect and/or due to being fed with less contaminated food compared to control chicks. We also predicted a significant influence of maternal wintering area on chicks' ∑OCs. Plasma ∑OCs of adults, assessed prior to the manipulation, significantly differed between wintering areas of birds. Chicks were weighed every 5 days and plasma ∑OCs were assessed at 20 days old. Based on nitrogen and carbon stable isotope analysis, the supplementary food contributed on average 20% of the dietary protein of the chicks. Although experimental chicks experienced better developmental conditions, supplementary food did not alleviate their organochlorine burden. Nevertheless, chicks whose mothers wintered in Europe showed ∑OCs 50% higher than chicks whose mothers wintered in Africa. Moreover, based on the positive relationship between ∑OCs of chicks and females, the contaminant load of Great skua chicks in Shetland appears to be more influenced by maternal transfer than by trophic transfer.