Marine Ecology - Progress Series, volume 755, pages 163-177

Current understanding of how climate change affects seabirds varies between regions and species in the North-East Atlantic

DT JOHNSTON 1
EM Humphreys 1
Jg Davies 1
T. Evans 2, 3
R. Howells 2
JW Pearce-Higgins 4, 5, 6
2
 
Marine Directorate, Victoria Quay, Leith, Edinburgh EH6 6QQ, UK
3
 
RSPB Scotland Headquarters, 2 Lochside View, Edinburgh Park, Edinburgh EH12 9DH, UK
4
 
British Trust for Ornithology, The Nunnery, Thetford IP24 2PU, UK
Publication typeJournal Article
Publication date2025-02-27
scimago Q1
SJR0.802
CiteScore5.3
Impact factor2.2
ISSN01718630, 16161599
Abstract

Climate change is regarded as a major threat to seabird populations globally. While the impacts on certain species are well-studied, the majority remain understudied, which may limit our understanding of the key mechanisms linking climate change to population dynamics. This information is important to identify vulnerable species and inform potential conservation responses. To address this gap, we reviewed literature on the impacts of climate change on the demography of 25 seabird species breeding in the UK and Republic of Ireland from relevant studies conducted within the wider North-East Atlantic. We found regional and species-specific variations in research effort, with the North Sea being the most studied area. The most frequently studied species were common guillemot Uria aalge (n = 35), black-legged kittiwake Rissa tridactyla (n = 29), and European shag Gulosus aristotelis (n = 25). Breeding productivity and population abundance were the primary demographic parameters investigated. The North Atlantic Oscillation and sea surface temperature were commonly used to describe climate influences, often linked to bottom-up mechanisms affecting demography through changes in prey availability. Breeding productivity was the most monitored demographic rate across species; this may lead to important mechanisms affecting other demographic rates, such as survival, being overlooked. Regional and species-specific disparities in research could bias the identification of key mechanisms. Despite these uncertainties, current literature highlights the importance of climate impacts through the food chain and severe storm events affecting seabirds. In response, climate change adaptation should prioritise management of fish stocks near breeding colonies and coastal management to protect vulnerable colonies from storm events.

Davies J., Humphreys E., Evans T., Howells R., O’Hara-Murray R., Pearce-Higgins J.
2023-12-21 citations by CoLab: 2 Abstract  
Estimating species’ vulnerability to climate change is a key challenge in conservation. Many seabird species are particularly sensitive to the negative effects of warming and are thus threatened by climate change. We projected seabird abundance in 2050 in Britain and Ireland, an internationally important region for seabirds, under climate change scenario RCP8.5. Our projections were based on fitted relationships between abundance and both climatic (air temperature, precipitation) and oceanographic (sea surface temperature, potential energy anomaly) variables, estimated using a Bayesian hurdle model with spatial random effects. The most consistent responses to environmental variables were negative relationships between species presence or abundance and breeding season maximum terrestrial air temperature. Although data and modelling issues meant that future projections could be made for only 15 of the 25 species and with low predictive power for spatial abundance, predictive power was high for spatial presence and moderate for temporal abundance change. Projections suggest that seabirds will generally fare poorly under climate change in Britain and Ireland, with 10 species projected to decline in abundance by 2050. Variable model fit and projection uncertainty limited confidence, which was generally greater for declines than for increases. Fulmar, puffin and Arctic tern were each projected to decline by over 70% with higher confidence. For a few species, colonisation of new areas may reduce projected losses, but this is potentially limited by low availability of suitable habitat. Projected abundance change was more negative for marine specialists than for generalists. Our findings highlight the vulnerability and conservation needs of seabirds in a changing climate.
Searle K., Butler A., Waggitt J., Evans P., Quinn L., Bogdanova M., Evans T., Braithwaite J., Daunt F.
2022-06-02 citations by CoLab: 9 Abstract  
Europe has set ambitious green energy targets, to which offshore renewable developments (ORDs) will make a significant contribution. Governments are legally required to deliver ORDs sustainably; however, they may have detrimental impacts on wildlife, especially those already experiencing declines due to climate change. Population viability analysis (PVA) is the standard method for forecasting population change in ORD assessments, but PVAs do not currently account for climate effects. We quantified climate effects on seabird breeding success for 8 UK species breeding in the North Sea. We assessed the potential for seabirds to mitigate climate-driven changes in breeding success by accessing wider resources through increased foraging ranges around colonies. We demonstrate strong links between breeding success and climate in 5 species. In 4 of these species, future climate projections indicated large declines in breeding success relative to current rates. Only one species was predicted to increase breeding success under future climate. In all 5 species, there was limited opportunity for species to increase breeding success by expanding foraging ranges to access more suitable future climatic conditions. Climate change will have significant ramifications for future breeding success of seabirds breeding in the North Sea, an area undergoing extensive and rapid offshore renewable energy development. We recommend 3 methods for including climate-driven changes to seabird breeding success within ORD assessments: development of predictive climate-driven habitat use models to estimate ORD-wildlife interactions; delivery of a new ORD assessment framework that includes dynamic predictions of climate-driven habitat use and demography of wildlife populations; and consideration of climate-driven changes in the implementation of compensatory measures.
Bowgen K.M., Kettel E.F., Butchart S.H., Carr J.A., Foden W.B., Magin G., Morecroft M.D., Smith R.K., Stein B.A., Sutherland W.J., Thaxter C.B., Pearce-Higgins J.W.
Biological Conservation scimago Q1 wos Q1
2022-05-01 citations by CoLab: 16 Abstract  
There is an urgent need to quantify the potential for conservation interventions to effectively manage the impacts of climate change on species' populations and ecological communities. In this first quantitative global assessment of biodiversity conservation interventions for climate change adaptation, we identified 77 peer-reviewed studies, including 443 cases describing the response of individual species' populations or assemblages to particular interventions, whilst also accounting for responses to climate change or particular climatic variables. Eighty-two percent of studies were from Europe or North America. In 30% of reported cases, interventions were regarded as beneficial (having a significant positive impact on a population also affected by a climatic variable). However, beneficial outcomes were more likely to be reported when fewer responses were analysed, suggesting a publication bias in the reporting of beneficial responses. Management focused on particular species (e.g. targeted habitat management and species recovery interventions) was modelled to have a higher probability (73%) of being beneficial than more generic interventions such as land and water management (22%) or protection (17%). Although more data on the effectiveness of climate change adaptation for species conservation are required, the diversity of examples reviewed suggests that climate change adaptation can successfully reduce negative impacts of, or enhance positive responses to, climate change. Targeted interventions maximise the persistence of the most vulnerable populations, whilst expanding habitat management and site protection interventions may benefit the largest number of species and ecosystems. The effective monitoring and evaluation of adaptation interventions is required to improve this evidence-base for future decision-making.
Whitehead E.A., Russell J.C., Hickey A.J., Taylor G.A., O'Reilly K.M., Della Penna A., Dunphy B.J.
2022-02-15 citations by CoLab: 2 Abstract  
Seabird breeding success is known to reflect oceanic conditions. Gray-faced petrels (Pterodroma gouldi) breeding on the east coast of Auckland, New Zealand, exhibit poor reproductive success and slow chick development compared to west coast conspecifics. This study mapped changes in physiological traits (corticosterone [CORT] and hematological parameters) indicative of sublethal stress in this Procellariiform species between the west coast (Ihumoana) and east coast (Hāwere) island colonies. We found adult gray-faced petrels on the east coast to be lighter and, unlike west coast birds, exhibited an attenuation of response CORT levels between incubation and chick-rearing phases. Such responses were also reflected in east coast chicks that were lighter and had higher feather CORT titers than west coast chicks. Measures of adult hematology and plasma biochemistry revealed significantly lower glucose levels in east coast birds and indicated that chick rearing is the most stressful phase of breeding for this species Combined; these results suggest that east coast birds are under greater nutritional stress and that parents appear to transfer the costs of poor foraging to their chicks to preserve their own condition, consequently increasing chick developmental stress. Our results suggest that any long-term decrease in ocean conditions and/or climatic shifts would be more acutely felt by east coast chicks and potentially their parents, resulting in years of poor breeding success rates on a local scale.
Choy E.S., O'Connor R.S., Gilchrist H.G., Hargreaves A.L., Love O.P., Vézina F., Elliott K.H.
Journal of Experimental Biology scimago Q1 wos Q2
2021-07-01 citations by CoLab: 28 Abstract  
ABSTRACT The Arctic is warming at approximately twice the global rate, with well-documented indirect effects on wildlife. However, few studies have examined the direct effects of warming temperatures on Arctic wildlife, leaving the importance of heat stress unclear. Here, we assessed the direct effects of increasing air temperatures on the physiology of thick-billed murres (Uria lomvia), an Arctic seabird with reported mortalities due to heat stress while nesting on sun-exposed cliffs. We used flow-through respirometry to measure the response of body temperature, resting metabolic rate, evaporative water loss and evaporative cooling efficiency (the ratio of evaporative heat loss to metabolic heat production) in murres while experimentally increasing air temperature. Murres had limited heat tolerance, exhibiting: (1) a low maximum body temperature (43.3°C); (2) a moderate increase in resting metabolic rate relative that within their thermoneutral zone (1.57 times); (3) a small increase in evaporative water loss rate relative that within their thermoneutral zone (1.26 times); and (4) a low maximum evaporative cooling efficiency (0.33). Moreover, evaporative cooling efficiency decreased with increasing air temperature, suggesting murres were producing heat at a faster rate than they were dissipating it. Larger murres also had a higher rate of increase in resting metabolic rate and a lower rate of increase in evaporative water loss than smaller murres; therefore, evaporative cooling efficiency declined with increasing body mass. As a cold-adapted bird, murres' limited heat tolerance likely explains their mortality on warm days. Direct effects of overheating on Arctic wildlife may be an important but under-reported impact of climate change.
Blagodatski A., Trutneva K., Glazova O., Mityaeva O., Shevkova L., Kegeles E., Onyanov N., Fede K., Maznina A., Khavina E., Yeo S., Park H., Volchkov P.
Pathogens scimago Q2 wos Q2 Open Access
2021-05-20 citations by CoLab: 102 PDF Abstract  
Avian influenza is one of the largest known threats to domestic poultry. Influenza outbreaks on poultry farms typically lead to the complete slaughter of the entire domestic bird population, causing severe economic losses worldwide. Moreover, there are highly pathogenic avian influenza (HPAI) strains that are able to infect the swine or human population in addition to their primary avian host and, as such, have the potential of being a global zoonotic and pandemic threat. Migratory birds, especially waterfowl, are a natural reservoir of the avian influenza virus; they carry and exchange different virus strains along their migration routes, leading to antigenic drift and antigenic shift, which results in the emergence of novel HPAI viruses. This requires monitoring over time and in different locations to allow for the upkeep of relevant knowledge on avian influenza virus evolution and the prevention of novel epizootic and epidemic outbreaks. In this review, we assess the role of migratory birds in the spread and introduction of influenza strains on a global level, based on recent data. Our analysis sheds light on the details of viral dissemination linked to avian migration, the viral exchange between migratory waterfowl and domestic poultry, virus ecology in general, and viral evolution as a process tightly linked to bird migration. We also provide insight into methods used to detect and quantify avian influenza in the wild. This review may be beneficial for the influenza research community and may pave the way to novel strategies of avian influenza and HPAI zoonosis outbreak monitoring and prevention.
Samplonius J.M., Atkinson A., Hassall C., Keogan K., Thackeray S.J., Assmann J.J., Burgess M.D., Johansson J., Macphie K.H., Pearce-Higgins J.W., Simmonds E.G., Varpe Ø., Weir J.C., Childs D.Z., Cole E.F., et. al.
Nature Ecology and Evolution scimago Q1 wos Q1
2020-12-14 citations by CoLab: 75 Abstract  
Climate warming has caused the seasonal timing of many components of ecological food chains to advance. In the context of trophic interactions, the match–mismatch hypothesis postulates that differential shifts can lead to phenological asynchrony with negative impacts for consumers. However, at present there has been no consistent analysis of the links between temperature change, phenological asynchrony and individual-to-population-level impacts across taxa, trophic levels and biomes at a global scale. Here, we propose five criteria that all need to be met to demonstrate that temperature-mediated trophic asynchrony poses a growing risk to consumers. We conduct a literature review of 109 papers studying 129 taxa, and find that all five criteria are assessed for only two taxa, with the majority of taxa only having one or two criteria assessed. Crucially, nearly every study was conducted in Europe or North America, and most studies were on terrestrial secondary consumers. We thus lack a robust evidence base from which to draw general conclusions about the risk that climate-mediated trophic asynchrony may pose to populations worldwide. Five key criteria are proposed to demonstrate robustly that temperature-mediated phenological asynchrony will negatively impact consumers, which the authors show are rarely met in the current literature.
Bedford J., Ostle C., Johns D.G., Atkinson A., Best M., Bresnan E., Machairopoulou M., Graves C.A., Devlin M., Milligan A., Pitois S., Mellor A., Tett P., McQuatters‐Gollop A.
Global Change Biology scimago Q1 wos Q1
2020-04-01 citations by CoLab: 60 Abstract  
Increasing direct human pressures on the marine environment, coupled with climate-driven changes, is a concern to marine ecosystems globally. This requires the development and monitoring of ecosystem indicators for effective management and adaptation planning. Plankton lifeforms (broad functional groups) are sensitive indicators of marine environmental change and can provide a simplified view of plankton biodiversity, building an understanding of change in lower trophic levels. Here, we visualize regional-scale multi-decadal trends in six key plankton lifeforms as well as their correlative relationships with sea surface temperature (SST). For the first time, we collate trends across multiple disparate surveys, comparing the spatially and temporally extensive Continuous Plankton Recorder (CPR) survey (offshore) with multiple long-term fixed station-based time-series (inshore) from around the UK coastline. These analyses of plankton lifeforms showed profound long-term changes, which were coherent across large spatial scales. For example, 'diatom' and 'meroplankton' lifeforms showed strong alignment between surveys and coherent regional-scale trends, with the 1998-2017 decadal average abundance of meroplankton being 2.3 times that of 1958-1967 for CPR samples in the North Sea. This major, shelf-wide increase in meroplankton correlated with increasing SSTs, and contrasted with a general decrease in holoplankton (dominated by small copepods), indicating a changing balance of benthic and pelagic fauna. Likewise, inshore-offshore gradients in dinoflagellate trends, with contemporary increases inshore contrasting with multi-decadal decreases offshore (approx. 75% lower decadal mean abundance), urgently require the identification of causal mechanisms. Our lifeform approach allows the collation of many different data types and time-series across the NW European shelf, providing a crucial evidence base for informing ecosystem-based management, and the development of regional adaptation plans.
Dias M.P., Martin R., Pearmain E.J., Burfield I.J., Small C., Phillips R.A., Yates O., Lascelles B., Borboroglu P.G., Croxall J.P.
Biological Conservation scimago Q1 wos Q1
2019-09-01 citations by CoLab: 543 Abstract  
We present the first objective quantitative assessment of the threats to all 359 species of seabirds, identify the main challenges facing them, and outline priority actions for their conservation. We applied the standardised Threats Classification Scheme developed for the IUCN Red List to objectively assess threats to each species and analysed the data according to global IUCN threat status, taxonomic group, and primary foraging habitat (coastal or pelagic). The top three threats to seabirds in terms of number of species affected and average impact are: invasive alien species, affecting 165 species across all the most threatened groups; bycatch in fisheries, affecting fewer species (100) but with the greatest average impact; and climate change/severe weather, affecting 96 species. Overfishing, hunting/trapping and disturbance were also identified as major threats to seabirds. Reversing the top three threats alone would benefit two-thirds of all species and c. 380 million individual seabirds (c. 45% of the total global seabird population). Most seabirds (c. 70%), especially globally threatened species, face multiple threats. For albatrosses, petrels and penguins in particular (the three most threatened groups of seabirds), it is essential to tackle both terrestrial and marine threats to reverse declines. As the negative effects of climate change are harder to mitigate, it is vital to compensate by addressing other major threats that often affect the same species, such as invasive alien species, bycatch and overfishing, for which proven solutions exist.
Church G.E., Furness R.W., Tyler G., Gilbert L., Votier S.C.
ICES Journal of Marine Science scimago Q1 wos Q1 Open Access
2018-12-10 citations by CoLab: 26 Abstract  
Abstract Understanding anthropogenic impacts are crucial to maintain marine ecosystem health. The North Sea has changed in recent decades, largely due to commercial fishing and climate change. Seabirds can act as useful indicators of these changes. By analyzing n = 20 013 pellets and n = 24 993 otoliths regurgitated by great skuas Stercorarius skua in northern Scotland over five decades from the 1970s to the 2010s (in 36 years 1973–2017), we reveal how the diet of this top predator has changed alongside the changing North Sea ecosystem. Sandeels Ammodytes spp. were the most common dietary item during the 1970s, but became virtually absent from the 1980s onward. Discarded whitefish dominated skua diets from the 1980s to the present day, despite long-term declines in North Sea discard production. However, the discarded fish eaten by great skuas has become smaller and the species composition changed. Skua pellets only rarely contained avian prey in the 1970s but this increased during the 1980s, and fluctuated between 10% and 20% from the 1990s to 2010s. There have also been changes in the avian prey in the diet—black-legged kittiwakes Rissa tridactyla generally being replaced by auks Alcid spp. and northern fulmars Fulmarus glacialis. The Shetland marine ecosystem has experienced steep declines in sandeel stocks and in seabirds that feed on them. Great skuas have been able to prey switch to respond to this change, supported by abundant discards, enabling them to maintain a favourable population status while other seabird species have declined.
Howells R.J., Burthe S.J., Green J.A., Harris M.P., Newell M.A., Butler A., Wanless S., Daunt F.
Marine Biology scimago Q1 wos Q2
2018-11-21 citations by CoLab: 14 Abstract  
Populations of marine top predators are exhibiting pronounced demographic changes due to alterations in prey availability and quality. Changes in diet composition is a key potential mechanism whereby alterations in prey availability can affect predator demography. Studies of long-term trends in diet have focused on the breeding season. However, long-term changes in non-breeding season diet is an important knowledge gap, since this is generally the most critical period of the year for the demography of marine top predators. In this study, we analysed 495,239 otoliths from 5888 regurgitated pellets collected throughout the annual cycle over three decades (1985–2014) from European shags Phalacrocorax aristotelis on the Isle of May, Scotland (56°11′N, 02°33′W). We identified dramatic reductions in the frequency of lesser sandeel Ammodytes marinus occurrence over the study, which was more pronounced during the non-breeding period (96% in 1988 to 45% in 2014), than the breeding period (91–67%). The relative numerical abundance of sandeel per pellet also reduced markedly (100–13% of all otoliths), with similar trends apparent during breeding and non-breeding periods. In contrast, the frequencies of Gadidae, Cottidae, Pleuronectidae and Gobiidae all increased, resulting in a doubling in annual prey richness from 6 prey types per year in 1988 to 12 in 2014. Our study demonstrates that the declining importance of the previously most prominent prey and marked increase in diet diversity is apparent throughout the annual cycle, suggesting that substantial temporal changes in prey populations have occurred, which may have important implications for seabird population dynamics.
Black C., Collen B., Lunn D., Filby D., Winnard S., Hart T.
Ecology and Evolution scimago Q1 wos Q2 Open Access
2018-07-23 citations by CoLab: 8 PDF Abstract  
Variation in the phenology of avian taxa has long been studied to understand how a species reacts to environmental changes over both space and time. Penguins (Sphenicidae) serve as an important example of how biotic and abiotic factors influence certain stages of seabird phenology because of their large ranges and the extreme, dynamic conditions present in their Southern Ocean habitats. Here, we examined the phenology of gentoo (Pygoscelis papua) and chinstrap penguins (Pygoscelis antarctica) at 17 sites across the Scotia arc, including the first documented monitoring of phenology on the South Sandwich Islands, to determine which breeding phases are intrinsic, or rather vary across a species range and between years. We used a novel method to measure seabird breeding phenology and egg and chick survival: time-lapse cameras. Contrary to the long-standing theory that these phases are consistent between colonies, we found that latitude and season had a predominant influence on the length of the nest establishment, incubation, and guard durations. We observe a trend toward longer incubation times occurring farther south, where ambient temperatures are colder, which may indicate that exposure to cold slows embryo growth. Across species, in colonies located farther south, parents abandoned nests later when eggs were lost or chicks died and the latest record of eggs or chicks in the nest occurred earlier during the breeding period. The variation in both space and time observed in penguin phenology provides evidence that the duration of phases within the annual cycle of birds is not fundamental, or genetic, as previously understood. Additionally, the recorded phenology dates should inform field researchers on the best timing to count colonies at the peak of breeding, which is poorly understood.
Keogan K., Daunt F., Wanless S., Phillips R.A., Walling C.A., Agnew P., Ainley D.G., Anker-Nilssen T., Ballard G., Barrett R.T., Barton K.J., Bech C., Becker P., Berglund P., Bollache L., et. al.
Nature Climate Change scimago Q1 wos Q1
2018-04-01 citations by CoLab: 81 Abstract  
Reproductive timing in many taxa plays a key role in determining breeding productivity 1 , and is often sensitive to climatic conditions 2 . Current climate change may alter the timing of breeding at different rates across trophic levels, potentially resulting in temporal mismatch between the resource requirements of predators and their prey 3 . This is of particular concern for higher-trophic-level organisms, whose longer generation times confer a lower rate of evolutionary rescue than primary producers or consumers 4 . However, the disconnection between studies of ecological change in marine systems makes it difficult to detect general changes in the timing of reproduction 5 . Here, we use a comprehensive meta-analysis of 209 phenological time series from 145 breeding populations to show that, on average, seabird populations worldwide have not adjusted their breeding seasons over time (−0.020 days yr−1) or in response to sea surface temperature (SST) (−0.272 days °C−1) between 1952 and 2015. However, marked between-year variation in timing observed in resident species and some Pelecaniformes and Suliformes (cormorants, gannets and boobies) may imply that timing, in some cases, is affected by unmeasured environmental conditions. This limited temperature-mediated plasticity of reproductive timing in seabirds potentially makes these top predators highly vulnerable to future mismatch with lower-trophic-level resources 2 . Time of reproduction may be altered as the climate changes. For seabirds, it is shown that there has not been an adjustment in timing as the climate changes and the sea surface warms. This lack of plasticity could result in a mismatch with food resources.
O'Hanlon N.J., James N.A., Masden E.A., Bond A.L.
Environmental Pollution scimago Q1 wos Q1
2017-12-01 citations by CoLab: 69 Abstract  
Marine plastic pollution is an increasing, and global, environmental issue. Numerous marine species are affected by plastic debris through entanglement, nest incorporation, and ingestion, which can lead to lethal and sub-lethal impacts. However, in the northeastern Atlantic Ocean, an area of international importance for seabirds, there has been little effort to date to assess information from studies of wildlife and plastic to better understand the spatiotemporal variation of how marine plastic affects different seabird species. To improve our understanding of seabirds and marine plastic in this region, we completed a synthesis of the published and grey literature to obtain information on all known documented cases of plastic ingestion and nest incorporation by this group. We found that of 69 seabird species that commonly occur in the northeastern Atlantic, 25 had evidence of ingesting plastic. However, data on plastic ingestion was available for only 49% of all species, with 74% of investigated species recorded ingesting plastic. We found only three published studies on nest incorporation, for the Northern Gannet (Morus bassanus) and Black-legged Kittiwake (Rissa tridactyla). For many species, sample sizes were small or not reported, and only 39% of studies were from the 21st century, whilst information from multiple countries and years was only available for 11 species. This indicates that we actually know very little about the current prevalence of plastic ingestion and nest incorporation for many species, several of them globally threatened. Furthermore, in the majority of studies, the metrics reported were inadequate to carry out robust comparisons among locations and species or perform meta-analyses. We recommend multi-jurisdictional collaboration to obtain a more comprehensive and current understanding of how marine plastic is affecting seabirds in the northeastern Atlantic Ocean.
Scheffers B.R., De Meester L., Bridge T.C., Hoffmann A.A., Pandolfi J.M., Corlett R.T., Butchart S.H., Pearce-Kelly P., Kovacs K.M., Dudgeon D., Pacifici M., Rondinini C., Foden W.B., Martin T.G., Mora C., et. al.
Science scimago Q1 wos Q1 Open Access
2016-11-11 citations by CoLab: 975 PDF Abstract  
Accumulating impacts Anthropogenic climate change is now in full swing, our global average temperature already having increased by 1°C from preindustrial levels. Many studies have documented individual impacts of the changing climate that are particular to species or regions, but individual impacts are accumulating and being amplified more broadly. Scheffers et al. review the set of impacts that have been observed across genes, species, and ecosystems to reveal a world already undergoing substantial change. Understanding the causes, consequences, and potential mitigation of these changes will be essential as we move forward into a warming world. Science , this issue p. 10.1126/science.aaf7671

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