An ecologically sound and participatory monitoring network for pan-Arctic seabirds.

In a warming Arctic, circumpolar long-term monitoring programs are key to advancing ecological knowledge and informing environmental policies. Calls for better involvement of Arctic peoples in all stages of the monitoring process are widespread, although such transformation of Arctic science is still in its infancy. Seabirds stand out as ecological sentinels of environmental changes, and priority has been given to implement the Circumpolar Seabird Monitoring Plan (CSMP). We assessed the representativeness of a pan-Arctic seabird monitoring network focused on the black-legged kittiwake (Rissa tridactyla) by comparing the distribution of environmental variables for all known versus monitored colonies. We found that with respect to its spatiotemporal coverage, this monitoring network does not fully embrace current and future environmental gradients. To improve the current scheme, we designed a method to identify colonies whose inclusion in the monitoring network will improve its ecological representativeness, limit logistical constraints, and improve involvement of Arctic peoples. We thereby highlight that inclusion of study sites in the Bering Sea, Siberia, western Russia, northern Norway, and southeastern Greenland could improve the current monitoring network and that their proximity to local populations might allow increased involvement of local communities. Our framework can be applied to improve existing monitoring networks in other ecoregions and sociological contexts.

Una red de monitoreo participativa y ecológica para las aves marinas del Ártico Resumen En un Ártico cada vez más cálido, los programas circumpolares de monitoreo a largo plazo son importantes para potenciar el conocimiento ecológico e informar las políticas ambientales. Existe un llamado generalizado para involucrar de mejor manera a los pueblos árticos en el proceso de monitoreo, aunque dicha transformación de la ciencia ártica todavía está en desarrollo. Las aves marinas resaltan como centinelas del cambio ambiental y se ha priorizado implementar el Plan Circumpolar de Monitoreo de Aves Marinas (CSMP). Comparamos la distribución de las variables ambientales de todas las colonias conocidas de la gaviota tridáctila (Rissa tridactyla) contra las colonias monitoreadas para evaluar la representación de una red pan­ártica de monitoreo enfocada en esta especie. Encontramos que esta red de monitoreo no considera del todo los gradientes ambientales actuales y futuros con respecto a la cobertura espaciotemporal. Para mejorar el esquema actual, diseñamos un método para identificar las colonias cuya inclusión en la red de monitoreo mejorará su representación ecológica, limitará las restricciones logísticas e incrementará la participación de los pueblos árticos. Por lo tanto, resaltamos que la inclusión de los sitios de estudio en el Mar de Bering, Siberia, Rusia occidental, el norte de Noruega y el sureste de Groenlandia mejorarían la red actual de monitoreo. También destacamos que la proximidad de los sitios de estudio con las poblaciones locales podría permitir una mayor participación de estas. Nuestro marco puede aplicarse para mejorar las redes de monitoreo existentes en otros contextos socioecológicos y ecoregiones.

Seabirds reveal mercury distribution across the North Atlantic.

Mercury (Hg) is a heterogeneously distributed toxicant affecting wildlife and human health. Yet, the spatial distribution of Hg remains poorly documented, especially in food webs, even though this knowledge is essential to assess large-scale risk of toxicity for the biota and human populations. Here, we used seabirds to assess, at an unprecedented population and geographic magnitude and high resolution, the spatial distribution of Hg in North Atlantic marine food webs. To this end, we combined tracking data of 837 seabirds from seven different species and 27 breeding colonies located across the North Atlantic and Atlantic Arctic together with Hg analyses in feathers representing individual seabird contamination based on their winter distribution. Our results highlight an east-west gradient in Hg concentrations with hot spots around southern Greenland and the east coast of Canada and a cold spot in the Barents and Kara Seas. We hypothesize that those gradients are influenced by eastern (Norwegian Atlantic Current and West Spitsbergen Current) and western (East Greenland Current) oceanic currents and melting of the Greenland Ice Sheet. By tracking spatial Hg contamination in marine ecosystems and through the identification of areas at risk of Hg toxicity, this study provides essential knowledge for international decisions about where the regulation of pollutants should be prioritized.

The challenges of opportunistic sampling when comparing prevalence of plastics in diving seabirds: A multi-species example from Norway.

There is a need for baseline information about how much plastics are ingested by wildlife and potential negative consequences thereof. We analysed the frequency of occurrence (FO) of plastics >1 mm in the stomachs of five pursuit-diving seabird species collected opportunistically. Atlantic puffins (Fratercula arctica) found emaciated on beaches in SW Norway had the highest FO of plastics (58.8 %), followed by emaciated common guillemots (Uria aalge; 9.1 %) also found beached in either SW or SE Norway. No plastics were detected in razorbills (Alca torda), great cormorants (Phalacrocorax carbo), and European shags (Gulosus aristotelis) taken as bycatch in northern Norway. This is the first study to report on plastic ingestion of these five species in northern Europe, and it highlights both the usefulness and limitations of opportunistic sampling. Small sample sizes, as well as an unbalanced sample design, complicated the interpretation of the results.

Hybridization of Atlantic puffins in the Arctic coincides with 20th-century climate change.

The Arctic is experiencing the fastest rates of global warming, leading to shifts in the distribution of its biota and increasing the potential for hybridization. However, genomic evidence of recent hybridization events in the Arctic remains unexpectedly rare. Here, we use whole-genome sequencing of contemporary and 122-year-old historical specimens to investigate the origin of an Arctic hybrid population of Atlantic puffins (Fratercula arctica) on Bjørnøya, Norway. We show that the hybridization between the High Arctic, large-bodied subspecies F. a. naumanni and the temperate, smaller-sized subspecies F. a. arctica began as recently as six generations ago due to an unexpected southward range expansion of F. a. naumanni. Moreover, we find a significant temporal loss of genetic diversity across Arctic and temperate puffin populations. Our observations provide compelling genomic evidence of the impacts of recent distributional shifts and loss of diversity in Arctic communities during the 20th century.

Consequences of cross-season demographic correlations for population viability.

Demographic correlations are pervasive in wildlife populations and can represent important secondary drivers of population growth. Empirical evidence suggests that correlations are in general positive for long-lived species, but little is known about the degree of variation among spatially segregated populations of the same species in relation to environmental conditions. We assessed the relative importance of two cross-season correlations in survival and productivity, for three Atlantic puffin (Fratercula arctica) populations with contrasting population trajectories and non-overlapping year-round distributions. The two correlations reflected either a relationship between adult survival prior to breeding on productivity, or a relationship between productivity and adult survival the subsequent year. Demographic rates and their correlations were estimated with an integrated population model, and their respective contributions to variation in population growth were calculated using a transient-life table response experiment. For all three populations, demographic correlations were positive at both time lags, although their strength differed. Given the different year-round distributions of these populations, this variation in the strength population-level demographic correlations points to environmental conditions as an important driver of demographic variation through life-history constraints. Consequently, the contributions of variances and correlations in demographic rates to population growth rates differed among puffin populations, which has implications for-particularly small-populations' viability under environmental change as positive correlations tend to reduce the stochastic population growth rate.

Raising offspring increases ageing: Differences in senescence among three populations of a long-lived seabird, the Atlantic puffin.

Actuarial senescence, the decline of survival with age, is well documented in the wild. Rates of senescence vary widely between taxa, to some extent also between sexes, with the fastest life histories showing the highest rates of senescence. Few studies have investigated differences in senescence among populations of the same species, although such variation is expected from population-level differences in environmental conditions, leading to differences in vital rates and thus life histories. We predict that, within species, populations differing in productivity (suggesting different paces of life) should experience different rates of senescence, but with little or no sexual difference in senescence within populations of monogamous, monomorphic species where the sexes share breeding duties. We compared rates of actuarial senescence among three contrasting populations of the Atlantic puffin Fratercula arctica. The dataset comprised 31 years (1990-2020) of parallel capture-mark-recapture data from three breeding colonies, Isle of May (North Sea), Røst (Norwegian Sea) and Hornøya (Barents Sea), showing contrasting productivities (i.e. annual breeding success) and population trends. We used time elapsed since first capture as a proxy for bird age, and productivity and the winter North Atlantic Oscillation Index (wNAO) as proxies for the environmental conditions experienced by the populations within and outside the breeding season, respectively. In accordance with our predictions, we found that senescence rates differed among the study populations, with no evidence for sexual differences. There was no evidence for an effect of wNAO, but the population with the lowest productivity, Røst, showed the lowest rate of senescence. As a consequence, the negative effect of senescence on the population growth rate (λ) was up to 3-5 times smaller on Røst (Δλ = -0.009) than on the two other colonies. Our findings suggest that environmentally induced differences in senescence rates among populations of a species should be accounted for when predicting effects of climate variation and change on species persistence. There is thus a need for more detailed information on how both actuarial and reproductive senescence influence vital rates of populations of the same species, calling for large-scale comparative studies.

Variation and correlation in the timing of breeding of North Atlantic seabirds across multiple scales.

Timing of breeding, an important driver of fitness in many populations, is widely studied in the context of global change, yet despite considerable efforts to identify environmental drivers of seabird nesting phenology, for most populations we lack evidence of strong drivers. Here we adopt an alternative approach, examining the degree to which different populations positively covary in their annual phenology to infer whether phenological responses to environmental drivers are likely to be (a) shared across species at a range of spatial scales, (b) shared across populations of a species or (c) idiosyncratic to populations. We combined 51 long-term datasets on breeding phenology spanning 50 years from nine seabird species across 29 North Atlantic sites and examined the extent to which different populations share early versus late breeding seasons depending on a hierarchy of spatial scales comprising breeding site, small-scale region, large-scale region and the whole North Atlantic. In about a third of cases, we found laying dates of populations of different species sharing the same breeding site or small-scale breeding region were positively correlated, which is consistent with the hypothesis that they share phenological responses to the same environmental conditions. In comparison, we found no evidence for positive phenological covariation among populations across species aggregated at larger spatial scales. In general, we found little evidence for positive phenological covariation between populations of a single species, and in many instances the inter-year variation specific to a population was substantial, consistent with each population responding idiosyncratically to local environmental conditions. Black-legged kittiwake Rissa tridactyla was the exception, with populations exhibiting positive covariation in laying dates that decayed with the distance between breeding sites, suggesting that populations may be responding to a similar driver. Our approach sheds light on the potential factors that may drive phenology in our study species, thus furthering our understanding of the scales at which different seabirds interact with interannual variation in their environment. We also identify additional systems and phenological questions to which our inferential approach could be applied.

North Atlantic winter cyclones starve seabirds.

Each winter, the North Atlantic Ocean is the stage for numerous cyclones, the most severe ones leading to seabird mass-mortality events called "winter wrecks."1-3 During these, thousands of emaciated seabird carcasses are washed ashore along European and North American coasts. Winter cyclones can therefore shape seabird population dynamics4,5 by affecting survival rates as well as the body condition of surviving individuals and thus their future reproduction. However, most often the geographic origins of impacted seabirds and the causes of their deaths remain unclear.6 We performed the first ocean-basin scale assessment of cyclone exposure in a seabird community by coupling winter tracking data for ∼1,500 individuals of five key North Atlantic seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia, and Rissa tridactyla) and cyclone locations. We then explored the energetic consequences of different cyclonic conditions using a mechanistic bioenergetics model7 and tested the hypothesis that cyclones dramatically increase seabird energy requirements. We demonstrated that cyclones of high intensity impacted birds from all studied species and breeding colonies during winter but especially those aggregating in the Labrador Sea, the Davis Strait, the surroundings of Iceland, and the Barents Sea. Our broad-scale analyses suggested that cyclonic conditions do not increase seabird energy requirements, implying that they die because of the unavailability of their prey and/or their inability to feed during cyclones. Our study provides essential information on seabird cyclone exposure in a context of marked cyclone regime changes due to global warming.8.

Complex population structure of the Atlantic puffin revealed by whole genome analyses.

The factors underlying gene flow and genomic population structure in vagile seabirds are notoriously difficult to understand due to their complex ecology with diverse dispersal barriers and extensive periods at sea. Yet, such understanding is vital for conservation management of seabirds that are globally declining at alarming rates. Here, we elucidate the population structure of the Atlantic puffin (Fratercula arctica) by assembling its reference genome and analyzing genome-wide resequencing data of 72 individuals from 12 colonies. We identify four large, genetically distinct clusters, observe isolation-by-distance between colonies within these clusters, and obtain evidence for a secondary contact zone. These observations disagree with the current taxonomy, and show that a complex set of contemporary biotic factors impede gene flow over different spatial scales. Our results highlight the power of whole genome data to reveal unexpected population structure in vagile marine seabirds and its value for seabird taxonomy, evolution and conservation.

Centennial relationships between ocean temperature and Atlantic puffin production reveal shifting decennial trends.

The current warming of the oceans has been shown to have detrimental effects for a number of species. An understanding of the underlying mechanisms may be hampered by the non-linearity and non-stationarity of the relationships between temperature and demography, and by the insufficient length of available time series. Most demographic time series are too short to study the effects of climate on wildlife in the classical sense of meteorological patterns over at least 30 years. Here we present a harvest time series of Atlantic puffins (Fratercula arctica) that goes back as far as 1880. It originates in the world's largest puffin colony, in southwest Iceland, which has recently experienced a strong decline. By estimating an annual chick production index for 128 years, we found prolonged periods of strong correlations between local sea surface temperature (SST) and chick production. The sign of decennial correlations switches three times during this period, where the phases of strong negative correlations between puffin productivity and SST correspond to the early 20th century Arctic warming period and to the most recent decades. Most of the variation (72%) in chick production is explained by a model in which productivity peaks at an SST of 7.1°C, clearly rejecting the assumption of a linear relationship. There is also evidence supporting non-stationarity: The SST at which puffins production peaked has increased by 0.24°C during the 20th century, although the increase in average SST during the same period has been more than three times faster. The best supported models indicate that the population's decline is at least partially caused by the increasing SST around Iceland.

Local prey shortages drive foraging costs and breeding success in a declining seabird, the Atlantic puffin.

As more and more species face anthropogenic threats, understanding the causes of population declines in vulnerable taxa is essential. However, long-term datasets, ideal to identify lasting or indirect effects on fitness measures such as those caused by environmental factors, are not always available. Here we use a single year but multi-population approach on populations with contrasting demographic trends to identify possible drivers and mechanisms of seabird population changes in the north-east Atlantic, using the Atlantic puffin, a declining species, as a model system. We combine miniature GPS trackers with camera traps and DNA metabarcoding techniques on four populations across the puffins' main breeding range to provide the most comprehensive study of the species' foraging ecology to date. We find that puffins use a dual foraging tactic combining short and long foraging trips in all four populations, but declining populations in southern Iceland and north-west Norway have much greater foraging ranges, which require more (costly) flight, as well as lower chick-provisioning frequencies, and a more diverse but likely less energy-dense diet, than stable populations in northern Iceland and Wales. Together, our findings suggest that the poor productivity of declining puffin populations in the north-east Atlantic is driven by breeding adults being forced to forage far from the colony, presumably because of low prey availability near colonies, possibly amplified by intraspecific competition. Our results provide valuable information for the conservation of this and other important North-Atlantic species and highlight the potential of multi-population approaches to answer important questions about the ecological drivers of population trends.

New tools to evaluate plastic ingestion by northern fulmars applied to North Sea monitoring data 2002-2018.

Monitoring plastic in stomachs of beached northern fulmars for OSPAR's Ecological Quality Objectives (EcoQOs) has been incorporated into the European Marine Strategy Framework Directive (MSFD). This paper aims to provide the appropriate tools to interpret the monitoring results. MSFD requires a data-derived threshold value (Fulmar-TV) representing 'Good Environmental Status'. Such Fulmar-TV was calculated from near-pristine Canadian Arctic data where 10.06% of fulmars exceeded the level of 0.1 g ingested plastic. This Fulmar-TV is almost identical to the earlier OSPAR EcoQO, arbitrarily set at 10%. The MSFD approach was evaluated for 2661 North Sea fulmars in 2002-2018. Between 2014 and 2018, 51% of 393 fulmars exceeded 0.1 g plastic, significantly above the proposed Fulmar-TV. Linear regression of individual ingested plastic mass over the 2009-2018 period indicates a significant decrease. Over the longer term 2002-2018, logistic regression of annual EcoQ% shows a significant decline and predicts compliance with the Fulmar-TV by 2054.

Ingested plastics in northern fulmars (Fulmarus glacialis): A pathway for polybrominated diphenyl ether (PBDE) exposure?

Although it has been suggested that plastic may act as a vector for pollutants into the tissue of seabirds, the bioaccumulation of harmful contaminants, such as polybrominated diphenyl ethers (PBDEs), released from ingested plastics is poorly understood. Plastic ingestion by the procellariiform species northern fulmar (Fulmarus glacialis) is well documented. In this study, we measured PBDEs levels in liver tissue of northern fulmars without and with (0.13-0.43 g per individual) stomach plastics. PBDE concentrations in the plastic sampled from the same birds were also quantified. Birds were either found dead on beaches in southern Norway or incidentally caught in longline fisheries in northern Norway. PBDEs were detected in all birds but high concentrations were only found in liver samples from beached birds, peaking at 2900 ng/g lipid weight. We found that body condition was a significant factor explaining the elevated concentration levels in livers of beached birds. BDE209 was found in ingested plastic particles and liver tissue of birds with ingested plastics but was absent in the livers of birds without ingested plastics. This strongly suggests a plastic-derived transfer and accumulation of BDE209 to the tissue of fulmars, levels of which might prove useful as a general indicator of plastic ingestion in seabirds.

Meeting Paris agreement objectives will temper seabird winter distribution shifts in the North Atlantic Ocean.

We explored the implications of reaching the Paris Agreement Objective of limiting global warming to <2°C for the future winter distribution of the North Atlantic seabird community. We predicted and quantified current and future winter habitats of five North Atlantic Ocean seabird species (Alle alle, Fratercula arctica, Uria aalge, Uria lomvia and Rissa tridactyla) using tracking data for ~1500 individuals through resource selection functions based on mechanistic modeling of seabird energy requirements, and a dynamic bioclimate envelope model of seabird prey. Future winter distributions were predicted to shift with climate change, especially when global warming exceed 2°C under a "no mitigation" scenario, modifying seabird wintering hotspots in the North Atlantic Ocean. Our findings suggest that meeting Paris agreement objectives will limit changes in seabird selected habitat location and size in the North Atlantic Ocean during the 21st century. We thereby provide key information for the design of adaptive marine-protected areas in a changing ocean.

Diverging phenological responses of Arctic seabirds to an earlier spring.

The timing of annual events such as reproduction is a critical component of how free-living organisms respond to ongoing climate change. This may be especially true in the Arctic, which is disproportionally impacted by climate warming. Here, we show that Arctic seabirds responded to climate change by moving the start of their reproduction earlier, coincident with an advancing onset of spring and that their response is phylogenetically and spatially structured. The phylogenetic signal is likely driven by seabird foraging behavior. Surface-feeding species advanced their reproduction in the last 35 years while diving species showed remarkably stable breeding timing. The earlier reproduction for Arctic surface-feeding birds was significant in the Pacific only, where spring advancement was most pronounced. In both the Atlantic and Pacific, seabirds with a long breeding season showed a greater response to the advancement of spring than seabirds with a short breeding season. Our results emphasize that spatial variation, phylogeny, and life history are important considerations in seabird phenological response to climate change and highlight the key role played by the species' foraging behavior.

Spatial and temporal variations in seabird bycatch: Incidental bycatch in the Norwegian coastal gillnet-fishery.

The general decline of seabird populations worldwide raises large concerns. Although multiple factors are interacting to cause the observed trends, increased mortality from incidental bycatch in fisheries has proven to be important for many species. However, the bulk of published knowledge is derived from longline fisheries, whereas bycatch in gillnet fisheries is less studied and even overlooked in some areas. We present seabird bycatch data from a 10-year time-series of fishery data from the large fleet of small-vessels fishing with gillnets along the Norwegian coast-a large area and fishery with no prior estimates of seabird bycatch. In general, we document high rates of incidental bycatch (averaging 0.0023 seabirds/net, or approximately 0.08 seabirds/fishing trip). This results in an estimated annual bycatch between 1580 and 11500 (95% CI) birds in this fishery. There was a surprisingly high percentage (43%) of surface-feeding seabirds in the bycatch, with northern fulmar being the most common species. Among the diving seabirds caught, common guillemot was most numerous. Our findings suggest that coastal gillnet fisheries represent a more general threat to a wider range of seabird populations, as opposed to longline fisheries where surface-feeding seabird species seem to dominate the bycatch. The bycatch estimates for the Norwegian gillnet-fishery varied in time, between areas, and with fishing depth and distance from the coast, but we found no clear trends in relation to the type of gillnets used. The results enabled us to identify important spatio-temporal trends in the seabird bycatch, which can allow for the development and implementation of more specific mitigation measures. While specific time closures might be an efficient option to reduce bycatch for diving seabirds, measures such as gear modification and reduction in release of wastewater during fishing operation are probably a more effective mitigation approach for reducing bycatch of surface-feeding seabirds.

Ocean-wide Drivers of Migration Strategies and Their Influence on Population Breeding Performance in a Declining Seabird.

Which factors shape animals' migration movements across large geographical scales, how different migratory strategies emerge between populations, and how these may affect population dynamics are central questions in the field of animal migration [1] that only large-scale studies of migration patterns across a species' range can answer [2]. To address these questions, we track the migration of 270 Atlantic puffins Fratercula arctica, a red-listed, declining seabird, across their entire breeding range. We investigate the role of demographic, geographical, and environmental variables in driving spatial and behavioral differences on an ocean-basin scale by measuring puffins' among-colony differences in migratory routes and day-to-day behavior (estimated with individual daily activity budgets and energy expenditure). We show that competition and local winter resource availability are important drivers of migratory movements, with birds from larger colonies or with poorer local winter conditions migrating further and visiting less-productive waters; this in turn led to differences in flight activity and energy expenditure. Other behavioral differences emerge with latitude, with foraging effort and energy expenditure increasing when birds winter further north in colder waters. Importantly, these ocean-wide migration patterns can ultimately be linked with breeding performance: colony productivity is negatively associated with wintering latitude, population size, and migration distance, which demonstrates the cost of competition and migration on future breeding and the link between non-breeding and breeding periods. Our results help us to understand the drivers of animal migration and have important implications for population dynamics and the conservation of migratory species.

Circumpolar dynamics of a marine top-predator track ocean warming rates.

Global warming is a nonlinear process, and temperature may increase in a stepwise manner. Periods of abrupt warming can trigger persistent changes in the state of ecosystems, also called regime shifts. The responses of organisms to abrupt warming and associated regime shifts can be unlike responses to periods of slow or moderate change. Understanding of nonlinearity in the biological responses to climate warming is needed to assess the consequences of ongoing climate change. Here, we demonstrate that the population dynamics of a long-lived, wide-ranging marine predator are associated with changes in the rate of ocean warming. Data from 556 colonies of black-legged kittiwakes Rissa tridactyla distributed throughout its breeding range revealed that an abrupt warming of sea-surface temperature in the 1990s coincided with steep kittiwake population decline. Periods of moderate warming in sea temperatures did not seem to affect kittiwake dynamics. The rapid warming observed in the 1990s may have driven large-scale, circumpolar marine ecosystem shifts that strongly affected kittiwakes through bottom-up effects. Our study sheds light on the nonlinear response of a circumpolar seabird to large-scale changes in oceanographic conditions and indicates that marine top predators may be more sensitive to the rate of ocean warming rather than to warming itself.

Modelled drift patterns of fish larvae link coastal morphology to seabird colony distribution.

Colonial breeding is an evolutionary puzzle, as the benefits of breeding in high densities are still not fully explained. Although the dynamics of existing colonies are increasingly understood, few studies have addressed the initial formation of colonies, and empirical tests are rare. Using a high-resolution larval drift model, we here document that the distribution of seabird colonies along the Norwegian coast can be explained by variations in the availability and predictability of fish larvae. The modelled variability in concentration of fish larvae is, in turn, predicted by the topography of the continental shelf and coastline. The advection of fish larvae along the coast translates small-scale topographic characteristics into a macroecological pattern, viz. the spatial distribution of top-predator breeding sites. Our findings provide empirical corroboration of the hypothesis that seabird colonies are founded in locations that minimize travel distances between breeding and foraging locations, thereby enabling optimal foraging by central-place foragers.