Diel rhythms synchronized to Earth's photic cycle are near-ubiquitous among animals living in regions with distinct day-night cues. Where such cues are reduced or absent, however, activity patterns may weaken, reorganize or become decoupled from the light-dark cycle, which may allow for more flexible behavioural expression. Using a dataset of >900 free-ranging black-legged kittiwakes from colonies spanning a broad latitudinal gradient (40-81°N), we show considerable population-level variation in diel activity patterns, from clear 24 h rhythms with distinct peaks at dawn and dusk at lower latitudes, to variable and non-diel rhythms with flattened activity patterns across the 24 h day at high latitudes. These patterns were consistent with the wide spectrum of light-dark cycles experienced across their breeding range, supported by our finding that activity was strongly predicted by sun elevation, suggesting alignment with the light-dark cycle. Together, our findings reveal substantial within-species variation in rhythmicity across latitudes and demonstrate that changes in the photic environment can reshape the temporal organization of activity even within a single, widely distributed species.
Climate change is playing a major role in the current global biodiversity crisis. However, despite climate change being most pronounced in the Arctic, its impacts on biodiversity in this region remains largely unknown. Here, we combined three decades of abundance data from various animal groups (from zooplankton to megafauna) and regions in the European Arctic and Greenland to assess recent changes in biodiversity within Arctic coastal communities. Our results support the "borealization" hypothesis in all regions and provide evidence that marine ecosystems in the North Atlantic Arctic are shifting toward a boreal (i.e. cold temperate) state. Arctic endemic species are generally declining in abundance, while boreal species are increasing. These changes in abundance are associated with an average increase in biodiversity (e.g. species richness), although there are important variations among animal groups. This increase might be transient and the long-term implications of the ongoing changes in Arctic coastal biodiversity on ecosystem functioning and services remain uncertain.
The behaviour of individual animals to return to a previously used location is referred to as site fidelity and is driven by predictable resource distributions in time and space. Understanding site fidelity in seabirds under global change is important as changes in environmental conditions may vary spatially. This study investigated site fidelity to wintering areas (wintering-site fidelity) of Brünnich's Guillemots Uria lomvia in three West Greenland breeding colonies (Kippaku, Sermilinnguaq, Kitsissut Avalliit) using light-based geolocation data collected over 9 years (2008-2012, 2019-2022). Considering the ocean-scale distribution and long migration of the species, we found that Brünnich's Guillemots from these colonies showed wintering-site fidelity. Mean inter-track distance (MID), as a measure of site fidelity, was 3.5 times greater among birds than among years for the same bird (log scale coefficient = -1.25, 95% confidence limits -1.42 – -1.069 ), suggesting wintering-site fidelity at an individual level. Kippaku and Sermilinnguaq Guillemots had within-individual MIDs of less than 200 km, while among-individual MIDs remained below 700 km. Based on empirical data, Kitsissut Avalliit had a within-individual MID of less than 258 km and an among-individual MID of 724 km. While wintering-site fidelity did not differ between colonies, Kippaku birds showed larger variation in space-use when compared to Sermilinnguaq birds. Wintering-site fidelity did not differ? between sexes. This work highlights that Brünnich’s Guillemots are site faithful to specific areas of the North Atlantic Ocean during their wintering period. The protection of these wintering areas is important for the continued conservation of the species, particularly for declining colonies.
We surgically implanted a 50 g PTT-100 (Platform Transmitter Terminal) in 21 Common Eiders (Somatria mollissima) and a 29 g PTT-100 in 10 Thick-billed Murres (Uria lomvia). After 2-4 months, one Common Eider implanted intracoelomically and two Thick-billed Murres implanted subcutaneously were harvested by local subsistence hunters and examined in the laboratory. External examination of the harvested birds did not reveal any morphological or pathological changes, while the surgical abdominal and cervical wounds seemed to have healed with granulation tissue in all three birds. Necropsy showed chronic inflammation and fatty necrosis in one of the murres, while the antenna Dacron cuffs were at skin level as originally attached for all three birds, with primary tissue healing and no signs of inflammation. In the eider, a few peritoneal adherences were found on the liver without additional signs of inflammation, while one murre had adherences and granulomatous tissue around the PTT with signs of severe inflammation and external rejection. These results indicate that birds can survive implantation of transmitters, even if inflammation develops around the implants. The study points to the importance of continuously refining the techniques for implanting devices in wild birds and performing necropsies on recovered birds.
Developing standardized methodology to allow efficient and cost-effective ecological data collection, particularly at scale, is of critical importance for understanding species' declines. Remote camera networks can enable monitoring across large spatiotemporal scales and at relatively low researcher cost, but manually analysing images and extracting biologically meaningful data is time-consuming. Citizen science image analysis could reduce researcher workload and increase output from large datasets, while actively raising awareness of ecological and conservation issues. Nevertheless, testing the validity of citizen science data collection and the retention of volunteers is essential before integrating these approaches into long-term monitoring programmes. In this study, we used data from a Zooniverse citizen science project, Seabird Watch, to investigate changes in breeding timing of a globally declining seabird species, the Black-legged Kittiwake Rissa tridactyla. Time-lapse cameras collected >200 000 images between 2014 and 2023 across 11 locations covering the species' North Atlantic range (51.7 degrees N-78.9 degrees N), with over 35000 citizen science volunteers 'tagging' adult and juvenile Kittiwakes in images. Most volunteers (81%) classified images for only a single day, and each volunteer classified a median of five images, suggesting that high volunteer recruitment rates are important for the project's continued success. We developed a standardized method to extract colony arrival and departure dates from citizen science annotations, which did not significantly differ from manual analysis by a researcher. We found that Kittiwake colony arrival was 2.6 days later and departure was 1.2 days later per 1 degrees increase in latitude, which was consistent with expectations. Year-round monitoring also showed that Kittiwakes visited one of the lowest latitude colonies, Skellig Michael (51.8 degrees N), during winter, whereas birds from a colony at similar latitude, Skomer Island (51.7 degrees N), did not. Our integrated time-lapse camera and citizen science system offers a cost-effective means of measuring changes in colony attendance and subsequent breeding timing in response to environmental change in cliff-nesting seabirds. This study is of wide relevance to a broad range of species that could be monitored using time-lapse photography, increasing the geographical reach and international scope of ecological monitoring against a background of rapidly changing ecosystems and challenging funding landscapes.
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 SeabirdMonitoring 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.
AimEcologically similar species living in sympatry are expected to segregate to reduce the effects of competition where resources are limiting. Segregation from heterospecifics commonly occurs in space, but it is often unknown whether such segregation has underlying environmental causes. Indeed, species could segregate because of different fundamental environmental requirements (i.e., 'niche divergence'), because competitive exclusion at sympatric sites can force species to either change the habitat use they would have at allopatric sites (i.e., 'niche displacement') or to avoid certain areas, independently of habitat (i.e., 'spatial avoidance'). Testing these hypotheses requires the comparison between sympatric and allopatric sites. Understanding the competitive mechanisms that underlie patterns of spatial segregation could improve predictions of species responses to environmental change, as competition might exacerbate the effects of environmental change.LocationNorth Atlantic and Arctic.TaxaCommon guillemots Uria aalge and Br & uuml;nnich's guillemots Uria lomvia.MethodsHere, we examine support for these explanations for spatial segregation in two closely-related seabird species, common guillemots (Uria aalge) and Br & uuml;nnich's guillemots (U. lomvia). For this, we collated a pan-Atlantic data set of breeding season foraging tracks from 1046 individuals, collected from 20 colonies (8 sympatric and 12 allopatric). These were analysed with habitat models in a spatially transferable framework to compare habitat preferences between species at sympatric and allopatric sites.ResultsWe found no effect of the distribution of heterospecifics on local habitat preferences of the focal species. We found differences in habitat preferences between species, but these were not sufficient to explain the observed levels of spatial segregation at sympatric sites.Main ConclusionsAssuming we did not omit any relevant environmental variables, these results suggest a mix of niche divergence and spatial avoidance produces the observed patterns of spatial segregation.
Scheduling between mates in species with long-term pair bonds can be essential for positive fitness. The annual cycle in photoperiod is the primary environmental cue used by many animals to synchronize behavior and physiology among members of a population, and animals that migrate must have similar annual schedules to ensure successful breeding. However, we know little about whether members of mated pairs in migratory species experience similar photic environments across the year, which could allow for synchronization in annual phenology. Here, we used light-based geolocation to estimate positions of mated pairs of Thick-billed Murres (Uria lomvia, a seabird a.k.a. Brünnich’s Guillemot) which bred above the northern polar circle in Greenland. We tested the hypothesis that individuals in mated pairs occur in more similar locations and photic environments than randomly matched females and males. We found no difference in the amount of spatial separation or in the photic environment between mates and randomized heterosexual pairings. In general, the distance between females and males ranged from 1,198.5 km during August to 737.4 km during January. The sexes remained in photic environments with highly correlated photoperiods and moderately correlated times of solar noon in UTC. The spatial separation of, but similar photic environments experienced by, female and male murres regardless of pair status is probably adaptive by facilitating the synchronization of annual schedules between sexes, while allowing individuals in mated pairs the freedom to pursue the best foraging opportunities during migration and overwintering independent of their mate.
Alteration and diminution in sea ice cover in the Arctic region will give rise to an intensification and expansion of fishing activities in the Arctic and associated marginal seas. Increased fishing activity, especially in the summer, could pose a direct threat to the millions of seabirds breeding in this region, as well as non-breeding migrants, and potentially result in an increase of bycatch mortality. To inform what conservation and management actions may be needed, an analysis of where seabirds/fisheries interaction are most likely to occur is required. Here, we establish what information would be required to effectively model circumpolar bycatch risk of seabirds in the Arctic, and then we assess the availability of the requisite data. The quality and availability of fishing effort, and bycatch monitoring effort data are not homogeneous among Arctic countries. Undertaking a true circumpolar analysis at this time would be difficult, and with the current data accessibility, many assumptions would have to be made, potentially leading to caveats in the results. Improved communications between the various agencies and institutes working on fisheries and seabirds would strengthen the quantitative basis for future analyses. We offer suggestions on how to improve bycatch estimates and the identification of high-risk areas for seabird bycatch in the Arctic.
Mercury (Hg) is a metallic trace element toxic for humans and wildlife that can originate from natural and anthropic sources. Hg spatial gradients have been found in seabirds from the Arctic and other oceans, suggesting contrasting toxicity risks across regions. Selenium (Se) plays a protective role against Hg toxicity, but its spatial distribution has been much less investigated than that of Hg. From 2015 to 2017, we measured spatial co-exposure of Hg and Se in blood samples of two seabird species, the Brünnich's guillemot (Uria lomvia) and the black-legged kittiwake (Rissa tridactyla) from 17 colonies in the Arctic and subarctic regions, and we calculated their molar ratios (Se:Hg), as a measure of Hg sequestration by Se and, therefore, of Hg exposure risk. We also evaluated concentration differences between species and ocean basins (Pacific-Arctic and Atlantic-Arctic), and examined the influence of trophic ecology on Hg and Se concentrations using nitrogen and carbon stable isotopes. In the Atlantic-Arctic ocean, we found a negative west-to-east gradient of Hg and Se for guillemots, and a positive west-to-east gradient of Se for kittiwakes, suggesting that these species are better protected from Hg toxicity in the European Arctic. Differences in Se gradients between species suggest that they do not follow environmental Se spatial variations. This, together with the absence of a general pattern for isotopes influence on trace element concentrations, could be due to foraging ecology differences between species. In both oceans, the two species showed similar Hg concentrations, but guillemots showed lower Se concentrations and Se:Hg than kittiwakes, suggesting a higher Hg toxicity risk in guillemots. Within species, neither Hg, nor Se or Se:Hg differed between both oceans. Our study highlights the importance of considering Se together with Hg, along with different species and regions, when evaluating Hg toxic effects on marine predators in international monitoring programs.
Substantial seabird bycatch occurs in North Atlantic lumpfish fisheries, potentially impacting seabird populations and raising concerns about the fisheries’ sustainability. Despite increased efforts to address this issue, existing approaches have yielded inconclusive and mixed results. In this study, we explored an alternative approach by comparing bycatch levels between conventional gillnets and LED-illuminated gillnets during the 2023 Greenland lumpfish fishing season. Results indicated that LED-illuminated nets showed almost no difference in bycatch compared to no-LED nets for the three main bird species (common eider, king eider, and long-tailed duck), with no statistically significant differences observed. Additionally, the LED-illuminated nets exhibited a slightly lower, but insignificant, catch rate of the targeted female lumpfish. In conclusion, our findings suggest that using LED as a bird deterrent in this fishery is not an optimal solution to bycatch mitigation.
Bycatch in gillnets is a global issue and mitigation measures that balance target species catch rates, bycatch reduction and fisher support are scarce. In the North Atlantic lumpfish fisheries, bycatch includes marine mammals and seabirds, and there are no permanent technical initiatives to reduce the bycatch. In the West Greenland fishery, common eider bycatch is several thousand individuals annually. We explored if bycatch in this fishery could be reduced by modifying standard lumpfish gillnets by adding a 45 cm high small-meshed net panel to the bottom part of the net. We tested the nets in combination with standard nets and estimated catch rates in a controlled setting in 2021 and in the commercial fishery in 2022. The modified nets had a 71% reduced bycatch rate for common eider and a 25% reduced catch rate for female lumpfish. A combination of the panel and increased seaweed entanglement was the most likely explanation for the effect. In addition to the effect of the net modification, the common eider bycatch decreased significantly during the season, and we recommend studying the net effect further and exploring the option of postponing the fishing season as a simpler way of reducing bycatch.
Marine traffic poses a growing threat to wildlife in the marine environment, including Arctic seabirds, which are exposed to high vessel densities when breeding in coastal areas. However, little is known about the magnitude of the problem. Here, we utilized underwater acoustic monitoring to quantify marine traffic and above-water disturbances at two thick-billed murre colonies in Greenland in 2016. We detected a total of 307 vessels, and only 4 % was known from automatic monitoring systems. Based on proximity, noise emission, and boating behavior, we classified 11 vessels as disturbing and an additional 12 as potentially disturbing for the seabirds. One colony facing population decline was located closest to the main boating route and experienced 2-5 times more disturbances than the other (increasing) colony, suggesting a negative impact of marine traffic. Our study shows that underwater acoustics can be a useful method to quantify above-water disturbances of seabird colonies.
Incidental bycatch is a well-known challenge in gillnet fisheries throughout the world, and the fishery for North Atlantic lumpfish Cyclopterus lumpus roe is no exception. In Greenland, the fishery was Marine Stewardship Council-certified in 2015 but has pending conditions related to bycatch quantification, enforcement and mitigation strategies. To improve this situation and to assess the potential impact of bycatch, we collected independent on-board observer data on non-target fish and seabirds over 2 seasons (2019 and 2021). We recorded 6 fish species, but the only species constituting >1% of the lumpfish landings was the spotted wolffish Anarhichas minor. The bycatch of fish likely had little impact on the involved fish stocks. We recorded 4 seabird species, of which common eider Somateria mollissima was most common. When extrapolated to the entire West Greenland lumpfish fishery, the estimated bycatch of common eider was considerably higher in 2019 (19938; 95% CI: 3486-59661) than in 2021 (9802: 1260-29940) due to a longer fishing season in 2019. On average, for 2019 and 2021, the bycatch was modelled to reduce the growth potential for the West Greenland winter population by 51%. In comparison, the current hunting level (16538 birds yr-1) reduced the growth potential by 30%. The larger impact of bycatch was mainly due to a larger proportion of adults and females being targeted. The common eider bycatch impacts mainly the breeding population in Canada and Southwest Greenland and less so in Northwest Greenland. As mitigation, we recommend temporal closures of the fishery unless modified gillnets, which markedly reduce bycatch, become available.
Density-dependent prey depletion around breeding colonies has long been considered an important factor controlling the population dynamics of colonial animals.1-4 Ashmole proposed that as seabird colony size increases, intraspecific competition leads to declines in reproductive success, as breeding adults must spend more time and energy to find prey farther from the colony.1 Seabird colony size often varies over several orders of magnitude within the same species and can include millions of individuals per colony.5,6 As such, colony size likely plays an important role in determining the individual behavior of its members and how the colony interacts with the surrounding environment.6 Using tracking data from murres (Uria spp.), the world's most densely breeding seabirds, we show that the distribution of foraging-trip distances scales to colony size0.33 during the chick-rearing stage, consistent with Ashmole's halo theory.1,2 This pattern occurred across colonies varying in size over three orders of magnitude and distributed throughout the North Atlantic region. The strong relationship between colony size and foraging range means that the foraging areas of some colonial species can be estimated from colony sizes, which is more practical to measure over a large geographic scale. Two-thirds of the North Atlantic murre population breed at the 16 largest colonies; by extrapolating the predicted foraging ranges to sites without tracking data, we show that only two of these large colonies have significant coverage as marine protected areas. Our results are an important example of how theoretical models, in this case, Ashmole's version of central-place-foraging theory, can be applied to inform conservation and management in colonial breeding species.
Since the last Arctic Monitoring and Assessment Programme (AMAP) effort to review biological effects of mercury (Hg) on Arctic biota in 2011 and 2018, there has been a considerable number of new Arctic bird studies. This review article provides contemporary Hg exposure and potential health risk for 36 Arctic seabird and shorebird species, representing a larger portion of the Arctic than during previous AMAP assessments now also including parts of the Russian Arctic. To assess risk to birds, we used Hg toxicity benchmarks established for blood and converted to egg, liver, and feather tissues. Several Arctic seabird populations showed Hg concentrations that exceeded toxicity benchmarks, with 50 % of individual birds exceeding the "no adverse health effect" level. In particular, 5 % of all studied birds were considered to be at moderate or higher risk to Hg toxicity. However, most seabirds (95 %) were generally at lower risk to Hg toxicity. The highest Hg contamination was observed in seabirds breeding in the western Atlantic and Pacific Oceans. Most Arctic shorebirds exhibited low Hg concentrations, with approximately 45 % of individuals categorized at no risk, 2.5 % at high risk category, and no individual at severe risk. Although the majority Arctic-breeding seabirds and shorebirds appeared at lower risk to Hg toxicity, recent studies have reported deleterious effects of Hg on some pituitary hormones, genotoxicity, and reproductive performance. Adult survival appeared unaffected by Hg exposure, although long-term banding studies incorporating Hg are still limited. Although Hg contamination across the Arctic is considered low for most bird species, Hg in combination with other stressors, including other contaminants, diseases, parasites, and climate change, may still cause adverse effects. Future investigations on the global impact of Hg on Arctic birds should be conducted within a multi-stressor framework. This information helps to address Article 22 (Effectiveness Evaluation) of the Minamata Convention on Mercury as a global pollutant.
The conservation of migratory marine species, including pelagic seabirds, is challenging because their movements span vast distances frequently beyond national jurisdictions. Here, we aim to identify important aggregations of seabirds in the North Atlantic to inform ongoing regional conservation efforts. Using tracking, phenology, and population data, we mapped the abundance and diversity of 21 seabird species. This revealed a major hotspot associated with a discrete area of the subpolar frontal zone, used annually by 2.9–5 million seabirds from ≥56 colonies in the Atlantic: the first time this magnitude of seabird concentrations has been documented in the high seas. The hotspot is temporally stable and amenable to site‐based conservation and is under consideration as a marine protected area by the OSPAR Commission. Protection could help mitigate current and future threats facing species in the area. Overall, our approach provides an exemplar data‐driven pathway for future conservation efforts on the high seas.
Understanding spatiotemporal variation in vital rates and population growth rates is a central aim of population ecology, and is critical to conservation of migratory species where different populations may spend the non-breeding season in sometimes widely separated areas. However, estimating those parameters and identifying the underlying drivers of variation for species migrating to remote areas is challenging. The thick-billed murre Uria lomvia is a colonial seabird with a pan-Arctic distribution. Previous studies have documented large-scale spatial variation in population growth in the species’ Atlantic range, with eastern populations declining and western populations being stable, and suggested that this variation was linked to conditions in the wintering areas. We analyse variation in breeding success and adult survival from colonies throughout the Atlantic range, construct region-specific stochastic population models, and compare model predictions to observed colony growth rates. Breeding success was uniformly high in Greenland and Iceland, lower and more variable in Spitsbergen, Bjørnøya and Hudson Bay, and low at Jan Mayen. Adult survival was average or high in all colonies during the 2010s. Observed colony growth rates were lower than modelled growth rates for most populations. This suggests that pre-breeding survival or breeding propensity must have been lower than assumed in many of these populations. Low pre-breeding survival could be linked to poor feeding conditions in fall and winter, possibly influenced by large-scale oceanographic variation. Our results support the idea that seabird populations not subject to breeder mortality due to anthropogenic causes are regulated mainly through variation in pre-breeding survival.
Gunshots are a worldwide source of anthropogenic disturbance, and knowledge about the potential effect on wildlife is central for conservation and sustainable management of affected species. This study contributes novel insight of the response behaviour and effect of gunshot disturbance on one of the most culturally important seabird species of the Arctic, the thick-billed murre (Brünnich's guillemot, Uria lomvia). We studied effects of gunshot disturbance at breeding colonies to explore the distance where effects started, the difference in the behavioural response between presumed disturbed colonies (e.g. by traffic and hunting) and largely undisturbed colonies, and among plots with varying seabird densities. We carried out two different types of controlled gunshot experiments – one measuring the distance at which murres first reacted to gunshots (flight initiation distance, ‘FID') and one measuring the proportion of murres remaining in the colony during and after repeated gunshots (20 gunshots within 78 min). FID varied from 0.5 to 5 km. The proportion of murres remaining in the plot after the repeated gunshots ranged from 0.44 to 0.8. Mainly murres not attending offspring took off when disturbed by gunshots, but occasionally also birds incubating their egg or brooding their chick took off. We found that density of seabirds (murres and black-legged kittiwakes Rissa tridactyla) at the plot area scale was the best explaining factor for FID, and to some extent also for the effect of repeated gunshots, both for disturbed and undisturbed colonies. Murres in denser colonies reacted earlier (had larger FIDs) and somewhat more strongly (higher proportions fleeing) to gunshots than murres in less densely populated colonies. FID clearly showed that the current legislation in Greenland regulating gunshots near breeding colonies is insufficient. We provide some recommendations for improved management of a popular game species under pressure.