Sex differences in foraging ecology of a zooplanktivorous little auk Alle alle during the pre-laying period: insights from remote sensing and animal-tracking.

BACKGROUND: Energy and time allocation in seabirds differ between consecutive stages of breeding given various requirements of particular phases of the reproductive period. Theses allocations may also be sex-specific considering differential energetic or nutritional requirements of males and females and/or sexual segregation in foraging niches and/or areas. In this study we investigated the foraging ecology of an Arctic, zooplanktivorous seabird, the little auk Alle alle during the pre-laying period using remote sensing of the environment and GPS-TDR loggers deployed on birds. We compared foraging trips range and habitats of birds with other stages of the breeding period and between sexes. RESULTS: We found that little auks during the pre-laying period foraged exclusively in cold sea surface temperature zones (with temperatures < 5 ºC) but in various sea depth zones. They dived to similar depths ranging from -4.0 to -10.9 m, exploring various thermal microhabitats (with mean temperatures values ranging from 2.2 °C in Shelf sea depth zone to 5.9 °C in Deep sea depth zone). The majority of foraging trips and dives characteristics were similar to subsequent phases of breeding. However, home ranges during the pre-laying trips were wider compared to the incubation period. As expected, females exhibited wider foraging niches compared to males (wider range of sea surface temperature and sea depth in foraging locations), which could be explained by sex specific energetic and/or nutritional requirements (females producing an egg). We also delineated local foraging areas important for little auks during their whole breeding season. Protection of these areas is crucial for sustaining the local marine biodiversity. CONCLUSIONS: We found that little auks females during the pre-laying period explored wider foraging niches compared to males. These differences may be attributed to sex-specific nutritional or/and energetical constraints at this stage of breeding. The results of this study also emphasize the importance of shelf Arctic-type water masses as the foraging areas for little auks during successive stages of breeding.

Experimental reduction in blood oxygen-carrying capacity alters foraging behaviour in a colonial waterbird.

Oxidative metabolism is a key component of organismal physiology and it is primarily determined by aerobic capacity, which depends on the capacity of blood to carry oxygen. However, experimental manipulations of blood oxygen-carrying capacity are rarely implemented to test ecophysiological hypotheses in vertebrate populations. Here, we combined an experimental manipulation of blood oxygen-carrying capacity with GPS tracking to test whether suboptimal (reduced) haematological performance affects foraging behaviour in a colonial waterbird, the black-headed gull, Chroicocephalus ridibundus. First, a validation of phenylhydrazine (PHZ) treatment in gulls revealed a 9-18% reduction in haematocrit and blood haemoglobin concentration (via oxidative denaturation and haemolysis of erythrocytes). Then, GPS tracking of experimental (PHZ-treated) and control (saline-treated) gulls during the incubation period provided no support for reduced or suspended engagement in energetically costly activities (long-distance foraging trips) by experimental birds. Instead, we found evidence for fine-scale alterations in foraging behaviour of PHZ-treated individuals, which resulted in fewer foraging trips per unit time, but trips that were longer in duration and distance compared with those of control birds. This suggests reduced foraging performance of experimental birds (e.g. lower capacity to find and collect food during trips) or evasion of social competition, although no differences in the total investment in foraging may also suggest compensatory physiological responses to haemolytic anaemia. Our study contributes to a better understanding of the physio-ecological nexus in non-diving colonial avian species. Whether behavioural effects of reduced aerobic capacity have any implications for gull condition and reproductive performance should be the subject of further investigation.

Identification of Factors Affecting Environmental Contamination Represented by Post-Hatching Eggshells of a Common Colonial Waterbird with Usage of Artificial Neural Networks.

Artificial Neural Networks are used to find the influence of habitat types on the quality of the environment expressed by the concentrations of toxic and harmful elements in avian tissue. The main habitat types were described according to the Corine Land Cover CLC2012 model. Eggs of free-living species of a colonial waterbird, the grey heron Ardea cinerea, were used as a biological data storing media for biomonitoring. For modeling purposes, pollution indices expressing the sum of the concentration of harmful and toxic elements (multi-contamination rank index) and indices for single elements were created. In the case of all the examined indices apart from Cd, the generated topologies were a multi-layer perceptron (MLP) with 1 hidden layer. Interestingly, in the case of Cd, the generated optimal topology was a network with a radial basis function (RBF). The data analysis showed that the increase in environmental pollution was mainly influenced by human industrial activity. The increase in Hg, Cd, and Pb content correlated mainly with the increase in the areas characterized by human activity (industrial, commercial, and transport units) in the vicinity of a grey heron breeding colony. The decrease in the above elements was conditioned by relative areas of farmland and inland waters. Pollution with Fe, Mn, Zn, and As was associated mainly with areas affected by industrial activities. As the location variable did not affect the quality of the obtained networks, it was removed from the models making them more universal.

Contrasting Spatial and Seasonal Trends of Methylmercury Exposure Pathways of Arctic Seabirds: Combination of Large-Scale Tracking and Stable Isotopic Approaches.

Despite the limited direct anthropogenic mercury (Hg) inputs in the circumpolar Arctic, elevated concentrations of methylmercury (MeHg) are accumulated in Arctic marine biota. However, the MeHg production and bioaccumulation pathways in these ecosystems have not been completely unraveled. We measured Hg concentrations and stable isotope ratios of Hg, carbon, and nitrogen in the feathers and blood of geolocator-tracked little auk Alle alle from five Arctic breeding colonies. The wide-range spatial mobility and tissue-specific Hg integration times of this planktivorous seabird allowed the exploration of their spatial (wintering quarters/breeding grounds) and seasonal (nonbreeding/breeding periods) MeHg exposures. An east-to-west increase of head feather Hg concentrations (1.74-3.48 µg·g-1) was accompanied by significant spatial trends of Hg isotope (particularly Δ199Hg: 0.96-1.13‰) and carbon isotope (δ13C: -20.6 to -19.4‰) ratios. These trends suggest a distinct mixing/proportion of MeHg sources between western North Atlantic and eastern Arctic regions. Higher Δ199Hg values (+0.4‰) in northern colonies indicate an accumulation of more photochemically impacted MeHg, supporting shallow MeHg production and bioaccumulation in high Arctic waters. The combination of seabird tissue isotopic analysis and spatial tracking helps in tracing the MeHg sources at various spatio-temporal scales.

Sex- and breeding stage-specific hormonal stress response of seabird parents.

Changes in corticosterone (CORT) and prolactin (PRL) levels are thought to provide complementary information on parental decisions in birds in the context of stressful situations. However, these endocrine mechanisms have yet to be fully elucidated, appearing to vary among avian species without any clear pattern. Here, we examined CORT and PRL stress responses in a small Arctic seabird, the little auk (Alle alle). We analysed the levels of these hormones (baseline, and stress response, i.e. the change in the baseline in response to stress) with respect to the breeding phase (mid incubation and mid chick rearing) and the sex of the birds. Baseline CORT concentrations were similar during both breeding phases but baseline PRL levels were higher during incubation than chick rearing. The CORT and PRL stress responses were stronger during incubation than chick rearing (although with respect to CORT the effect was only marginally significant). There were also some sex-specific baseline levels and stress responses for both hormones (during the incubation period males compared to females exhibited higher CORT stress response and lower baseline PRL; during the chick rearing period males exhibited higher PRL stress response). Our results suggest that in the case of the little auk, both the incubation and the chick rearing periods may represent similar levels of physiological stress. However, the birds may be more sensitive to stress during incubation than during chick rearing, possibly because of inter-phase differences in predation pressure. The sex differences suggest differential exposure of males and females to stressors.

Flexibility of foraging strategies of the great skua Stercorarius skua breeding in the largest colony in the Barents Sea region.

BACKGROUND: Foraging strategies of seabird species often vary considerably between and within individuals. This variability is influenced by a multitude of factors including age, sex, stage of annual life cycle, reproductive status, individual specialization and environmental conditions. RESULTS: Using GPS-loggers, we investigated factors affecting foraging flight characteristics (total duration, maximal range, total distance covered) of great skuas Stercorarius skua of known sex breeding on Bjørnøya, Svalbard, the largest colony in the Barents Sea region. We examined influence of sex (females are larger than males), phase of breeding (incubation, chick-rearing), reproductive status (breeders, failed breeders) and bird ID (they are known for individual foraging specialization). Our analyses revealed that only bird ID affected foraging flight characteristics significantly, indicating a high degree of plasticity regardless of sex, reproductive status or phase of breeding. We recognized three main groups of individuals: 1) those preying mainly on other seabirds in the breeding colonies (6%), 2) those foraging at sea (76%) and kleptoparasiting other seabirds and/or foraging on fish and/or offal discarded by fishing vessels, and 3) those alternating between preying on other seabirds in breeding colonies and foraging at sea (18%). Despite marked size sexual dimorphism, we found no apparent sex differences in flight characteristics. Birds after egg- or chick-loss and thus not constrained as central foragers did not modify their foraging flights. CONCLUSIONS: Great skuas breeding on Bjørnøya displayed a high degree of plasticity regardless of sex, reproductive status or phase of breeding. We recognized groups of individuals regularly preying in the seabird colonies, foraging at sea, and alternating between both strategies. This suggests foraging specialization of some individuals.

Factors Affecting Element Concentrations in Eggshells of Three Sympatrically Nesting Waterbirds in Northern Poland.

Avian eggshells are convenient samples in biomonitoring studies, because they are easily accessible, especially from colonially or semicolonially breeding birds. In the present study, concentrations of 17 elements, including heavy metals and essential elements in post-hatch eggshells, were compared among three species of waterbirds of differing strategies for gaining reserves for egg production and diet: mallard, Anas platyrhynchos (ML, a capital breeder, mainly herbivorous), common tern, Sterna hirundo (CT, an income breeder, piscivorous) and black-headed gull, Chroicocephalus ridibundus (BHG, mixed strategy, omnivorous) and breeding sympatrically in three sites in North Poland. Analyses revealed that Fe, Zn, and Cu levels differed the most in the studied species, which may be explained by various contributions of fish, aquatic plants, and soil invertebrates in their diets. Generally, the studied species' eggshells accumulated amounts of elements comparable to those reported for other waterbirds without putting the growth and development of the embryo at risk. The only exception was very high levels of Cr in ML and CT, which may be explained by their foraging on aquatic organisms in waterbodies polluted by this element. Intersite differences in eggshell concentrations of Ni, Sr, Hg and Cr in CT (an income breeder) may be explained by the influence of local pollution sources (small factories, polluted river).

Factors determining presence of passerines breeding within White Stork Ciconia ciconia nests.

Nests of White Stork Ciconia ciconia are commonly used by various passerines as nesting sites. In this study, we investigated factors determining presence and number of pairs of species breeding within White Stork nests in an extensive farmland in NE Poland. In 133 (57%) out of 233 White Stork nests, we found at least one breeding pair of passerine bird. These were from three species: House Sparrows Passer domesticus (68% of 133 nests with co-breeding), Tree Sparrows Passer montanus (65%), and Starlings Sturnus vulgaris (30%). The probability of breeding passerines within White Stork nests increased with increasing nest thickness, and was significantly higher in currently occupied nests. Sparrows were more likely to breed in White Stork nests located on electricity poles, situated closer to settlements and surrounded mainly by arable fields where meadows were not prevalent. In this paper, we show that White Stork nests are favorable nesting sites for passerines, as they are well insulated and provide an anti-predatory shield.

Trace element concentrations in livers of Common Buzzards Buteo buteo from eastern Poland.

In this study, our aim was to determine the common sources of origin of 18 elements in the livers of Common Buzzards collected during the breeding season in an extensive agricultural landscape in south-east Poland with respect to age (adults and immatures) and sex (males and females). In all 34 specimens collected, the element concentrations followed the pattern of S > Na > Fe > Mg > Zn > Si > Cu > Mn > Ba > Se > B > Pb > Hg > Cd > Cr > Ni > Sr > V. Among the heavy metals examined, only the concentration of Pb was relatively high. Given the prevalence of farmland in the studied area (and the wide use of fertilizers), common use of lead-hunting ammunition and moderate concentration of Pb in fertilizers, the indirect influence of hunting ammunition ingested with food or as gastroliths was apparently responsible for the elevated levels of Pb in the livers of Common Buzzards. In our study, no significant sex-related differences were detected in the hepatic concentrations of any element. However, a significant age effect was observed for three elements, which had elevated levels in adults (Hg) and immature birds (B, Pb), and a significant age x sex interaction was found for S and Fe. These results might be explained by the importance of these elements in bone growth in immature birds (B), variable strategies of foraging between adults and immature birds (Pb), and possible intersex differences in the immature cohort in response to the presence of lead (S, Fe).

Foraging Habitat Availability and the Non-Fish Diet Composition of the Grey Heron (Ardea cinerea) at Two Spatial Scales.

Habitat structure on foraging ground is one of the crucial factors determining diet diversity in colonially breeding avian predators. Quantifying the habitat and diet composition at different spatial scales (regional and local inter-colonial) can help provide understanding of the drivers of diet composition. In this study, we examined the composition of the non-fish diet of an opportunistic predator, the Grey Heron (Ardea cinerea), based on pellets. We compared pellets from 21 colonies in two different regions of Poland that differing in composition of foraging habitats. Multivariate statistical techniques were used to analyze the relationship between diet and habitat compositions in a 20 km radius around these colonies. Significant inter-regional and inter-colony differences in Grey Heron diet and habitat composition were detected in foraging areas. However, some prey were present in the diet only from one of the regions. Around the heronries with a predominance of farmland habitats, the European water vole (Arvicola amphibius) and terrestrial invertebrates were present in relatively low frequencies, and the striped field mouse (Apodemus agrarius) was present in high frequencies. Voles (Microtus sp.) were more frequent in colonies with prevalence of non-irrigated arable land around the colony, in contrast to the bank vole (Myodes glareolus), which was less common in heronries with a higher contribution of this habitat type. Remains of aquatic invertebrates were less abundant in colonies surrounded by extensive forests. The results of our research indicate the opportunistic character of the non-fish part of the diet of the Grey Heron adapted to the local foraging habitat and prey availability.

The first evidence of alloparental feeding in a crevice-nesting seabird, the little auk.

An alloparent is an individual that cares for a young individual, but it is not its genetic parent. This behaviour is known in many species of animals, but some groups are still underreported. Here, we documented, in camera footage, the alloparental feeding of two chicks of the little auk, a crevice-nesting seabird. This is the first evidence of this behaviour in the little auk despite similar monitoring undertaken between 2016 and 2022 and the second record for a crevice/burrow-nesting seabird. We compared chicks that were fed by an alloparent to other chicks from the same year and explored reasons for the behaviour in the context of seabird breeding biology.

Concentration of metals and metalloids in livers of birds of various foraging guilds collected during the autumn migration period in Poland.

During migration, birds explore various habitats at stopover sites that differ in food resources and contamination levels. In this study, hepatic concentrations of 21 elements (metals and metalloids) in 11 species of birds, representing various foraging habitats (such as aquatic, aquatic/terrestrial, and terrestrial) and migration modes (migratory and sedentary) representing various foraging guilds (omnivores, piscivores, and molluscivores), were analyzed. The samples (N = 84) were collected during the autumn migration period in Poland. The concentrations of elements determined in this study exhibited high inter-species variability, reflecting the diversity in contamination levels depending on food resources used by specific bird groups. Many of the investigated individuals from different species showed exceeded levels of subclinical toxicity and moderate clinical poisoning due to Cd and Hg. Higher concentrations of As, Hg, and Ba and lower V concentrations were found in migratory birds as compared to sedentary birds. Species foraging in terrestrial habitat had different concentrations of some elements compared to aquatic and aquatic/terrestrial species. Some specific inter-species differences in hepatic elemental concentrations were found. Differences in elemental concentrations among various groups can primarily be attributed to their foraging guilds, with certain elements, particularly As, V, and Hg, playing a significant role in the dissimilarity of elemental concentrations between foraging habitat groups and migratory mode groups. The data collected confirmed the limited ability of As to enter ecosystem pathways. The results of this study contribute to understanding the year-round exposure of migratory birds to environmental contamination, which can have carry-over effects on their performance in wintering and breeding grounds.

Decline in the West Greenland population of a zooplanktivorous seabird, the little auk Alle alle.

The warming of the Arctic had lead to a diverse range of impacts on local biota, including northward shifts of some species range. Here, we report past and present distribution and abundance of an Arctic zooplanktivorous seabird, the little auk Alle alle in West Greenland south of 74° N, and examine the changes in sea surface temperature (SST) and sea ice concentration (SIC) in the birds foraging areas in 1850-2007. We estimated the little auk population in the studied region to be 5,200 pairs in the 1930s, 6,000-6,500 pairs in the 1940-1970s and 70-80 pairs by the 2000s. We found that periods with increased SST and reduced SIC, especially in the last few decades, coincided with little auk population declines. Besides, years with little auk presence in breeding sites were characterized by either low SST and low to moderate SIC or higher SST but moderate to high SIC. Observed contraction of the breeding range and a decrease in abundance of the little auk may be attributed to more complex climate-driven changes in the marine ecosystem at finer spatial and temporal scales and/or cannot be easily detected given the coarseness of data used. It is possible that the population in this region has never been very numerous being subjected to local impacts such as disease, bycatch, predation, etc. The climate warming that is currently being observed, along with corresponding shifts in zooplankton communities, may lead to extirpation of the studied little auk populations.

Coordination of parental performance is breeding phase-dependent in the Dovekie (Alle alle), a pelagic Arctic seabird.

Currently, parental care is becoming increasingly perceived as male and female cooperation, instead of being primarily shaped by sexual conflict. Most studies examining cooperating performance consider coordination of parental activities, and doing so focuses on a short time-window including only one stage of breeding (i.e., incubation or chick rearing period). Here, we considered the cooperation of breeding partners, investigating the coordination of parental care in a long-lived seabird species with long and extensive biparental care, the Dovekie (or Little Auk), Alle alle, and looked at the issue throughout the breeding season. Previous studies on this species revealed coordinated chick provisioning, but parental coordination during incubation remains unstudied. Using video recordings collected over the course of two breeding seasons, we tested whether coordination was subject to small-scale changes within each stage and whether there was a relationship between coordination levels across the two stages. We found that the level of parental coordination is overall high and increases during the incubation period but decreases through the chick rearing phases. There were some inter-annual differences in the coordination level both at the incubation and chick rearing stages. We also found some dependency between the coordination during the incubation and chick rearing periods. All these results suggest that coordination is not a fixed behavior but breeding-phase dependent. The present study thus provides insights into how parental care and parents' cooperation is shaped by brood needs and conditions. It also highlights a relationship between coordination levels during chick rearing and incubation periods, suggesting some extent of temporal dependence in coordination of parental performance within the breeding season.

Exploring avian exposure to parent polycyclic aromatic hydrocarbons (PAHs): Using the common eider Somateria mollissima in a global context.

Compared to other organic contaminants, birds are rarely studied for their exposure to polycyclic aromatic hydrocarbons (PAHs), mainly due to their effective metabolization of parent PAHs. However, as some studies suggest, exposure to PAHs may result in adverse health effects including decreased survival, especially following oil spills. In the present study, we analyzed samples from a sea duck, the common eider Somateria mollissima including feathers, preen oil, blood, liver and bile, to evaluate whether non- lethally collected samples could be reliably used for avian biomonitoring strategies. Phenanthrene was the only individual PAH detected across sample types, with the highest concentration found in preen gland and the lowest in blood. Significant differences in concentrations were observed between bile vs preen gland and liver vs preen gland, while for most compounds neither blood nor feathers showed detectable levels of parent PAHs. Therefore, the utility of those sample types for PAH exposure assessment may be limited and should be interpreted with caution, moreover as several physiological factors may affect them. Additionally, we also provide a comparison with the available literature to review current avian PAH exposure assessment and outline future research focused needs.

Diversity of major histocompatibility complex of II B gene and mate choice in a monogamous and long-lived seabird, the Little Auk (Alle alle).

The major histocompatibility complex (MHC) plays a key role in the adaptive immune system of vertebrates, and is known to influence mate choice in many species. In birds, the MHC has been extensively examined but mainly in galliforms and passerines while other taxa that represent specific ecological and evolutionary life-histories, like seabirds, are underexamined. Here, we characterized diversity of MHC Class II B exon 2 in a colonial pelagic seabird, the Little Auk (or Dovekie Alle alle). We further examined whether MHC variation could be maintained through balancing selection and disassortative mating. We found high polymorphism at the genotyped MHC fragment, characterizing 99 distinct alleles across 140 individuals from three populations. The alleles frequencies exhibited a similar skewed distribution in both sexes, with the four most commonly occurring alleles representing approximately 35% of allelic variation. The results of a Bayesian site-by-site selection analysis suggest evidence of balancing selection and no direct evidence for MHC-dependent disassortative mating preferences in the Little Auk. The latter result might be attributed to the high overall polymorphism of the examined fragment, which itself may be maintained by the large population size of the species.

Behavioural and hormonal stress responses during chick rearing do not predict brood desertion by female in a small Arctic seabird.

We examined behavioural and hormonal stress responses in a small seabird (little auk, Alle alle), which exhibits a transition from biparental to male-only care towards the end of the nesting period, in order to understand the mechanisms underlying this parental strategy. We hypothesized that the male staying with the chick should be less sensitive to stressors. As such the male might offer the offspring more efficient protection during the fledging period than the female. We tested this hypothesis by observing male and female behaviour in a neophobia test. We also measured the birds' baseline and stress-induced levels of corticosterone and prolactin using the standardized capture-and-restraint protocol. Both sexes respond rapidly to foreign objects, delaying the entry time to the nest with food, consuming the food load, and/or temporarily abandoning feeding. However, we did not find any differences between the sexes in the frequency of each behaviour or in the time of the first reaction to the experimental treatment. Level of both corticosterone and prolactin increased after the experimental treatment. However, we did not find sex differences in baseline and stress-induced hormone levels. The results indicate that the males are as much sensitive to the stress situation as the females. Thus, the pattern of male and female behavioural and hormonal responses to stress does not predict their behaviour at the final breeding stage.

Differential autumn migration of the aquatic warbler Acrocephalus paludicola.

Relatively little attention has been paid to sex differences in the migration of birds in autumn. We studied the autumn migration strategy of molecularly sexed males and females in the globally threatened aquatic warbler Acrocephalus paludicola. We captured 176 birds at a stopover site in the Loire estuary at Donges, France. The median date for the passage of adults was 8 days earlier in males than females, although the timing of migration in first-year males and females was similar. This indicates that males, who are without parental duties, can start their migration earlier than females and first-year birds. Adults were significantly heavier than immature birds but did not have higher fat scores. In both age categories, more males (two to three times more) were captured. However, various factors (including tape-luring) can affect observed sex ratio.

Mine or my neighbours' offspring: an experimental study on parental discrimination of offspring in a colonial seabird, the little auk Alle alle.

Individual recognition (IR) abilities may result from various ecological and naturally selected features of a species. Complex IR mechanisms should develop when the risk of misidentification of a chick is high. For colonial seabirds, the ability to identify their own brood is crucial to ensure parental fitness. Vocalizations seem to be a key component of most parent-offspring interactions, although few studies have assessed the interindividual differences in seabird chick calls and their potential usage in IR. The little auk (Alle alle), which breeds in dense colonies, constitutes a perfect model for testing IR. In this study, we (1) examined chick calls at different stages of the nesting period, and (2) cross-fostered chicks to examine the rate of acceptance/nonacceptance of chicks by parents. We found significant interindividual differences in chick begging and fledging calls. Surprisingly, all cross-fostered chicks in our experiments were accepted by their foster parents, and male parents were as equally likely to accept cross-fostered chicks as females, even though the sexes would be expected to differ in offspring recognition due to different postfledging interactions with the chick. The revealed individuality of chick calls suggests the potential for chick vocal recognition in the studied species, but parent birds may disregard the individual characteristics enabling chick discrimination. This may take place as long as the chick is found in the nest because of the high likelihood that the chick present there is the focal one. However, IR during and after fledging requires further study. Studying the complexity of IR mechanisms is important for better understanding various avian social relationships and interactions.

Molecular tools prove little auks from Svalbard are extremely selective for Calanus glacialis even when exposed to Atlantification.

Two Calanus species, C. glacialis and C. finmarchicus, due to different life strategies and environmental preferences act as an ecological indicators of Arctic Atlantification. Their high lipid content makes them important food source for higher trophic levels of Arctic ecosystems including the most abundant Northern Hemisphere's seabird, the little auk (Alle alle). Recent studies indicate a critical need for the use of molecular methods to reliably identify these two sympatric Calanus species. We performed genetic and morphology-based identification of 2600 Calanus individuals collected in little auks foraging grounds and diet in summer seasons 2019-2021 in regions of Svalbard with varying levels of Atlantification. Genetic identification proved that 40% of Calanus individuals were wrongly classified as C. finmarchicus according to morphology-based identification in both types of samples. The diet of little auks consisted almost entirely of C. glacialis even in more Atlantified regions. Due to the substantial bias in morphology-based identification, we expect that the scale of the northern expansion of boreal C. finmarchicus may have been largely overestimated and that higher costs for birds exposed to Atlantification could be mostly driven by a decrease in the size of C. glacialis rather than by shift from C. glacialis to C. finmarchicus.