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Climate warming is rapidly altering marine ecosystems towards a more temperate state on the European side of the Arctic. However, this “Atlantification” has rarely been confirmed, as long-term datasets on Arctic marine organisms are scarce. We present a 19-year time series (1982–2016) of diet samples from black-legged kittiwakes as an indicator of the changes in a high Arctic marine ecosystem (Kongsfjorden, Svalbard). Our results highlight a shift from Arctic prey dominance until 2006 to a more mixed diet with high contribution of Atlantic fishes. Capelin, an Atlantic species, dominated the diet composition in 2007, marking a shift in the food web. The occurrence of polar cod, a key Arctic fish species, positively correlated with sea ice index, whereas Atlantic species demonstrated the opposite correlation indicating that the diet shift was likely connected with recent climate warming. Kittiwakes, which gather available fish and zooplankton near the sea surface to feed their chicks, can act as messengers of ecosystem change. Changes in their diet reveal that the Kongsfjord system has drifted in an Atlantic direction over the last decade.

Summary Reproduction is energetically expensive, and daily energy expenditure (DEE) often peaks during the period of rearing young. The ‘potentiation’ hypothesis predicts that high DEE needs to be sustained by a corresponding up‐regulation of metabolic machinery; thus, a concomitant increase in the resting metabolic rate (RMR) is expected. Alternatively, the ‘compensation’ hypothesis predicts that DEE and RMR are regulated independently and animals may maintain low RMR to maximize the energy available for reproduction. This might particularly be the case if DEE was limited, for example, by extrinsic food supply or intrinsic physiological factors. We tested these hypotheses in free‐living seabirds by manipulating their energy demands (experimentally reduced or increased brood size) and supplies (providing supplemental food), and simultaneously measuring their DEE and RMR (by the doubly labelled water method and an indirect hormonal proxy, respectively). In support of the ‘compensation’ hypothesis, metabolic rates were adjusted independently and in opposite directions with an increase in DEE and a decrease in the hormonal proxy for RMR in individuals rearing young compared to birds with removed broods. Energy expenditure of unfed birds with chicks appeared to be limited as experimental brood enlargement did not cause an increase in DEE. Supplemental feeding did not allow DEE to exceed this apparent limitation. We propose that a reduction in the resting metabolism is a strategy to increase allocation of energy to reproduction when DEE is constrained and that this constraint is unlikely to be related to food supply. Lay Summary

1. The ability to store energy endogenously is an important ecological mechanism that allows animals to buffer predictable and unpredictable variation in allostatic load. The secretion of glucocorticoids, which reflects changes in allostatic load, is suggested to play a major role in the adjustment of endogenous stores to these varying conditions. 2. Although crucially important, the relationship between allostatic load and energy stores remains largely unexplored. Two contrasting hypotheses describe how stores may be adjusted: animals may use low allostatic loads to increase stores to a maximum possible ('fat and fit'), or they can attain a lean physique due to fitness advantages of a low body mass ('lean and fit'). 3. We compiled observational and experimental data available for a long-lived seabird to examine the relationship between glucocorticoids and stored energy at two life history stages (incubation and chick-rearing). Data were collected across multiple years and colonies in the North Pacific, thereby reflecting the wide range of environmental conditions birds' encounter in the marine environment. During experimental manipulations, allostatic load was minimized by supplementing food to free-living birds. 4. We found that the relationship between allostatic load and energy stores was clearly curvilinear at both life history stages. Observational data suggested that energy stores remained relatively stable under low allostatic load and decreased under high loads. Experimental data showed that birds did not maximize energy stores under favourable conditions but maintained energy stores below a physiologically attainable level. 5. Energy stores remained consistently lower during chick-rearing compared to incubation across the wide range of variations in allostatic load suggesting that stage-specific trade-offs limit the accumulation of energy during favourable environmental conditions. Secretion of glucocorticoids did not appear to mediate this shift in energy stores between the life history stages. 6. Overall, results of this study support the 'lean and fit' hypothesis. We conclude that increased energy stores may not necessarily reflect better environmental conditions experienced by individuals or predict their higher fitness. A major advantage of adopting a lean physique when environmental conditions allow may be the avoidance of additional energetic costs for moving a heavy body. In breeding seabirds, this advantage may be more important during chick-rearing. In the focal species, the secretion of glucocorticoids might be involved in regulation of energy stores within a life history stage but does not appear to mediate an adaptive shift in energy stores between the incubating and chick-rearing stages of reproduction.

Global warming, combined with an increasing influence of Atlantic Waters in the European Arctic, are causing a so-called Atlantification of the Arctic. This phenomenon is affecting the plankton biomass and communities with potential consequences for the upper trophic levels. Using long-term data (2005-2020) from a high Arctic zooplanktivorous seabird, the little auk (Alle alle), we tested the hypothesis that the Atlantification affects its diet, body condition and demography. We based our study on data collected in three fjords in West Spitsbergen, Svalbard, characterized by distinct oceanographic conditions. In all three fjords, we found a positive relationship between the inflow of Atlantic Waters and the proportion of Atlantic prey, notably of the copepod Calanus finmarchicus, in the little auk chick diet. A high proportion of Atlantic prey was negatively associated with adult body mass (though the effect size was small) and with chick survival (only in one fjord where chick survival until 21 days was available). We also found a negative and marginally significant effect of the average proportion of Atlantic prey in the chick diet on chick growth rate (data were available for one fjord only). Our results suggest that there are fitness costs for the little auk associated with the Atlantification of West Spitsbergen fjords. These costs seem especially pronounced during the late phase of the chick rearing period, when the energetic needs of the chicks are the highest. Consequently, even if little auks can partly adapt their foraging behaviour to changing environmental conditions, they are negatively affected by the ongoing changes in the Arctic marine ecosystems. These results stress the importance of long-term monitoring data in the Arctic to improve our understanding of the ongoing Atlantification and highlight the relevance of using seabirds as indicators of environmental change.

Thyroid hormones (TH) are known to stimulate in vitro oxygen consumption of tissues in mammals and birds. Hence, in many laboratory studies a positive relationship between TH concentrations and basal metabolic rate (BMR) has been demonstrated whereas evidence from species in the wild is scarce. Even though basal and field metabolic rates (FMR) are often thought to be intrinsically linked it is still unknown whether a relationship between TH and FMR exists. Here we determine the relationship between the primary thyroid hormone triiodothyronine (T3) with both BMR and FMR in a wild bird species, the black-legged kittiwake (Rissa tridactyla). As predicted we found a strong and positive relationship between plasma concentrations of T3 and both BMR and mass-independent BMR with coefficients of determination ranging from 0.36 to 0.60. In contrast there was no association of T3 levels with either whole-body or mass-independent FMR (R(2) = 0.06 and 0.02, respectively). In accordance with in vitro studies our data suggests that TH play an important role in modulating BMR and may serve as a proxy for basal metabolism in wild birds. However, the lack of a relationship between TH and FMR indicates that levels of physical activity in kittiwakes are largely independent of TH concentrations and support recent studies that cast doubt on a direct linkage between BMR and FMR.

In birds with facultative brood reduction, survival of the junior chick is thought to be regulated primarily by food availability. In black-legged kittiwakes (Rissa tridactyla) where parents and chicks are provided with unlimited access to supplemental food during the breeding season, brood reduction still occurs and varies interannually. Survival of the junior chick is therefore affected by factors in addition to the amount of food directly available to them. Maternally deposited yolk androgens affect competitive dynamics within a brood, and may be one of the mechanisms by which mothers mediate brood reduction in response to a suite of environmental and physiological cues. The goal of this study was to determine whether food supplementation during the pre-lay period affected patterns of yolk androgen deposition in free-living kittiwakes in two years (2003 and 2004) that varied in natural food availability. Chick survival was measured concurrently in other nests where eggs were not collected. In both years, supplemental feeding increased female investment in eggs by increasing egg mass. First-laid (\"A\") eggs were heavier but contained less testosterone and androstenedione than second-laid (\"B\") eggs across years and treatments. Yolk testosterone was higher in 2003 (the year with higher B chick survival) across treatments. The difference in yolk testosterone levels between eggs within a clutch varied among years and treatments such that it was relatively small when B chick experienced the lowest and the highest survival probabilities, and increased with intermediate B chick survival probabilities. The magnitude of testosterone asymmetry in a clutch may allow females to optimize fitness by either predisposing a brood for reduction or facilitating survival of younger chicks.

1. Parental investment strategies are considered to represent a trade-off between the benefits of investment in current offspring and costs to future reproduction. Due to their high residual reproductive value, long-lived organisms are predicted to be more reluctant to increase parental effort. 2. We tested the hypothesis that breeding little auks (Alie alle) have a fixed level of reproductive investment, and thus reduce parental effort when costs associated with reproduction increase. 3. To test this hypothesis we experimentally increased the flight costs of breeding little auks via feather clipping. In 2005 we examined changes in the condition of manipulated parents, of the mates of manipulated parents, and of their chick as direct measures of change in parental resource allocation between self-maintenance and current reproduction. In 2007 we increased sample sizes to determine whether there was a physiological cost (elevated corticosterone, CORT) associated with the manipulation. 4. We found that:(i) clipped birds and their mates lost more body mass than controls, but there was no difference in mass loss between members of a pair; (ii) clipped birds had higher CORT levels than control birds; (iii) there were no inter-annual differences in body mass and CORT levels between clipped individuals and their mates at recapture, and (iv) chicks with a clipped parent had lower peak and fledging mass, and higher CORT levels than control chicks in both years. 5. Contrary to our hypothesis, the reduction in body mass of partners to clipped birds suggests that little auks can increase parental effort to some extent. Nonetheless, the lower fledging mass and higher CORT of chicks with a clipped parent indicates provisioning rates may not have been fully maintained. 6. As predicted by life-history theory, there may be a threshold to the additional reproductive costs breeders will accept, with parents prioritizing self-maintenance over increased provisioning effort when foraging costs become too high.

1. The rate at which free-living animals can expend energy is limited but the causes of this limitation are not well understood. Theoretically, energy expenditure may be intrinsically limited by physiological properties of the animal constraining its capacity to process energy. Alternatively, the limitation could be set extrinsically by the amount of energy available in the environment or by a fitness trade-off in terms of reduced future survival associated with elevated metabolism. 2. We measured daily energy expenditure (DEE) using the doubly labelled water method in chick-rearing black-legged kittiwakes (Rissa tridactyla) at a study site close to the northern limit of their breeding range over 5 years. We measured breeding success, foraging trip duration and diet composition as proxies of resource availability during these years and estimated the probability of parent kittiwakes to return to the colony in relation to their energy expenditure in order to determine whether kittiwakes adjust their DEE in response to variation in prey availability and whether elevated DEE is associated with a decrease in adult survival. 3. We found that DEE was strikingly similar across all five study years. There was no evidence that energy expenditure was limited by resource availability that varied considerably among study years. Furthermore, there was no evidence of a negative effect of DEE on adult return rate, which does not support the hypothesis of a survival cost connected to elevated energy expenditure. 4. The additional lack of variation in DEE with respect to ambient temperature, brood size or between sexes suggests that kittiwakes at a time of peak energy demands may operate close to an intrinsic metabolic ceiling independent of extrinsic factors.

Quantifying patterns of genetic diversity and differentiation among populations of Arctic birds is fundamental for understanding past and ongoing population processes in the Arctic. However, the genetic differentiation of many important Arctic species remains uninvestigated. Here, phylogeography and population genetics were examined in the worldwide population of a small seabird, the little auk (dovekie, Alle alle )—the most numerous avian species of the Arctic ecosystem. Blood samples or feathers were collected from 328 little auks (325 from the nominate subspecies and 3 from the A. a. polaris ) in nine main breeding aggregations in the northern Atlantic and one location from the Pacific Ocean. The mtDNA haplotypes of the two subspecies were not segregated into separate groups. Also, no genetic structure was found within the nominate race based on microsatellite markers. The level of genetic differentiation among populations was low yet significant (mean F ST  = 0.005). Some pairwise F ST comparisons revealed significant differences, including those involving the most distant Pacific colony as well as among some Atlantic populations. Weak population differentiation following the model of isolation by distance in the little auk is similar to the patterns reported in other high-Arctic bird species, indicating that a lack of distinct genetic structure is a common phenomenon in the Arctic avifauna.

The number of offshore wind farms in Europe and elsewhere has substantially increased in recent years. This rapid development has raised concerns about potential impacts on marine wildlife, particularly on seabirds, as these can be negatively affected through collision and displacement. While collision risk has been the focus of a number of studies, information about displacement of seabirds is scarce. Here we present data from an extensive survey program that aimed at determining the effects on seabirds of the first German offshore wind farm, ‘alpha ventus’. Data were collected by line transect surveys during the first 3 yr of operation. We found significant displacement of 5 species with 75–92% lower abundance inside compared to outside the wind farm. For 3 species, the response distance to the outermost turbines was estimated to exceed 1 km. Two gull species were attracted to the wind farm site. Our results and a review of the available literature revealed good agreement with respect to the sign of the response (avoidance vs. attraction) but considerable differences in the strength of the response and the spatial extent of the disturbance outside the footprint of wind farms. While it seems unlikely that small-scale displacement by single wind farms would have an impact at the population level, the extent of the proposed development of offshore wind energy warrants further research into cumulative effects and their biological significance for seabird populations.