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To monitor the rapid changes occurring in Arctic ecosystems and predict their direction, basic information about the current number and structure of the main components of these systems is necessary. Using boat-based surveys, we studied the numbers and distribution of seabirds foraging in Hornsund (SW Spitsbergen) during three summer seasons. The average number of seabirds foraging concurrently in the whole fjord was estimated at 28,000. Little Auks Alle alle were the most numerous, followed by Northern Fulmars Fulmarus glacialis, Brünnich’s Guillemots Uria lomvia and Black-legged Kittiwakes Rissa tridactyla. The pelagic zone was exploited by some 75% of the birds. Their density was the highest (> 400 ind. km−2) in the tidewater glacier bays, where kittiwakes were predominant, and the lowest in the coastal glacier bays. The seabirds in Hornsund daily consumed c. 12.7 tons of food, i.e. c. 0.2% of the summer mesozooplankton and fish standing stocks available in the fjord. This food consisted primarily of copepods, amphipods and molluscs (c. 70%), whereas fish made up

Using GPS-tracked individuals, we compared foraging ecology and reproductive output of a High-Arctic zooplanktivorous seabird, the little auk Alle alle, between three years differing in environmental conditions (sea surface temperature). Despite contrasting environmental conditions, average foraging fights distance and duration were generally similar in all studied years. Also, in all years foraging locations visited by the little auk parents during short trips (ST, for chick provisioning) were significantly closer to the colony compared to those visited during long trips (LTs, mainly for adults’ self-maintenance). Nevertheless, we also found some differences in the little auk foraging behaviour: duration of LTs was the longest in the coldest year suggesting more time for resting for adults compared to warmer years. Besides, birds foraged closer to the colony and in significantly colder water in the coldest year. Interestingly, these differences did not affect chick diet: in all the years, the energy content of food loads was similar, with the Arctic copepod, Calanus glacialis copepodite stage V being the most preferred prey item (>73% of items by number and >67% by energy content). Also chick survival was similar in all the study years. However, when examining chicks growth rate we found that their peak body mass was lower in warmer years suggesting that overall conditions in the two warm years were less favourable. While our results, demonstrate a great foraging flexibility by little auks, they also point out their vulnerability to changing environmental conditions.

Here, we model current and future distribution of a foraging Arctic endemic species, the little auk (Alle alle), a small zooplanktivorous Arctic seabird. We characterized environmental conditions [sea depth, sea surface temperature (SST), marginal sea ice zone (MIZ)] at foraging positions of GPS-tracked individuals from three breeding colonies in Svalbard: one located at the southern rim of the Arctic zone (hereafter ‘boreo-Arctic’) and two in the high-Arctic zone on Spitsbergen (‘high-Arctic’). The birds from one ‘high-Arctic’ colony, influenced by cold Arctic water, foraged in the shallow shelf zone near the colony. The birds from remaining colonies foraged in a wider range of depths, in a higher SST zone (‘boreo-Arctic’) or in the productive but distant MIZ (second ‘high-Arctic’ colony). Given this flexible foraging behaviour, little auks may be temporarily resilient to moderate climate changes. However, our fuzzy logic models of future distribution under scenarios of 1 °C and 2 °C SST increase predict losses of suitable foraging habitat for the majority of little auk colonies studied. Over longer time scales negative consequences of global warming are inevitable. The actual response of little auks to future environmental conditions will depend on the range of their plasticity and pace of ecosystem changes.

Summary Two high latitude fjords of Spitsbergen (Hornsund 77°N and Kongsfjorden 79°N) are regarded as being highly productive (70 g and 50 gC m −2 year −1 ) and having organic-rich sediments. Hornsund has more organic matter in its sediments (8%), nearly half of it of terrestrial origin, while most of that in Kongsfjorden (5%) comes from fresh, marine sources (microplankton). Analysis of the carbon sources in both fjords shows that a major difference is the much larger seabird population in Hornsund-dominated with over 100 thousands pairs of plankton feeding little auks in Hornsund versus 2 thousand pairs in Kongsfjorden, and marine food consumption estimated as 5573 tonnes of carbon in Hornsund, versus 3047 tonnes in Kongsfjorden during one month of chick feeding period. Seabird colonies supply rich ornithogenic tundra (595 tonnes of C, as against only 266 tonnes of C in the Kongsfjorden tundra). No much of the terrestrial carbon, flushed out or wind-blown to the fjord, is consumed on the seabed – a state of affairs that is reflected by the low metabolic activity of bacteria and benthos and the lower benthic biomass in Hornsund than in Kongsfjorden.

Different phenological responses to climate changes by species representing preys and predators may lead to mismatch between functionally dependent components of an ecosystem, with important effects on its structure and functioning. Here, we investigate within-season variation in zooplankton availability, chick diet composition and breeding performance of a small planktivorous seabird, the little auk (Alle alle) in two large colonies in Hornsund and Magdalenefjorden, Spitsbergen, differing in synchrony of breeding (11-day vs. 22-day hatching period, respectively). Assuming similar zooplankton phenology and existing differences in duration of the little auk breeding period, we expected lower availability of the preferred food in the less synchronized colony in Magdalenefjorden and in consequence a negative effect on nestling body mass and survival. We found that in both colonies Calanus glacialis (copepodite stage CV) was the most important prey item in the chick diet making up 68–87 % of the biomass and energy of all prey items. The only exception was the end of the chick-rearing period in Magdalenefjorden, when contribution of this prey item was significantly lower (24–26 %). Thus, late breeders in Magdalenefjorden were apparently mismatched regarding C. glacialis CV availability. However, the hatching date did not affect birds fitness (reproductive output and chick pre-fledging mass) significantly. Results of our study indicate that little auks breeding on Spitsbergen can respond to a wide range of environmental conditions and prey availabilities through the plasticity of their foraging behaviour, which may help them to maintain their optimum fitness level in changing and unpredictable environments.

The diversity of cyanobacteria and algae from various microhabitats in Spitsbergen is comparatively well known. However, the relationships between environmental factors and the structure of microflora communities remain largely unclear. This study was conducted in Hornsund Bay, which exhibits large variability in the physicochemical characteristics of habitats, particularly with regard to the availability of nitrogen and phosphorus. This variability, to a large degree, is caused by seabird colonies, which fertilize nutrient-poor terrestrial ecosystems near their nesting areas. The large variations in ecological conditions and vegetation types in the study area aid assessment of habitats representing different combinations of factors potentially influencing the formation of cyanobacterial and algal assemblages. The aim of this study was to examine the influence of physicochemical parameters on the taxonomic composition and diversity of green algae and cyanobacteria (particularly the coccoid, oscillatorialean, and heterocystous taxa). The study encompassed two groups of habitats – soil surface habitats and water-saturated habitats, both characterized by diverse influences of seabird colonies, vegetation cover, and moisture. Our results showed that taxonomic diversity and composition of cyanobacteria and algae were mainly influenced by P–PO43−, N–NH4+ and Ca2+ (soil surface habitats), and NO3−, as well as moisture (index of wetness) and pH (water-saturated habitats). The variability of these physicochemical properties was largely due to the variability of the seabird colony influence. Taken together, our findings aid in understanding the processes of formation of phycoflora assemblages in Arctic tundra.

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.

It is well known that polar bears feed on vegetation. Here, I report novel observations of polar bears grazing on polar scurvy grass (Cochlearia groenlandica) at the foot of a large seabird colony on a cliff on Spitsbergen, Svalbard, in the summers of 2005, 2006, 2009, 2014 and 2015. Why they choose such energy-costly climbing to feed on plants is not clear. One possibility is that they may be suffering from vitamin C deficiency and are searching for this particular plant, which has a high level of this vitamin. Another, but not exclusive reason, is that vegetation containing scurvy grass is abundant enough to be efficiently grazed by such unspecialized plant-eaters as polar bears only on such relatively inaccessible, steep slopes below seabird colonies. Most of the lowland and gently sloping tundra areas in Svalbard are overgrazed by geese and reindeer, the populations of which have increased considerably as a consequence of climate amelioration. Large seabird colonies are known to attract animals from different trophic levels, but this is the first description of their attractiveness to polar bears as grazing areas.

Observed large-scale changes in climate and oceanography, which are especially pronounced in the Arctic, represent profound challenges for upper trophic predators. Knowledge about the extent to which marine predators are able to adjust to environmental variability is essential in order to assess the impact of changing oceanic conditions on the Arctic ecosystem. The main goal of this study was to investigate the impact of oceanographic conditions that varied on an inter-annual and inter-colony basis on the foraging and breeding strategy of a zooplanktivorous alcid, the little auk Alle alle. The study was conducted in two little auk breeding colonies in Spitsbergen (Hornsund and Magdalenefjord) located in different hydrographical regimes in two contrasting seasons (2009 and 2010). This article provides a comprehensive study linking a number of variables describing foraging ground quality, measured directly at sea, with the birds' foraging strategy, parental efforts, body condition and survival of ne...

In the resource-limited Arctic environment, vegetation developing near seabird colonies is exception- ally luxuriant. Nevertheless, there are very few detailed quantitative studies of any specific plant species responses to ornithogenic manuring. Therefore, we studied variability of polar scurvygrass Cochlearia groenlandica individual biomass and leaf width along a seabird influenced gradient determining environmental conditions for vegetation in south-west Spitsbergen. We found seabird colony effect being a paramount factor responsible for augmented growth of C. groenlandica. The species predominated close to the colony and reached the highest mean values of individual biomass (1.4 g) and leaf width (26.6 mm) 10 m below the colony. Its abundance and size declined towards the coast. Both C. groenlandica individual traits signifi- cantly decreased with distance from the colony, soil water and organic matter content and increased with guano deposition, soil d 15 N, conductivity, acidity and nitrate, phosphate and potassium ion content. Our study supports the hypothesis that seabirds have fundamental importance for vegetation growth in poor Arctic environment. Highly plastic species such as C. groenlandica may be a useful instrument in detecting habitat condition changes, for instance resulting from climate change.

Prolonged chasing of an adult reindeer (Rangifer tarandus) by a polar bear (Ursus maritimus) was observed both on land and in the sea, in Magdalenefjorden, northwest Spitsbergen. Polar bears were also observed catching black guillemot (Cepphus grylle) in the sea in northwest Spitsbergen and feeding on chicks in the arctic tern (Sterna paradisea) colony in Hornsund, southwest Spitsbergen. While feeding on seabird species is unsurprising, the prolonged chasing of adult reindeer is unusual for polar bear hunting behaviour. The few documented cases of polar bear hunting reindeer consist of either surprising and killing resting/sleeping prey or stalking and a short rapid chase of the reindeer. Our observations describe new feeding habits of polar bears that may be in response to decreasing seal availability due to shrinking sea-ice cover in the Arctic.

The role of seabirds as sea-land biovectors of nutrients is well documented. However, no studies have examined whether and how colonial seabirds that differ in diet may influence terrestrial vegetation. Therefore, the purpose of the study was to describe and compare plant communities located in the vicinity of the two most common types of seabird colonies in Arctic, occupied by piscivorous or planktivorous species. Within 46 plots arranged in four transects in the vicinity of planktivorous (little auk, Alle alle) and piscivorous colonies (mixed colony of Brunnich’s guillemot, Uria lomvia, and black-legged kittiwake, Rissa tridactyla) we measured the following: guano deposition, physical and chemical characteristics of soil, total nitrogen and its stable isotope signatures in soil and plants, ground vegetation cover of vascular plants and mosses, and the occurrence of lichens, algae and cyanobacteria. Using LINKTREE analysis, we distinguished five plant communities, which reflected declining influence along a birds fertilization gradient measured as guano deposition. SIMPROOF test revealed that these communities differed significantly in species composition, with the differences related to total soil nitrogen content and δ15N, distinctive levels of phosphates, potassium and nitrates, and physical soil properties, i.e., pH, conductivity and moisture. The communities were also clearly distinguished by distance from the bird colony. The two colony types promoted development of specific plant communities: the immediate vicinity of the planktivorous colony characterized by a Deschampsia alpina–Cerastium arcticum community while under the piscivorous colony a Cochlearia groenlandica–Poa alpina community was present. Despite the similar size of the colonies and similar magnitude of guano input, differences between ornithogenic communities were connected mostly to phosphate content in the soil. Our results show that the guano input from seabirds which have different diets can affect High Arctic vegetation in specific and more complex ways than previously realized.

The crucial role of seabirds in the enrichment of nutrient-poor polar terrestrial ecosystem is well-known. However, no studies have examined the potentially different impacts associated with piscivorous and planktivorous bird colonies on the surrounding tundra soils. Therefore, we compared guano deposition and physical and chemical parameters of soil near two large seabird colonies, one of planktivorous little auks (Alle alle) and the other comprising piscivorous Brunnich’s guillemots (Uria lomvia) and kittiwakes (Rissa tridactyla). The two colonies generated similar levels of guano deposition, with the intensity of deposition decreasing away from the colony. Guano deposition adjacent to both colonies was considerably higher than that in control areas. The increased guano supply around colonies significantly enhanced soil conductivity, nitrogen (NO3−, NH4+), potassium (K+), and phosphate (PO43−) ion concentrations and led to reduced pH values. Guano deposition explained 84 % (piscivorous colony) and 67 % (planktivorous colony) of the total variation in the tested soil parameters. Planktivore and piscivore colonies affected adjacent tundra in different ways. The phosphate content and pH value of soil influenced by piscivores were significantly higher than values measured in planktivore-influenced soil. The gradient of guano deposition and associated ion content in the soil decreased more rapidly with distance from the piscivore colony. Climate-induced changes in populations of planktivorous and piscivorous seabirds are expected in the study region and may therefore have substantial consequential effects on Arctic terrestrial ecosystems.

Despite a widespread recognition of the role of seabird colonies in the fertilization of nutrient-poor polar terrestrial ecosystems, qualitative and quantitative data documenting any consequential influence on soil invertebrate communities are still lacking. Therefore, we studied community structure and abundance of springtails (Collembola) in ornithogenic tundra near two large seabird colonies in Hornsund, south-west Spitsbergen. We found considerably (5–20×) higher densities and biomass of Collembola in the vicinities of both colonies (the effect extending up to ca. 50 m from the colony edge) than in comparable control areas of tundra not influenced by allochtonous nutrient input. The most common springtails observed in the seabird-influenced areas were Folsomia quadrioculata, Hypogastrura viatica and Megaphorura arctica. The latter species appeared the most resistant to ornithogenic nutrient input and was found commonly closest to the bird colonies. Collembolan abundance decreased with increasing distance from the seabird colonies. However, relationships between collembolan density and specific physicochemical soil parameters and vegetation characteristics were weak. The most important factors were the cover of the nitrophilous green alga Prasiola crispa, total plant biomass and soil solution conductivity, all of which were correlated with distance from the colony and estimated amount by guano deposition. Community composition and abundance of springtails showed no evidence of being influenced of seabird diet, with no differences apparent between communities found in ornithogenic tundra developing in the vicinity of planktivorous and piscivorous seabird colonies. The study provides confirmation of the influence of marine nutrient input by seabirds on soil microfaunal communities.

Many arctic terrestrial ecosystems suffer from a permanent deficiency of nutrients. Marine birds that forage at sea and breed on land can transport organic matter from the sea to land, and thus help to initiate and sustain terrestrial ecosystems. This organic matter initiates the emergence of local tundra communities, increasing primary and secondary production and species diversity. Climate change will influence ocean circulation and the hydrologic regime, which will consequently lead to a restructuring of zooplankton communities between cold arctic waters, with a dominance of large zooplankton species, and Atlantic waters in which small species predominate. The dominance of large zooplankton favours plankton-eating seabirds, such as the little auk (Alle alle), while the presence of small zooplankton redirects the food chain to plankton-eating fish, up through to fish-eating birds (e.g., guillemots Uria sp.). Thus, in regions where the two water masses compete for dominance, such as in the Barents Sea, plankton-eating birds should dominate the avifauna in cold periods and recess in warmer periods, when fish-eaters should prevail. Therefore under future anthropogenic climate scenarios, there could be serious consequences for the structure and functioning of the terrestrial part of arctic ecosystems, due in part to changes in the arctic marine avifauna. Large colonies of plankton-eating little auks are located on mild mountain slopes, usually a few kilometres from the shore, whereas colonies of fish-eating guillemots are situated on rocky cliffs at the coast. The impact of guillemots on the terrestrial ecosystems is therefore much smaller than for little auks because of the rapid washing-out to sea of the guano deposited on the seabird cliffs. These characteristics of seabird nesting sites dramatically limit the range of occurrence of ornithogenic soils, and the accompanying flora and fauna, to locations where talus-breeding species occur. As a result of climate warming favoring the increase of ichthyiofagous cliffnesting seabirds, we can expect that large areas of ornithogenic tundra around the colonies of plankton-eating seabirds situated far from the sea may disappear, while areas of tundra in the vicinity of cliffs inhabited by fish-eating seabirds, with low total production and supporting few large herbivores, will likely increase, but only imperceptibly. This may lead to habitat fragmentation with negative consequences for populations of tundra-dependent birds and mammals, and the possibility of a substantial decrease in biodiversity of tundra plant and animal communities. r 2007 Elsevier Ltd. All rights reserved.

Seabirds are among the most important vectors transferring biogenic compounds from the sea onto land in the polar regions and, consequently, influencing the properties of soil and vegetation. We studied the influence of bird colonies (Adelie penguin Pygoscelis adeliae, gentoo penguin P. papua and giant petrels Macronectes giganteus) on soil properties and plant communities on King George Island, Maritime Antarctic. We designated seven transects, each starting from the colony edge and running to a natural boundary feature, which were divided into contiguous sample plots where we identified specific plant taxa (Prasiola crispa, Deschampsia antarctica, Colobanthus quitensis, Usnea sp.), as well as hydrophilous and xerophilous ecological groups of mosses. Based on percentage contributions of each of these taxa, we distinguished six distinct vegetation zones along the transects, in which we measured physical (moisture, conductivity and pH) and chemical (NO3 −, NO2 −, NH4 +, K+ and PO4 3− content) soil parameters. Our study confirmed that, with increasing distance from bird colonies, the concentration of nutrients and soil conductivity decreased, while pH increased. The vegetation zones were clearly related to this gradient of seabird colony influence and occurred in the same sequence for all three bird species examined, although the largest colony of Adelie penguins had the strongest effect on vegetation. Similarly, the physical and chemical soil properties did not differ significantly between the colonies.

Variation in body size among subpopulations of the same species may reflect phenotypic or genetic responses to environmental gradients or geographical distance. Here, we examine geographical variation in the body size of the dovekie (Alle alle), the most numerous high-Arctic seabird. Locations of dovekie breeding sites are largely restricted to the high-Arctic zone of the Atlantic. We compared wing length, head-bill length, body mass, and a body size index of 1,076 birds from nine main colonies spanning a large part of the breeding range of the species. Results suggest morphological variation across the studied populations of dovekies, with a longitudinal increase in body size from west to east. The smallest birds breed in the western part of the population (Greenland and Jan Mayen), middle-sized individuals on Svalbard, and the largest birds (A. a. polaris subspecies) breed in the eastern part of the studied area, Franz Josef Land. Environmental (air temperature, wind speed, and sea surface temperature) and geographical (intercolonial distance) parameters were analyzed to explore potential mechanisms driving differences in body size. The body size of birds increased significantly with decreasing air temperature, but only when the two subspecies were considered. We did not find a relationship between sea surface temperature and body size of birds. Also, no close relationship was revealed between birds' body size and the geographical distance between colonies. Whether the body size variation of dovekie can be explained by phenotypic plasticity in response to environmental conditions in wintering areas or a pattern of distance-independent gene flow between colonies remains to be explored.

International audience; The timing of breeding is a life-history trait that can greatly affect fitness, because successful reproduction depends on the match between the food requirements for raising young and the seasonal peak in food availability. We analysed phenology (hatch dates) in relation to climate change for 2 seabird species breeding in the high-Arctic, little auks Alle alle and black-legged kittiwakes Rissa tridactyla, for the periods 1963–2008 and 1970–2008, respectively. We show that spring climate has changed during the study period, with a strong increase in both air temperature (TEMP) and sea surface temperature (SST) and a decrease in sea ice concentration. Little auks showed a trend for earlier breeding over the study period, while kittiwakes showed a non-significant trend for later breeding, demonstrating different phenological responses in these 2 species. Little auks and kittiwakes adjusted their timing of breeding to different environmental signals. Spring TEMP was the best predictor of little auk phenology, with a significant negative effect. Spring SST was the strongest predictor of kittiwake phenology, with a non-significant negative effect. Spring sea ice concentration and the North Atlantic Oscillation (NAO) winter index had a low relative variable importance. Furthermore, in kittiwakes, years with late breeding were associated with low clutch size and mean annual breeding success, indicating poor investment and food availability. This study identifies some spring environmental factors important for regulating the timing of breeding in the high-Arctic, most likely through effects on snow cover limiting access to nest sites and the development of the polar marine food web. It remains to be investigated whether environmental factors are reliable predictors of marine prey phenology, and whether the decision to start breeding is constrained by food availability.

During the second half of August 1992 polar bears were observed feeding intensively in the little auk Alle alle colony on Rubini Rock, Hooker Island, Frans Josef Land. They dug out nests eating eggs, chicks and adult birds. Their activity results in both worsening the breeding conditions and decreasing the breeding success of the little auks. The polar bear has not earlier been known as a little auk predator.

Glucocorticoids participate in the control of whole body homoeostasis and an organism’s response to stress. Corticosterone, which is the principal glucocorticoid in birds, has been shown to increase in response to different energetic demands and perturbations that individuals have to cope with. In this study, a non-invasive method to examine the corticosterone secretion by measuring faecal corticosterone metabolites (FCM) has been established for an Arctic seabird, the Little Auk (Alle alle). A group-specific immunoassay was successfully validated for adults and chicks using an adrenocorticotropic challenge test. Then, FCM levels were investigated under different energetic and physiological demands, determined by weather conditions, week of chick rearing in adults, and age in chicks. The amount of rainfall had no effect on FCM levels in adults, whereas it negatively affected FCM levels in chicks. There was no variation in FCM concentrations among weeks of chick rearing in adults. In chicks, the FCM levels increased with age. Moreover, chicks with higher FCM levels had lower body mass and fledged later than chicks with lower FCM levels. This study demonstrates that environmental stress such as poor weather conditions can trigger significant changes in corticosterone levels in seabird chicks. Furthermore, the results indicate that corticosterone may be involved in the physiological and behavioural adjustments necessary for successful fledging and post-fledging survival.