A major feeding ground for cetaceans and seabirds in the south-western Greenland Sea

In the frame of our long-term study of the distribution of seabirds and marine mammals in polar seas, observers from this team participated in the European Arctic expeditions of icebreaking RV Polarstern during summer 2008. The main aims were to obtain more information on the mechanisms underlying the at-sea distribution of the ??higher trophic levels?? and to detect possible temporal and spatial changes in numbers, especially in function of climatic changes and decreasing ice coverage in the Arctic. In total, 1,175 half-an-hour transect counts were devoted to seabird and marine mammal distribution in the Greenland and Norwegian Seas, from 14 June to 15 August, 2008. A major feeding ground for cetaceans and seabirds was detected in the south-western Greenland Sea and the Denmark Strait in the Polar Front between Polar and Arctic Water masses. For the main cetacean and seabird species, almost all individuals recorded during the whole expedition were encountered in this limited zone of 33 counts: 135 humpback whales Megaptera novaeangliae and 1,940 common guillemots Uria aalge. Moreover, 4,400 fulmars Fulmarus glacialis light morph were observed in this zone, i.e. close to half of the total. All were actively feeding, making this zone the major feeding ground for the higher trophic levels in the European Arctic Seas and so confirming the great importance of fronts (upwellings) for the biological productivity of the oceans. Such data are essential for a successful conservation and management of the Denmark Strait and adjacent areas.     

Variation in ice conditions has strong effects on the breeding of marine birds at Prince Leopold Island, Nunavut

The maximum extent of sea ice in the northern hemisphere has been contracting for several decades, with implications for all ice-associated biota. To determine how variation in ice conditions affects reproduction in marine birds, we studied the effects of ice conditions on breeding of four species of seabirds over four years at Prince Leopold Island, Nunavut, a colony close to the limits of ice conditions where breeding is feasible. In 2000 and 2003, open water was present close to the colony in June, when the birds began to lay eggs. In 2001 and 2002, the ice edge in June was >200 km to the east of the colony, forcing birds to commute long distances to open water to feed. Egg-laying by thick-billed murres, black-legged kittiwakes and glaucous gulls was delayed and eggs and clutches were smaller in 2001 and 2002. However, northern fulmars laid at the same time in all years, although their incubation shifts were longer in 2001 and 2002 than in 2003. Open water was present close to the colony by the time of hatching in all years. Despite this, nestling survival of northern fulmars, body condition and nestling growth of thick-billed murres and body condition and nestling survival of black-legged kittiwakes were lower in 2001 and 2002 than in –2000 and 2003. All these indicators suggest that feeding conditions in the years of late ice break-up continued to be worse than usual even after open water was available at the colony. Our study suggests that current trends towards earlier ice break-up in the Arctic may be beneficial for marine birds at Prince Leopold Island, at least in the short-term.     

Variable sea‐ice conditions influence trophic dynamics in an Arctic community of marine top predators

Sea‐ice coverage is a key abiotic driver of annual environmental conditions in Arctic marine ecosystems and could be a major factor affecting seabird trophic dynamics. Using stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) in eggs of thick‐billed murres (Uria lomvia), northern fulmars (Fulmarus glacialis), glaucous gulls (Larus hyperboreus), and black‐legged kittiwakes (Rissa tridactyla), we investigated the trophic ecology of prebreeding seabirds nesting at Prince Leopold Island, Nunavut, and its relationship with sea‐ice conditions. The seabird community of Prince Leopold Island had a broader isotopic niche during lower sea‐ice conditions, thus having a more divergent diet, while the opposite was observed during years with more extensive sea‐ice conditions. Species' trophic position was influenced by sea ice; in years of lower sea‐ice concentration, gulls and kittiwakes foraged at higher trophic levels while the opposite was observed for murres and fulmars. For murres and fulmars over a longer time series, there was no evidence of the effect of sea‐ice concentration on species' isotopic niche. Results suggest a high degree of adaptation in populations of high Arctic species that cope with harsh and unpredictable conditions. Such different responses of the community isotopic niche also show that the effect of variable sea‐ice conditions, despite being subtle at the species level, might have larger implications when considering the trophic ecology of the larger seabird community. Species‐specific responses in foraging patterns, in particular trophic position in relation to sea ice, are critical to understanding effects of ecosystem change predicted for a changing climate.     

Diet of black guillemots and northern fulmars breeding beside a High Arctic polynya

In the Arctic, the availability of early-season open water (shore leads, polynyas) is a key factor influencing annual reproduction by marine birds, as these relatively productive locations provide the only sites where migrating and breeding birds can feed. We examined the diet of two marine birds, black guillemots (Cepphus grylle) and northern fulmars (Fulmarus glacialis), feeding in the Hell Gate–Cardigan Strait Polynya in the Canadian High Arctic during the breeding seasons of 1980–1984. There was little overlap in prey items consumed between these two species, except that both relied on similar sympagic amphipods. Guillemots fed principally on benthic prey, and compared to earlier High Arctic studies, they consumed proportionally more benthic fish, mysids and decapods. Fulmars appeared to feed on swarms of pelagic or surface prey, and in this polynya they consumed proportionally fewer fish and more polychaetes than has been reported for fulmars at other High Arctic locations. Given that climate change may alter marine food webs by changing the types and availability of prey, our results set a baseline of dietary information for these predators against which future conditions can be compared.     

Climate as a driver of phenological change in southern seabirds

Seabirds are one of the most threatened groups of birds globally and, overall, their conservation status is deteriorating rapidly. Southern hemisphere countries are over-represented in the number of species of conservation concern yet long-term phenological data on seabirds in the southern hemisphere is limited. A better understanding of the implications of changes in the marine and terrestrial environments to seabird species is required in order to improve their management and conservation status. Here we conducted a meta-analysis of the phenological drivers and trends among southern hemisphere seabirds. Overall there was a general trend towards later phenological events over time (34 % of all data series, N?=?47; 67 % of all significant trends), though this varied by taxa and location. The strongest trends towards later events were for seabirds breeding in Australia, the Laridae (gulls, noddies, terns) and migratory southern polar seabirds. In contrast, earlier phenologies were more often observed for the Spheniscidae (penguins) and for other seabirds breeding in the Antarctic and subantarctic. Phenological changes were most often associated with changes in oceanographic conditions, with sea-ice playing an important role for more southerly species. For some species in some locations, such as the Little Penguin Eudyptula minor in south-eastern Australia, warmer oceans projected under various climate change scenarios are expected to correspond to increased seabird productivity, manifested through earlier breeding, heavier chicks, an increased chance of double brooding, at least in the short-term.     

Ectoparasites of northern fulmars Fulmarus glacialis (Procellariiformes: Procellariidae) from the Canadian Arctic

We studied the prevalence and intensity of infestation of ectoparasites on northern fulmars (Fulmarus glacialis L.) from a breeding colony in Arctic Canada in June–August 2003. No fleas or ticks were found on any fulmars, but three species of chewing lice (Phthiraptera) were recorded: Ischnocera: Perineus nigrolimbatus (Giebel 1874), Ischnocera: Saemundssonia occidentalis (Kellogg 1896), and Amblycera: Ancistrona vagelli (Fabricius 1787). Non-breeding birds had a higher prevalence of lice than breeding birds, and prevalence varied markedly among louse species. Our study is an important baseline for the occurrence of ectoparasites on northern fulmars in the high Arctic, a region undergoing extensive environmental change due to global warming, and an area where parasites are expected to extend ranges or increase in prevalence under changing annual temperature regimes.     

Chlamydiaceae in North Atlantic Seabirds Admitted to a Wildlife Rescue Center in Western France

Birds are the primary hosts of Chlamydia psittaci, a bacterium that can cause avian chlamydiosis in birds and psittacosis in humans. Wild seabirds are frequently admitted to wildlife rescue centers (WRC) at European Atlantic coasts, for example, in connection with oil spills. To investigate the extent of chlamydial shedding by these birds and the resulting risk for animals in care and the medical staff, seabirds from a French WRC were sampled from May 2011 to January 2014. By use of a quantitative PCR (qPCR), 195 seabirds belonging to 4 orders, 5 families and 13 species were examined, of which 18.5% proved to be Chlamydiaceae positive. The highest prevalence of shedders was found in northern gannets (Morus bassanus) (41%), followed by European herring gulls (Larus argentatus) (14%) and common murres (Uria aalge) (7%). Molecular characterization and phylogenetic analysis of qPCR-positive northern gannet samples revealed two variants of a strain closely related to C. psittaci. In European herring gulls and in one common murre, strains showing high sequence similarity to the atypical Chlamydiaceae-like C122 previously found in gulls were detected. Our study shows that seabirds from the northeastern Atlantic Ocean carry several chlamydial organisms, including C. psittaci-related strains. The staff in WRCs should take protective measures, particularly in the case of mass admissions of seabirds.     

Changes in body composition during breeding: Reproductive strategies of three species of seabirds under poor environmental conditions

Seabirds differ dramatically in life history traits and breeding strategies. For example, gulls have short incubation shifts (several hours) and high metabolic rates, auks have medium-length incubation shifts (12-24h) and high metabolic rates, and petrels have long incubation shifts (days) and low metabolic rates. How these different strategies affect the dynamics of body components is poorly known. We compared body, organ and lipid mass changes among three different seabirds (gull: black-legged kittiwake Rissa tridactyla; auk: thick-billed murre Uria lomvia; petrel: northern fulmar Fulmarus glacialis) at Prince Leopold Island, Nunavut, Canada during 2002 (a year with low reproductive success and poor chick growth across all three species). This study is among the first to compare mass and lipid dynamics among different species foraging in the same food web and at similar trophic levels during the same breeding season (same environmental conditions). In fulmars and murres, most of decreases in body mass reflected decreases in lipid mass while in kittiwakes the increase in body mass reflected an increase in lean mass, especially the muscle. The species with the longest fasting endurance (incubation shift length) had the highest percent body lipids during incubation (fulmars: 13.3%, murres: 7.3%, kittiwakes: 6.9%), the highest variability in body lipids, tended to regulate body mass primarily through lipid stores and tended to regulate exercise and digestive organs separately. In contrast, in the species with the highest metabolic rate, all organ systems were adjusted similarly and in relation to body mass, and in a similar manner between incubation (stress due to heavy ice conditions) and chick-rearing (lower stress due to ice-free conditions). In high metabolic rate species, we suggest that organ size varies in response to environmental stress. We conclude that the organ dynamics of seabirds are set by a combination of key life history traits (like incubation shift and metabolic rate) and environmental conditions.     

ANALYSIS OF MECHANISMS OF MICROEVOLUTIONARY CHANGE IN CEPPHUS GUILLEMOTS USING PATTERNS OF CONTROL REGION VARIATION

We surveyed population-level sequence variation in part of the mitochondrial control region for three species including eight subspecies of Cepphus guillemots (Charadriiformes: Alcidae) to test specific predictions about mechanisms of population differentiation. We found that sequences of spectacled guillemots (C. carbo) were more closely related to those of pigeon guillemots (C. columba; both found in the Pacific Ocean) than to those of black guillemots (C. grylle; Arctic and Atlantic Oceans), despite dissimilarities in plumage between spectacled guillemots and the other species. Distributions of species and timing of divergence events suggest that speciation involved allopatric and microallopatric populations isolated by Pleistocene glaciers. Control region sequences were significantly differentiated among populations within species and suggest that gene flow is low; however, populations are probably not in genetic equilibrium, so these results probably reflect historical isolation of colonies. In contrast, phylogenetic relationships among sequences within species were poorly resolved, probably because of a combination of incomplete lineage sorting and contemporary gene flow. Indices of genetic diversity provided no suggestion of recent bottlenecks in most populations, although two populations apparently underwent recent severe bottlenecks. Genetic divergence among populations was not correlated with geographic distance, which argues against isolation by distance. Results of these analyses, combined with breeding distributions and timing of divergence events, suggest that populations diverged during isolation in glacial refugia. Our results are consistent with earlier hypotheses posed by Storer and Udvardy.     

Influence of weather on reproductive success of northern fulmars in the Canadian high Arctic

The northern fulmar (Fulmarus glacialis) is a common seabird of the North Atlantic Ocean, with breeding colonies broadly dispersed between 45°N and 80°N. At higher latitudes, breeding fulmars experience extensive sea-ice and presumably snow and low temperatures which do not affect fulmars in the southern part of the breeding range. We studied the relationship between weather and reproductive success of northern fulmars breeding at two colonies in the Canadian high Arctic. Collectively, hatching success, fledging success, and productivity (chicks fledged per egg laid) were similar between our study and results from colonies located south of the Arctic. However, a larger proportion of fulmars at apparently occupied sites (AOS) in high Arctic colonies appeared to forego egg-laying, resulting in lower proportions of chicks fledged per AOS. Extreme inclement weather was the major factor influencing nesting success, resulting in pulses of egg or chick loss during or immediately following major storms, although the mechanism of effects appeared to differ between the two colonies. For Arctic fulmars, the risks of nest failure due to stochastic, deleterious weather events may be offset by the predictable abundance of food supplies during chick-rearing in Arctic waters.     

An important late summer aggregation of fin whales Balaenoptera physalus,little auks Alle alle and Brünnich’s guillemots Uria lomvia in the eastern Greenland Sea and Fram Strait: influence of hydrographic structures

The distribution at sea of upper trophic levels—seabirds and marine mammals—is depending on their food availability: high concentrations reflect high prey abundance and thus high biological production. Polar marine ecosystems are characterized by low biodiversity and high biological patchiness. The distribution of predators, as a consequence, shows a similar patchiness. During two expeditions of icebreaking RV Polarstern in June–July 2011, biodiversity in the arctic marine zone north of 70°N was very low, with low numbers of species: 20 seabirds, eight cetaceans, five pinnipeds and polar bear. Moreover, a few species accounted for the majority in numbers: four bird species for 95 % of the total of 23,000 seabirds recorded during 700 transect counts: fulmar Fulmarus glacialis, kittiwake Rissa tridactyla, Brünnich’s guillemot Uria lomvia and little auk Alle alle. Among the marine mammals, 250 fin whales Balaenoptera physalus accounted for 80 % of the identified large cetaceans, 270 white-beaked dolphin Lagenorhynchus albirostris for 100 % of the small cetaceans and 180 harp seals Pagophilus groenlandica for 80 % of the identified pinnipeds. Their quantitative distribution was depending on water masses and oceanic fronts, large cetaceans—mainly fin whales—showing an important aggregation on the shelf slope off western Spitsbergen, as well as little auks and Brünnich’s guillemots. So that this zone, shelf slope and front of mixed Arctic/Atlantic Waters, showed unusually high seabird and cetacean concentrations. Seasonal factors possibly influencing their distribution are addressed.     

Cryptic introgression between murre sister species (Uria spp.) in the Pacific low Arctic: frequency, cause, and implications

As southern species undergo northward range expansions, reports of hybridization between temperate and arctic taxa are increasing, which may have important implications for the evolution, conservation, and management of arctic species. The extent of hybridization between temperate common murres (Uria aalge) and arctic thick-billed murres (U. lomvia), seabirds in the family Alcidae, has been the subject of debate. In a previous survey of variation in mitochondrial DNA (mtDNA) in common and thick-billed murres sampled from throughout the North Pacific and low Arctic, 12 of 327 common murres (~4%) were found to possess DNA sequences characteristic of thick-billed murres. In the present study, we surveyed variation in three nuclear introns in 230 common murres and 56 thick-billed murres and report that these putative hybrids carry various combinations of intron alleles from common and thick-billed murres. Analysis using the program STRUCTURE indicated that nine of these individuals possessed high proportions of thick-billed murre intron alleles, two possessed alleles in F1 and F2 proportions, and one individual possessed predominantly common murre intron alleles. We propose that the asymmetric mtDNA introgression we observed is most likely the result of mate choice at mixed colonies based on differences in male mating behaviours. Our results highlight that hybridization between thick-billed and common murres is more prevalent than previously thought, which may have important implications for the conservation and management of arctic-dwelling thick-billed murres as common murres expand northward.     

Interactive effects of body mass changes and species‐specific morphology on flight behavior of chick‐rearing Antarctic fulmarine petrels under diurnal wind patterns

For procellariiform seabirds, wind and morphology are crucial determinants of flight costs and flight speeds. During chick‐rearing, parental seabirds commute frequently to provision their chicks, and their body mass typically changes between outbound and return legs. In Antarctica, the characteristic diurnal katabatic winds, which blow stronger in the mornings, form a natural experimental setup to investigate flight behaviors of commuting seabirds in response to wind conditions. We GPS‐tracked three closely related species of sympatrically breeding Antarctic fulmarine petrels, which differ in wing loading and aspect ratio, and investigated their flight behavior in response to wind and changes in body mass. Such information is critical for understanding how species may respond to climate change. All three species reached higher ground speeds (i.e., the speed over ground) under stronger tailwinds, especially on return legs from foraging. Ground speeds decreased under stronger headwinds. Antarctic petrels (Thalassoica antarctica; intermediate body mass, highest wing loading, and aspect ratio) responded stronger to changes in wind speed and direction than cape petrels (Daption capense; lowest body mass, wing loading, and aspect ratio) or southern fulmars (Fulmarus glacialoides; highest body mass, intermediate wing loading, and aspect ratio). Birds did not adjust their flight direction in relation to wind direction nor the maximum distance from their nests when encountering headwinds on outbound commutes. However, birds appeared to adjust the timing of commutes to benefit from strong katabatic winds as tailwinds on outbound legs and avoid strong katabatic winds as headwinds on return legs. Despite these adaptations to the predictable diurnal wind conditions, birds frequently encountered unfavorably strong headwinds, possibly as a result of weather systems disrupting the katabatics. How the predicted decrease in Antarctic near‐coastal wind speeds over the remainder of the century will affect flight costs and breeding success and ultimately population trajectories remains to be seen.     

The stress hormone corticosterone in a marine top predator reflects short‐term changes in food availability

In many seabird studies, single annual proxies of prey abundance have been used to explain variability in breeding performance, but much more important is probably the timing of prey availability relative to the breeding season when energy demand is at a maximum. Until now, intraseasonal variation in prey availability has been difficult to quantify in seabirds. Using a state‐of‐the‐art ocean drift model of larval cod Gadus morhua, an important constituent of the diet of common guillemots Uria aalge in the southwestern Barents Sea, we were able to show clear, short‐term correlations between food availability and measurements of the stress hormone corticosterone (CORT) in parental guillemots over a 3‐year period (2009–2011). The model allowed the extraction of abundance and size of cod larvae with very high spatial (4 km) and temporal resolutions (1 day) and showed that cod larvae from adjacent northern spawning grounds in Norway were always available near the guillemot breeding colony while those from more distant southerly spawning grounds were less frequent, but larger. The latter arrived in waves whose magnitude and timing, and thus overlap with the guillemot breeding season, varied between years. CORT levels in adult guillemots were lower in birds caught after a week with high frequencies of southern cod larvae. This pattern was restricted to the two years (2009 and 2010) in which southern larvae arrived before the end of the guillemot breeding season. Any such pattern was masked in 2011 by already exceptionally high numbers of cod larvae in the region throughout chick‐rearing period. The findings suggest that CORT levels in breeding birds increase when the arrival of southern sizable larvae does not match the period of peak energy requirements during breeding.     

Population differentiation and evolution in the common guillemot Uria aalge

Common (Uria aalge) and Brünnich's guillemots (U. Iomvia) are colonial seabirds that nest in temperate to arctic oceans throughout the Northern hemisphere. They are very similar in the characteristics of ecology, demography and life history that are thought to determine the extent of differentiation among populations, yet geographic variation in morphology is notably greater in common guillemots. Despite evidence of strong natal philopatry, previous analyses of allozymes and the mitochondrial cytochrome b gene revealed little genetic differentiation among North Atlantic colonies of Brünnich's guillemots. To determine if the more extensive morphological variability in common guillemots reflects greater genetic variability, we sequenced part of the cytochrome b gene for 160 common guillemots from 10 colonies distributed throughout the Northern hemisphere. Genotype frequencies and phylogenetic relationships among genotypes both indicated that Atlantic and Pacific populations are genetically distinct. Genetic divergence among genotypes suggested that differentiation of these populations has resulted from separation by Pleistocene glaciers and the Bering Landbridge, as well as by currently unsuitable breeding habitat in the Arctic Ocean. Cytochrome b genotype frequencies also differed among Atlantic colonies, and appeared to define a cline similar to that described for morphological characters. Analyses of sequence variation suggested that this variation probably results from secondary contact between two refugial populations from the Pleistocene glaciations, rather than from isolation by distance or selection. In contrast, the Atlantic population of Brünnich's guillemots appears to have arisen through recent expansion of a single homogeneous refugial population.     

Guano deposition and nutrient enrichment in the vicinity of planktivorous and piscivorous seabird colonies in Spitsbergen

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 (PO₄ ^3?) 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.     

Seabirds and sandeels: the conflict between exploitation and conservation in the northern North Sea

Sandeels, especiallyAmmodytes marinus are a major component of the diet of many predatory fish, seabirds and seals. The industrial fishery for sandeels is now the largest of the North Sea fisheries. A sandeel fishery in the Shetland area began in 1974 but has recently declined. This change was accompanied by dramatic declines in the breeding success of certain seabirds, particularly Arctic terns and kittiwakes. Current information on seabirds and sandeels in the Shetland area is reviewed and areas where further research is needed, highlighted. The Shetland problem illustrates the difficulties of reconciling conservation and exploitation when fundamental ecological and behavioural knowledge is lacking, and also the need to obtain further information on the ecological impact of industrial fisheries.     

Species-specific population trends detected for penguins,gulls and cormorants over 20 years in sub-Antarctic Fuegian Archipelago

Understanding the dynamics and causes of population trends are essential for seabird conservation. Long-term studies of seabirds at high-latitude (Antarctic, sub-Antarctic and Arctic) regions have shown contrasting species-specific trends in population size in response to climate change and anthropogenic pressures. We have studied for the last 20 years (1992–2012) the population trends of seven seabird species that breed in the Beagle Channel, south-eastern Tierra del Fuego and at Staten Island, a sub-Antarctic region in southern Argentina. The numbers of Magellanic and Gentoo Penguins increased significantly since 1992 (by >15 % year^?1). In comparison, the populations of Imperial Cormorants, Dolphin Gulls and Kelp Gulls increased at slower rates (<5 % year^?1), while the Rock Cormorant population even decreased by 1.3 % year^?1. At Staten Island, the numbers of Rockhopper Penguins decreased by 24 % between the censuses of 1998 and 2010, whereas the population of Magellanic Penguins increased by 227 % during the same period. Over the study period, air and sea-surface temperatures remained stable in our study area, suggesting that the detected population changes are not driven by the climate. This finding contrasts with the detected links between increasing temperature trends and seabird population changes reported from Antarctic and Arctic regions. The level of tourism and size of the permanent human population has increased in the Beagle Channel area during the last 20 years and could be responsible for the increase of gull populations. The seabird species that received the highest number of visitors (Imperial Cormorants and penguin species) seem to be adapted or at least indifferent to pressures exerted by tourism, as their populations increased during the study period. In addition, increasing numbers of seabirds in the area may generally be leading to higher abundances of scavenging species (e.g. gulls).     

Energetic consequences of contrasting winter migratory strategies in a sympatric Arctic seabird duet

At the onset of winter, warm‐blooded animals inhabiting seasonal environments may remain resident and face poorer climatic conditions, or migrate towards more favourable habitats. While the origins and evolution of migratory choices have been extensively studied, their consequences on avian energy balance and winter survival are poorly understood, especially in species difficult to observe such as seabirds. Using miniaturized geolocators, time‐depth recorders and a mechanistic model, we investigated the migratory strategies, the activity levels and the energy expenditure of the closely‐related, sympatrically breeding Brünnich's guillemots Uria lomvia and common guillemots Uria aalge from Bjørnøya, Svalbard. The two guillemot species from this region present contrasting migratory strategies and wintering quarters: Brünnich's guillemots migrate across the North Atlantic to overwinter off southeast Greenland and Faroe Islands, while common guillemots remain resident in the Barents, the Norwegian and the White Seas. Results show that both species display a marked behavioural plasticity to respond to environmental constraint, notably modulating their foraging effort and diving behaviour. Nevertheless, we provide evidence that the migratory strategy adopted by guillemots can have important consequences for their energy balance. Overall energy expenditure estimated for the non‐breeding season is relatively similar between both species, suggesting that both southward migration and high‐arctic winter residency are energetically equivalent and suitable strategies. However, we also demonstrate that the migratory strategy adopted by Brünnich's guillemots allows them to have reduced daily energy expenditures during the challenging winter period. We therefore speculate that ‘resident’ common guillemots are more vulnerable than ‘migrating’ Brünnich's guillemots to harsh winter environmental conditions.     

Determining seabird body condition using nonlethal measures

Energy stores are critical for successful breeding, and longitudinal studies require nonlethal methods to measure energy stores ("body condition"). Nonlethal techniques for measuring energy reserves are seldom verified independently. We compare body mass, size-corrected mass (SCM), plasma lipids, and isotopic dilution with extracted total body lipid content in three seabird species (thick-billed murres Uria lomvia, all four measures; northern fulmars Fulmarus glacialis, three measures; and black-legged kittiwakes Rissa tridactyla, two measures). SCM and body mass were better predictors of total body lipids for the species with high percent lipids (fulmars; R2 = 0.5-0.6) than for the species with low percent lipids (murres and kittiwakes; R2 = 0.2-0.4). The relationship between SCM and percent body lipids, which we argue is often a better measure of condition, was also poor (R2 < 0.2) for species with low lipids. In a literature comparison of 17 bird species, percent lipids was the only predictor of the strength of the relationship between mass and total body lipids; we suggest that SCM be used as an index of energy stores only when lipids exceed 15% of body mass. Across all three species we measured, SCM based on the ordinary least squares regression of mass on the first principal component outperformed other measures. Isotopic dilution was a better predictor of both total body lipids and percent body lipids than were mass, SCM, or plasma lipids in murres. Total body lipids decreased through the breeding season at both sites, while total and neutral plasma lipid concentrations increased at one site but not another, suggesting mobilization of lipid stores for breeding. A literature review showed substantial variation in the reliability of plasma markers, and we recommend isotopic dilution (oxygen-18, plateau) for determination of energy reserves in birds where lipid content is below 15%.