Climate change in the Arctic: Testing the poleward expansion of ticks and tick-borne diseases.

Climate change is most strongly felt in the polar regions of the world, with significant impacts on the species that live there. The arrival of parasites and pathogens from more temperate areas may become a significant problem for these populations, but current observations of parasite presence often lack a historical reference of prior absence. Observations in the high Arctic of the seabird tick Ixodes uriae suggested that this species expanded poleward in the last two decades in relation to climate change. As this tick can have a direct impact on the breeding success of its seabird hosts and vectors several pathogens, including Lyme disease spirochaetes, understanding its invasion dynamics is essential for predicting its impact on polar seabird populations. Here, we use population genetic data and host serology to test the hypothesis that I. uriae recently expanded into Svalbard. Both black-legged kittiwakes (Rissa tridactyla) and thick-billed murres (Uria lomvia) were sampled for ticks and blood in Kongsfjorden, Spitsbergen. Ticks were genotyped using microsatellite markers and population genetic analyses were performed using data from 14 reference populations from across the tick's northern distribution. In contrast to predictions, the Spitsbergen population showed high genetic diversity and significant differentiation from reference populations, suggesting long-term isolation. Host serology also demonstrated a high exposure rate to Lyme disease spirochaetes (Bbsl). Targeted PCR and sequencing confirmed the presence of Borrelia garinii in a Spitsbergen tick, demonstrating the presence of Lyme disease bacteria in the high Arctic for the first time. Taken together, results contradict the notion that I. uriae has recently expanded into the high Arctic. Rather, this tick has likely been present for some time, maintaining relatively high population sizes and an endemic transmission cycle of Bbsl. Close future observations of population infestation/infection rates will now be necessary to relate epidemiological changes to ongoing climate modifications.

Melatonin and corticosterone profiles under polar day in a seabird with sexually opposite activity-rhythms.

The 24 h geophysical light-dark cycle is the main organizer of daily rhythms, scheduling physiology and behavior. This cycle attenuates greatly during the continuous light of summer at polar latitudes, resulting in species-specific and even individual-specific patterns of behavioral rhythmicity, but the physiological mechanisms underlying this variation are poorly understood. To address this knowledge gap and to better understand the roles of the hormones melatonin and corticosterone in rhythmic behavior during this 'polar day', we exploited the behavior of thick-billed murres (Uria lomvia), a charadriiform seabird with sexually opposite ('antiphase') activity-rhythms that have a duration of 24 h. Melatonin concentration in the plasma of inactive males was unexpectedly high around midday and subsequently fell during a sudden decrease in light intensity as the colony became shaded. Corticosterone concentration in plasma did not vary with time of day or activity in either sex. While the reasons for these unusual patterns remain unclear, we propose that a flexible melatonin response and little diel variation of corticosterone may be adaptive in thick-billed murres, and perhaps other polar birds and mammals, by stabilizing glucocorticoids' role of modulating energy storage and mobilization across the diel cycle and facilitating the appropriate reaction to unexpected stimuli experienced across the diel cycle while attending the colony.

Muscle myonuclear domain, but not oxidative stress, decreases with age in a long-lived seabird with high activity costs.

In birds, many physiological parameters appear to remain constant with increasing age, showing no deterioration until 'catastrophic' mortality sets in. Given their high whole-organism metabolic rate and the importance of flight in foraging and predator avoidance, flight muscle deterioration and accumulated oxidative stress and tissue deterioration may be an important contributor to physiological senescence in wild birds. As a by-product of aerobic respiration, reactive oxygen species are produced and can cause structural damage within cells. The anti-oxidant system deters oxidative damage to macromolecules. We examined oxidative stress and muscle ultrastructure in thick-billed murres aged 8 to 37 years (N=50) in pectoralis muscle biopsies. When considered in general linear models with body mass, body size and sex, no oxidative stress parameter varied with age. In contrast, there was a decrease in myonuclear domain similar to that seen in human muscle aging. We conclude that for wild birds with very high flight activity levels, muscle ultrastructural changes may be an important contributor to demographic senescence. Such gradual, linear declines in muscle morphology may eventually contribute to 'catastrophic' failure in foraging or predator avoidance abilities, leading to demographic senescence.

Earlier colony arrival but no trend in hatching timing in two congeneric seabirds (Uria spp.) across the North Atlantic.

A global analysis recently showed that seabird breeding phenology (as the timing of egg-laying and hatching) does not, on average, respond to temperature changes or advance with time (Keogan et al. 2018 Nat. Clim. Change8, 313-318). This group, the most threatened of all birds, is therefore prone to spatio-temporal mismatches with their food resources. Yet, other aspects of the breeding phenology may also have a marked influence on breeding success, such as the arrival date of adults at the breeding site following winter migration. Here, we used a large tracking dataset of two congeneric seabirds breeding in 14 colonies across 18° latitudes, to show that arrival date at the colony was highly variable between colonies and species (ranging 80 days) and advanced 1.4 days/year while timing of egg-laying remained unchanged, resulting in an increasing pre-laying duration between 2009 and 2018. Thus, we demonstrate that potentially not all components of seabird breeding phenology are insensitive to changing environmental conditions.

Do concentrations in eggs and liver tissue tell the same story of temporal trends of mercury in high Arctic seabirds?

Mercury (Hg) remains a key contaminant of concern in Arctic biota, and monitoring of Hg concentrations in seabird tissues will be an effective approach to track the effects of implementing the Minamata Convention. We examined trends in total Hg (THg) in liver and egg tissues of two Arctic seabirds, thick-billed murres (Uria lomvia) and northern fulmars (Fulmarus glacialis), between 1976 and 2013 to assess whether both tissues showed similar patterns of Hg change. Hepatic THg was consistently higher than egg THg, and both species had similar egg THg concentrations, but fulmars had higher hepatic THg than murres. Murre THg concentrations showed more relative variation through time than fulmars. We suggest that egg THg better reflects exposure of birds to THg in local, Arctic prey, whereas liver THg may incorporate longer term, year-round THg exposure. Additional analysis of THg distribution in Arctic seabirds post-laying would help inform interpretation of long-term trends.

Variation in Growth Drives the Duration of Parental Care: A Test of Ydenberg's Model.

The duration of parental care in animals varies widely, from none to lifelong. Such variation is typically thought to represent a trade-off between growth and safety. Seabirds show wide variation in the age at which offspring leave the nest, making them ideal to test the idea that a trade-off between high energy gain at sea and high safety at the nest drives variation in departure age (Ydenberg's model). To directly test the model assumptions, we attached time-depth recorders to murre parents (fathers [which do all parental care at sea] and mothers; [Formula: see text] of each). Except for the initial mortality experienced by chicks departing from the colony, the mortality rate at sea was similar to the mortality rate at the colony. However, energy gained by the chick per day was ∼2.1 times as high at sea compared with at the colony because the father spent more time foraging, since he no longer needed to spend time commuting to and from the colony. Compared with the mother, the father spent ∼2.6 times as much time diving per day and dived in lower-quality foraging patches. We provide a simple model for optimal departure date based on only (1) the difference in growth rate at sea relative to the colony and (2) the assumption that transition mortality from one life-history stage to the other is size dependent. Apparently, large variation in the duration of parental care can arise simply as a result of variation in energy gain without any trade-off with safety.

Body size affects individual winter foraging strategies of thick-billed murres in the Bering Sea.

Foraging and migration often require different energetic and movement strategies. Though not readily apparent, constraints during one phase might influence the foraging strategies observed in another. For marine birds that fly and dive, body size constraints likely present a trade-off between foraging ability and migration as smaller bodies reduce flight costs, whereas larger bodies are advantageous for diving deeper. This study examines individual wintering strategies of deep diving thick-billed murres (Uria lomvia) breeding at three colonies in the south-eastern Bering Sea: St Paul, St George and Bogoslof. These colonies, arranged north to south, are located such that breeding birds forage in a gradient from shelf to deep-water habitats. We used geolocation time-depth recorders and stable isotopes from feathers to determine differences in foraging behaviour and diet of murres during three non-breeding periods, 2008-2011. Body size was quantified by a principal component analysis (wing, culmen, head+bill and tarsus length). A hierarchical cluster analysis identified winter foraging strategies based on individual movement, diving behaviour and diet (inferred from stable isotopes). Structural body size differed by breeding island. Larger birds from St Paul had higher wing loading than smaller birds from St George. Larger birds, mainly from St Paul, dove to deeper depths, spent more time in the Bering Sea, and likely consumed higher trophic-level prey in late winter. Three winter foraging strategies were identified. The main strategy, employed by small birds from all three breeding colonies in the first 2 years, was characterized by high residency areas in the North Pacific south of the Aleutians and nocturnal diving. In contrast, 31% of birds from St Paul remained in the Bering Sea and foraged mainly during the day, apparently feeding on higher trophic-level prey. Throat feather stable isotopes indicated that individuals exhibited flexibility in the use of this colony-specific foraging strategy. The third strategy only occurred in 2010/2011, when birds dove more and deeper, suggesting limited prey resources. Foraging strategies partitioned with respect to annual differences, presumably in response to shifts in distribution of prey, and were linked to body size. The presence of a colony-specific wintering strategy suggests the potential for overwinter survival differences between these populations.

The Arctic: Glacial Refugium or Area of Secondary Contact? Inference from the Population Genetic Structure of the Thick-Billed Murre (Uria lomvia), with Implications for Management.

Quaternary glaciations affected the distribution of many species. Here, we investigate whether the Arctic represented a glacial refugium during the Last Glacial Maximum or an area of secondary contact following the ice retreat, by analyzing the genetic population structure of the thick-billed murre (Uria lomvia), a seabird that breeds throughout the North Atlantic, North Pacific and Arctic Oceans. The thick-billed murre is a species of socio-economic importance and faces numerous threats including hunting, oil pollution, gill netting, and climate change. We compared variation in the mitochondrial DNA (mtDNA) control region (n = 424), supplemented by 4 microsatellite loci (n = 445), among thick-billed murres sampled throughout their range. MtDNA data indicated that colonies comprise 4 genetically differentiated groups (Φst = 0.11-0.81): 1) Atlantic Ocean plus New Siberian Islands region, 2) Cape Parry, 3) Chukchi Sea, and 4) Pacific Ocean. Microsatellite variation differed between Atlantic and Pacific populations. Otherwise, little substructure was found within either ocean. Atlantic and Pacific populations appear to have been genetically isolated since the last interglacial period and should be considered separate evolutionary significant units for management. The Chukchi Sea and Cape Parry appear to represent areas of secondary contact, rather than arctic refugial populations.

The prudent parent meets old age: a high stress response in very old seabirds supports the terminal restraint hypothesis.

The reproductive success of wild animals usually increases with age before declining at the end of life, but the proximate mechanisms underlying those patterns remain elusive. Young animals are expected to invest less in current reproduction due to high prospects for future reproduction (the "restraint" hypothesis). The oldest animals may also show restraint when conditions are sub-optimal where even a small increase in reproductive investment may lead to death ("terminal restraint"). Alternatively, reproduction may be constrained by lack of experience and senescence (the "constraint" hypothesis). In two species of breeding seabirds, behavioural (time to return the offspring, calmness during restraint) and physiological (metabolism, glucose and corticosterone) parameters responded similarly to stress with advancing age, implying a generalized stress response. Across those parameters, birds were "shy" (high stress response) when young or old, and "bold" (low stress response) when middle-aged. Specifically, free corticosterone, the principal avian glucocorticoid responsible for directing energy away from reproduction and towards immediate survival following stress, was highest in both young and very old stressed birds. All age groups had a similar adrenal capacity to produce corticosterone, implying that middle-aged birds were showing restraint. Because the stress response, was highest at ages when the probability of current reproduction was lowest rather than at ages when the probability of future reproduction was highest we concluded that birds restrained reproductive investment based on current conditions rather than potential future opportunities. In particular, old birds showed terminal restraint when stressed. Hormonal cues promoted investment in adult survival over reproductive output at both the start and end of life consistent with the restraint hypothesis.

Age-related variation in energy expenditure in a long-lived bird within the envelope of an energy ceiling.

Energy expenditure in wild animals can be limited (i) intrinsically by physiological processes that constrain an animal's capacity to use energy, (ii) extrinsically by energy availability in the environment and/or (iii) strategically based on trade-offs between elevated metabolism and survival. Although these factors apply to all individuals within a population, some individuals expend more or less energy than other individuals. To examine the role of an energy ceiling in a species with a high and individually repeatable metabolic rate, we compared energy expenditure of thick-billed murres (Uria lomvia) with and without handicaps during a period of peak energy demand (chick-rearing, N = 16). We also compared energy expenditure of unencumbered birds (N = 260) across 8 years exhibiting contrasting environmental conditions and correlated energy expenditure with fitness (reproductive success and survival). Murres experienced an energy ceiling mediated through behavioural adjustments. Handicapped birds decreased time spent flying/diving and chick-provisioning rates such that overall daily energy expenditure remained unchanged across the two treatments. The energy ceiling did not reflect energy availability or trade-offs with fitness, as energy expenditure was similar across contrasting foraging conditions and was not associated with reduced survival or increased reproductive success. We found partial support for the trade-off hypothesis as older murres, where prospects for future reproduction would be relatively limited, did overcome an energy ceiling to invest more in offspring following handicapping by reducing their own energy reserves. The ceiling therefore appeared to operate at the level of intake (i.e. digestion) rather than expenditure (i.e. thermal constraint, oxidative stress). A meta-analysis comparing responses of breeding animals to handicapping suggests that our results are typical: animals either reduced investment in themselves or in their offspring to remain below an energy ceiling. Across species, whether a handicapped individual invested in its own energy stores or its offspring's growth was not explained by life history (future vs. current reproductive potential). Many breeding animals apparently experience an intrinsic energy ceiling, and increased energy costs lead to a decline in self-maintenance and/or offspring provisioning.

Quantifying marine traffic intensity in Northwest Greenland and the potential disturbance of two seabird colonies.

Marine traffic poses a growing threat to wildlife in the marine environment, including Arctic seabirds, which are exposed to high vessel densities when breeding in coastal areas. However, little is known about the magnitude of the problem. Here, we utilized underwater acoustic monitoring to quantify marine traffic and above-water disturbances at two thick-billed murre colonies in Greenland in 2016. We detected a total of 307 vessels, and only 4 % was known from automatic monitoring systems. Based on proximity, noise emission, and boating behavior, we classified 11 vessels as disturbing and an additional 12 as potentially disturbing for the seabirds. One colony facing population decline was located closest to the main boating route and experienced 2-5 times more disturbances than the other (increasing) colony, suggesting a negative impact of marine traffic. Our study shows that underwater acoustics can be a useful method to quantify above-water disturbances of seabird colonies.

Behavioural flexibility in an Arctic seabird using two distinct marine habitats to survive the energetic constraints of winter.

BACKGROUND: Homeothermic marine animals in Polar Regions face an energetic bottleneck in winter. The challenges of short days and cold temperatures are exacerbated for flying seabirds with small body size and limited fat stores. We use biologging approaches to examine how habitat, weather, and moon illumination influence behaviour and energetics of a marine bird species, thick-billed murres (Uria lomvia). METHODS: We used temperature-depth-light recorders to examine strategies murres use to survive winter in the Northwest Atlantic, where contrasting currents create two distinct marine habitats: cold (-0.1 ± 1.2 °C), shallower water along the Labrador Shelf and warmer (3.1 ± 0.3 °C), deep water in the Labrador Basin. RESULTS: In the cold shelf water, murres used a high-energy strategy, with more flying and less diving each day, resulting in high daily energy expenditure and also high apparent energy intake; this strategy was most evident in early winter when day lengths were shortest. By contrast, murres in warmer basin water employed a low-energy strategy, with less time flying and more time diving under low light conditions (nautical twilight and night). In warmer basin water, murres increased diving at night when the moon was more illuminated, likely taking advantage of diel vertically migrating prey. In warmer basin water, murres dove more at night and foraging efficiency increased under negative North Atlantic Oscillation (calmer ocean conditions). CONCLUSIONS: The proximity of two distinct marine habitats in this region allows individuals from a single species to use dual (low-energy/high-energy) strategies to overcome winter energy bottlenecks.

A comparison of techniques for classifying behavior from accelerometers for two species of seabird.

The behavior of many wild animals remains a mystery, as it is difficult to quantify behavior of species that cannot be easily followed throughout their daily or seasonal movements. Accelerometers can solve some of these mysteries, as they collect activity data at a high temporal resolution (<1 s), can be relatively small (<1 g) so they minimally disrupt behavior, and are increasingly capable of recording data for long periods. Nonetheless, there is a need for increased validation of methods to classify animal behavior from accelerometers to promote widespread adoption of this technology in ecology. We assessed the accuracy of six different behavioral assignment methods for two species of seabird, thick-billed murres (Uria lomvia) and black-legged kittiwakes (Rissa tridactyla). We identified three behaviors using tri-axial accelerometers: standing, swimming, and flying, after classifying diving using a pressure sensor for murres. We evaluated six classification methods relative to independent classifications from concurrent GPS tracking data. We used four variables for classification: depth, wing beat frequency, pitch, and dynamic acceleration. Average accuracy for all methods was >98% for murres, and 89% and 93% for kittiwakes during incubation and chick rearing, respectively. Variable selection showed that classification accuracy did not improve with more than two (kittiwakes) or three (murres) variables. We conclude that simple methods of behavioral classification can be as accurate for classifying basic behaviors as more complex approaches, and that identifying suitable accelerometer metrics is more important than using a particular classification method when the objective is to develop a daily activity or energy budget. Highly accurate daily activity budgets can be generated from accelerometer data using multiple methods and a small number of accelerometer metrics; therefore, identifying a suitable behavioral classification method should not be a barrier to using accelerometers in studies of seabird behavior and ecology.

Outlier analyses to test for local adaptation to breeding grounds in a migratory arctic seabird.

Investigating the extent (or the existence) of local adaptation is crucial to understanding how populations adapt. When experiments or fitness measurements are difficult or impossible to perform in natural populations, genomic techniques allow us to investigate local adaptation through the comparison of allele frequencies and outlier loci along environmental clines. The thick-billed murre (Uria lomvia) is a highly philopatric colonial arctic seabird that occupies a significant environmental gradient, shows marked phenotypic differences among colonies, and has large effective population sizes. To test whether thick-billed murres from five colonies along the eastern Canadian Arctic coast show genomic signatures of local adaptation to their breeding grounds, we analyzed geographic variation in genome-wide markers mapped to a newly assembled thick-billed murre reference genome. We used outlier analyses to detect loci putatively under selection, and clustering analyses to investigate patterns of differentiation based on 2220 genomewide single nucleotide polymorphisms (SNPs) and 137 outlier SNPs. We found no evidence of population structure among colonies using all loci but found population structure based on outliers only, where birds from the two northernmost colonies (Minarets and Prince Leopold) grouped with birds from the southernmost colony (Gannet), and birds from Coats and Akpatok were distinct from all other colonies. Although results from our analyses did not support local adaptation along the latitudinal cline of breeding colonies, outlier loci grouped birds from different colonies according to their non-breeding distributions, suggesting that outliers may be informative about adaptation and/or demographic connectivity associated with their migration patterns or nonbreeding grounds.

First Characterization of Avian Influenza Viruses from Greenland 2014.

In late February 2014, unusually high numbers of wild thick-billed murres (Uria lomvia) were found dead on the coast of South Greenland. To investigate the cause of death, 45 birds were submitted for laboratory examination in Denmark. Avian influenza viruses (AIVs) with subtypes H11N2 and low pathogenic H5N1 were detected in some of the birds. Characterization of the viruses by full genome sequencing revealed that all the gene segments belonged to the North American lineage of AIVs. The seemingly sparse and mixed subtype occurrence of low pathogenic AIVs in these birds, in addition to the emaciated appearance of the birds, suggests that the murre die-off was due to malnutrition as a result of sparse food availability or inclement weather. Here we present the first characterization of AIVs isolated in Greenland, and our results support the idea that wild birds in Greenland may be involved in the movement of AIV between North America and Europe.

Ingestion of plastic marine debris by Common and Thick-billed Murres in the northwestern Atlantic from 1985 to 2012.

Plastic ingestion by seabirds is a growing conservation issue, but there are few time series of plastic ingestion with large sample sizes for which one can assess temporal trends. Common and Thick-billed Murres (Uria aalge and U. lomvia) are pursuit-diving auks that are legally harvested in Newfoundland and Labrador, Canada. Here, we combined previously unpublished data on plastic ingestion (from the 1980s to the 1990s) with contemporary samples (2011-2012) to evaluate changes in murres' plastic ingestion. Approximately 7% of murres had ingested plastic, with no significant change in the frequency of ingestion among species or periods. The number of pieces of plastic/bird, and mass of plastic/bird were highest in the 1980s, lowest in the late 1990s, and intermediate in contemporary samples. Studying plastic ingestion in harvested seabird populations links harvesters to conservation and health-related issues and is a useful source of large samples for diet and plastic ingestion studies.