High latitude northern bats (Eptesicus nilssonii) reveal adaptations to both high and low ambient temperatures.

Insectivorous bats at northern latitudes need to cope with long periods of no food for large parts of the year. Hence, bats which are resident at northern latitudes throughout the year will need to undergo a long hibernation season and a short reproductive season where foraging time is limited by extended daylight periods. Eptesicus nilssonii is the northernmost occurring bat species worldwide and hibernates locally when ambient temperatures (Ta) limit prey availability. Therefore, we investigated the energy spent maintaining normothermy at different Ta, as well as how much bats limit energy expenditure while in torpor. We found that, despite being exposed to Ta as low as 1.1°C, bats did not increase torpid metabolic rate, thus indicating that E. nilssonii can survive and hibernate at low ambient temperatures. Furthermore, we found a lower critical temperature (Tlc) of 27.8°C, which is lower than in most other vespertilionid bats, and we found no indication of any metabolic response to Ta up to 37.1°C. Interestingly, carbon dioxide production increased with increasing Ta above the Tlc, presumably caused by a release of retained CO2 in bats that remained in torpor for longer and aroused at Ta above the Tlc. Our results indicate that E. nilssonii can thermoconform at near-freezing Ta, and hence maintain longer torpor bouts with limited energy expenditure, yet also cope with high Ta when sun exposed in roosts during long summer days. These physiological traits are likely to enable the species to cope with ongoing and predicted climate change.

State dependence of arousal from torpor in brown long-eared bats (Plecotus auritus).

To cope with periods of low food availability and unsuitable environmental conditions (e.g., short photoperiod or challenging weather), many heterothermic mammals can readily go into torpor to save energy. However, torpor also entails several potential costs, and quantitative energetics can, therefore, be influenced by the individual state, such as available energy reserves. We studied the thermal energetics of brown long-eared bats (Plecotus auritus) in the northern part of its distributional range, including torpor entry, thermoregulatory ability during torpor and how they responded metabolically to an increasing ambient temperature (Ta) during arousal from torpor. Torpor entry occurred later in bats with higher body mass (Mb). During torpor, only 10 out of 21 bats increased oxygen consumption (V̇O2) to a greater extent above the mean torpor metabolic rates (TMR) when exposed to low Ta. The slope of the torpid thermoregulatory curve was shallower than that of resting metabolic rate (RMR) during normothermic conditions, indicating a higher thermal insulation during torpor. During exposure to an increasing Ta, all bats increased metabolic rate exponentially, but the bats with higher Mb aroused at a lower Ta than those with lower Mb. In bats with low Mb, arousal was postponed to an Ta above the lower critical temperature of the thermoneutral zone. Our results demonstrate that physiological traits, which are often considered fixed, can be more flexible than previously assumed and vary with individual state. Thus, future studies of thermal physiology should to a greater extent take individual state-dependent effects into account.

Variation and correlation in the timing of breeding of North Atlantic seabirds across multiple scales.

Timing of breeding, an important driver of fitness in many populations, is widely studied in the context of global change, yet despite considerable efforts to identify environmental drivers of seabird nesting phenology, for most populations we lack evidence of strong drivers. Here we adopt an alternative approach, examining the degree to which different populations positively covary in their annual phenology to infer whether phenological responses to environmental drivers are likely to be (a) shared across species at a range of spatial scales, (b) shared across populations of a species or (c) idiosyncratic to populations. We combined 51 long-term datasets on breeding phenology spanning 50 years from nine seabird species across 29 North Atlantic sites and examined the extent to which different populations share early versus late breeding seasons depending on a hierarchy of spatial scales comprising breeding site, small-scale region, large-scale region and the whole North Atlantic. In about a third of cases, we found laying dates of populations of different species sharing the same breeding site or small-scale breeding region were positively correlated, which is consistent with the hypothesis that they share phenological responses to the same environmental conditions. In comparison, we found no evidence for positive phenological covariation among populations across species aggregated at larger spatial scales. In general, we found little evidence for positive phenological covariation between populations of a single species, and in many instances the inter-year variation specific to a population was substantial, consistent with each population responding idiosyncratically to local environmental conditions. Black-legged kittiwake Rissa tridactyla was the exception, with populations exhibiting positive covariation in laying dates that decayed with the distance between breeding sites, suggesting that populations may be responding to a similar driver. Our approach sheds light on the potential factors that may drive phenology in our study species, thus furthering our understanding of the scales at which different seabirds interact with interannual variation in their environment. We also identify additional systems and phenological questions to which our inferential approach could be applied.

A U-Turn for Mercury Concentrations over 20 Years: How Do Environmental Conditions Affect Exposure in Arctic Seabirds?

Mercury (Hg) is highly toxic in its methylated form (MeHg), and global change is likely to modify its bioavailability in the environment. However, it is unclear how top predators will be impacted. We studied blood Hg concentrations of chick-rearing black-legged kittiwakes Rissa tridactyla (2000-2019) in Svalbard (Norway). From 2000 to 2019, Hg concentrations followed a U-shaped trend. The trophic level, inferred from nitrogen stable isotopes, and chlorophyll a (Chl a) concentrations better predicted Hg concentrations, with positive and U-shaped associations, respectively. As strong indicators of primary productivity, Chl a concentrations can influence production of upper trophic levels and, thus, fish community assemblage. In the early 2000s, the high Hg concentrations were likely related to a higher proportion of Arctic prey in kittiwake's diet. The gradual input of Atlantic prey in kittiwake diet could have resulted in a decrease in Hg concentrations until 2013. Then, a new shift in the prey community, added to the shrinking sea ice-associated release of MeHg in the ocean, could explain the increasing trend of Hg observed since 2014. The present monitoring provides critical insights about the exposure of a toxic contaminant in Arctic wildlife, and the reported increase since 2014 raises concern for Arctic seabirds.

Assessment of shoulder rotation strength, muscle co-activation and shoulder pain in tetraplegic wheelchair athletes - A methodological study.

Objective: To develop a feasible protocol for testing maximum shoulder rotation strength in tetraplegic wheelchair athletes, and investigate concurrent validity of maximum isometric handheld dynamometer (HHD) towards maximum isokinetic dynamometer (ID) strength measurements; secondly, to study shoulder muscle activation during maximum shoulder rotation measurements, and the association between shoulder strength and shoulder pain.Design: Descriptive methodological.Setting: Danish Wheelchair Rugby (WCR) association for WCR tetraplegic athletes from local WCR-clubs.Participants: Twelve adult tetraplegics.Interventions: N/A.Outcome measures: Wheelchair User's Shoulder Pain Index (WUSPI) and Visual Analog Scale (VAS) measured shoulder pain, isometric HHD and ID (60°/s) measured maximum internal (IR) and external (ER) shoulder rotation strength. Surface Electromyography normalized to maximum EMG measured muscle activity (mm Infraspinatus and Latissimus Dorsi) during maximum shoulder rotation strength.Results: Concurrent validity of isometric HHD towards ID showed Concordance Correlation Coefficients of left and right arms 0.90 and 0.86 (IR), and 0.89 and 0.91 (ER), with no difference in muscle activity between isometric HHD and ID, but larger co-activation during ER. There was no association between shoulder strength and pain, except for significantly weak negative associations between ID and pain during ER for left and right arms (P = 0.03; P = 0.04).Conclusion: Standardized feasible protocol for tetraplegic wheelchair athletes for measuring maximum shoulder rotation strength was established. Isometric HHD is comparable with ID on normalized peak torques and muscle activity, but with larger co-activation. Strength was not clearly associated with shoulder pain.

Quantitative mismatch between empirical temperature-size rule slopes and predictions based on oxygen limitation.

In ectotherms, adult body size commonly declines with increasing environmental temperature, a pattern known as the temperature-size rule. One influential hypothesis explaining this observation is that the challenge of obtaining sufficient oxygen to support metabolism becomes greater with increasing body size, and more so at high temperatures. Yet, previous models based on this hypothesis do not account for phenotypic plasticity in the physiology of organisms that counteracts oxygen limitation at high temperature. Here, we model the predicted strength of the temperature-size response using estimates of how both the oxygen supply and demand is affected by temperature when allowing for phenotypic plasticity in the aquatic ectotherm Daphnia magna. Our predictions remain highly inconsistent with empirical temperature-size responses, with the prior being close to one order of magnitude stronger than the latter. These results fail to provide quantitative support for the hypothesis that oxygen limitation drives temperature-size clines in aquatic ectotherms. Future studies into the role of oxygen limitation should address how the strength of the temperature-size response may be shaped by evolution under fluctuating temperature regimes. Finally, our results caution against applying deterministic models based on the oxygen limitation hypothesis when predicting future changes in ectotherm size distributions under climate change.

Incubation temperature and physiological aging in the zebra finch.

In birds, incubation temperature has received increased attention as an important source of phenotypic variability in offspring. A lower than optimal incubation temperature may negatively affect aspects of nestling physiology, such as body growth and energy metabolism. However, the long-term effects of sub-optimal incubation temperature on morphology and physiology are not well understood. In a previous study, we showed that zebra finches from eggs incubated at a low temperature (35.9°C) for 2/3 of the total incubation time suffered a lower post-fledging survival compared to individuals that had been incubated at higher temperatures (37.0 and 37.9°C). In the present study, we investigated whether these variations in incubation temperature could cause permanent long-lasting differences in body mass, body size, or basal metabolic rate. Furthermore, we tested whether the observed differences in survival between treatment groups would be reflected in the rate of physiological deterioration, assessed through oxidative damage and decreased metabolic rate with age (i.e. 'metabolic aging'). Incubation temperature did not significantly affect embryonic or nestling body growth and did not influence final adult body mass or body size. Nor was there any long-term effect on basal metabolic rate. Birds from eggs incubated at the lowest temperature experienced an accumulation of oxidative damage with age, although this was not accompanied by an accelerated rate of metabolic aging. The present results suggest that the low survival in these birds was possibly mediated by increased oxidative stress, but independent of body growth and the basal metabolic rate.

Energetics of whiskered bats in comparison to other bats of the family Vespertilionidae.

Bats inhabit a variety of climate types, ranging from tropical to temperate zones, and environmental differences may therefore affect the basal metabolic rate (BMR) of bats from different populations. In the present study, we provide novel data on the energetics of whiskered bats (Myotis mystacinus), which is the smallest species within Chiroptera measured to date. We investigated the thermoregulatory strategies of M. mystacinus close to the northern limits of this species' distribution range and compared these data to other vespertilionid bats living in different climates. As mammals living in colder areas experience elevated thermoregulatory costs, often leading to an increase in BMR, we hypothesised that BMR of this northern population of whiskered bats would be higher than that of bats from climates with warm environmental temperatures. From a systematic literature search we obtained BMR estimates (N=47) from 24 species within Vespertilionidae. Our metabolic measurements of M. mystacinus in Norway (body mass of 4.4 g; BMR of 1.48 ml O2 g-1 h-1) were not different from other vespertilionid bats, based on the allometric equation obtained from the systematic literature search. Further, there was no effect of environmental temperature on BMR within Vespertilionidae. How these tiny bats adapt metabolically to high latitude living is thus still an open question. Bats do have a suite of physiological strategies used to cope with the varying climates which they inhabit, and one possible factor could be that instead of adjusting BMR they could express more torpor. This article has an associated First Person interview with the first author of the paper.

The standard metabolic rate of a land snail (Cepaea hortensis) is a repeatable trait and influences winter survival.

Phenotypic selection on physiological parameters is an underrepresented topic in studies of evolutionary biology. There is especially a lack of studies involving invertebrate organisms. We studied the repeatability of the standard metabolic rate (SMR) and the effect of individual variation in SMR on the subsequent winter survival in a terrestrial shell-bearing mollusc, the white-lipped snail (Cepaea hortensis) in mid-Norway. SMR was measured twice during the autumn and - after an experimental overwintering at controlled conditions - twice during the following spring. We found a significant repeatability of SMR over all three time periods tested, with a clear effect of time, with a high repeatability of 0.56 over 4 days during spring, 0.44 over 12 days in the autumn and 0.17 over 194 days from autumn to spring. That SMR is a repeatable physiological trait across the winter period during which a possible selection might occur, suggests that SMR could be a potential target of natural selection. We indeed found that the autumn SMR significantly influenced the probability of survival during the winter period, with a combination of a positive linear (P = .011) and a quadratic stabilizing (P = .001) effect on SMR. Our results hence support the view that metabolic rate is an important physiological component influencing the fitness of an organism.

How to quantify thermal acclimation capacity?

The capacity of organisms to acclimate will influence their ability to cope with ongoing global changes in thermal regimes. Here we highlight methodological issues associated with recent attempts to quantify variation in acclimation capacity among taxa and environments, and describe how these may introduce bias to conclusions. We then propose a measure of thermal acclimation capacity that more directly quantifies the process of acclimation. Future studies of variation in acclimation capacity should critically evaluate whether their chosen empirical metric accurately reflects the theoretical concept of acclimation.

Black-legged kittiwakes as messengers of Atlantification in the Arctic.

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.

Perfluorinated substances and telomeres in an Arctic seabird: Cross-sectional and longitudinal approaches.

Telomeres are non-coding DNA repeats located at the termini of eukaryotic chromosomes, regulated by dynamic processes balancing shortening and maintenance. Despite a mechanism to slow-down telomere shortening, cell division leads to progressive attrition of chromosomes, leading to the onset of cellular senescence or apoptosis. However, telomere restoration based on telomerase activity is the primary mechanism for telomere maintenance. Telomere length is associated to health and survival and can be impacted by a broad panel of environmental factors. However, the effect of contaminants on telomeres is poorly known for living organisms. The aim of this study was to investigate relationships between some poly- and perfluoroalkyl substances (PFASs), body condition and telomere length by using both a cross-sectional and longitudinal approach in adult breeding Black-legged kittiwakes (Rissa tridactyla) from Svalbard. First, we examined the associations between absolute telomere length and PFASs contamination in a given year (cross-sectional approach). Second, we investigated the relationships between telomere dynamics and PFASs contamination within a two years' time frame (longitudinal approach). Our results did not show any significant relationships of PFASs and body condition with absolute telomere length in a given year. Surprisingly, we found a positive and significant relationship between PFASs and telomere dynamics in both sexes with elongated telomere in birds bearing the highest concentrations of PFASs. Our study underlines (i) the need to investigate PFAS effects on telomere dynamics with a longitudinal approach and (ii) a potential positive effect of these contaminants on telomere length, with the most contaminated birds showing the slowest rate of telomere shortening or even displaying elongated ones. Our study is the first to report a relationship between PFASs and telomere length in free-living vertebrates. A possible underlying mechanism and other potential confounding factors are discussed.

Temporal variation in circulating concentrations of organochlorine pollutants in a pelagic seabird breeding in the high Arctic.

The present study explored short-term temporal variations in circulating concentrations of 3 legacy organochlorines with different physicochemical properties (polychlorinated biphenyl 153 [PCB-153], p,p'-dichlorodiphenyldichloroethylene [DDE], and hexachlorobenzene [HCB]) in breeding kittiwakes (Rissa tridactyla) in a colony in Svalbard (78°N), Norwegian Arctic. Concentrations were measured in blood of a large number (n = 412-521 blood samples, depending on the data analyses) of prebreeding, incubating, and chick-rearing birds over a period of 5 yr (2007-2011). The PCB-153 concentrations were equal in male and female blood in the prebreeding period, whereas females had significantly lower concentrations during incubation and chick rearing, probably because of their ability to eliminate organochlorines through egg laying. A similar temporal pattern was observed with DDE, although the lower concentrations in incubating females were not significant. Males and females had similar concentrations of HCB over all reproductive stages. The concentrations of all 3 compounds varied greatly between years. The concentrations of PCB-153 tended to decline over the study period, whereas concentrations of HCB showed an increasing trend, especially among chick-rearing males late in the season. Concentrations of PCB-153 increased approximately 2.5 times from the prebreeding to the chick-rearing period, concurrent with mobilization of body lipids (reduced body mass). A similar, but less pronounced trend was found for HCB. For DDE, however, kittiwakes had the highest concentrations in the prebreeding period, suggesting relatively high exposure in their winter areas. The present study documented large variations in circulating concentrations of legacy organochlorines among and within breeding seasons in kittiwakes, but the alterations within seasons were relatively consistent from year to year. Environ Toxicol Chem 2017;36:442-448. © 2016 SETAC.

Is there plasticity in developmental instability? The effect of daily thermal fluctuations in an ectotherm.

Diversified bet-hedging (DBH) by production of within-genotype phenotypic variance may evolve to maximize fitness in stochastic environments. Bet-hedging is generally associated with parental effects, but phenotypic variation may also develop throughout life via developmental instability (DI). This opens for the possibility of a within-generation mechanism creating DBH during the lifetime of individuals. If so, DI could in fact be a plastic trait itself; if a fluctuating environment indicates uncertainty about future conditions, sensing such fluctuations could trigger DI as a DBH response. However, this possibility has received little empirical attention. Here, we test whether fluctuating environments may elicit such a response in the clonally reproducing crustacean Daphnia magna. Specifically, we exposed genetically identical individuals to two environments of different thermal stability (stable vs. pronounced daily realistic temperature fluctuations) and tested for effects on DI in body mass and metabolic rate shortly before maturation. Furthermore, we also estimated the genetic variation in DI. Interestingly, fluctuating temperatures did not affect body mass, but metabolic rate decreased. We found no evidence for plasticity in DI in response to environmental fluctuations. The lack of plasticity was common to all genotypes, and for both traits studied. However, we found considerable evolvability for DI, which implies a general evolutionary potential for DBH under selection for increased phenotypic variance.

Exposure to oxychlordane is associated with shorter telomeres in arctic breeding kittiwakes.

Telomeres are DNA-protein complexes located at the end of chromosomes, which play an important role in maintaining the genomic integrity. Telomeres shorten at each cell division and previous studies have shown that telomere length is related to health and lifespan and can be affected by a wide range of environmental factors. Among them, some persistent organic pollutants (POPs) have the potential to damage DNA. However, the effect of POPs on telomeres is poorly known for wildlife. Here, we investigated the relationships between some legacy POPs (organochlorine pesticides and polychlorobiphenyls) and telomere length in breeding adult black-legged kittiwakes (Rissa tridactyla), an arctic seabird species. Our results show that among legacy POPs, only blood concentration of oxychlordane, the major metabolite of chlordane mixture, is associated with shorter telomere length in females but not in males. This suggests that female kittiwakes could be more sensitive to oxychlordane, potentially explaining the previously reported lower survival rate in most oxychlordane-contaminated kittiwakes from the same population. This study is the first to report a significant and negative relationship between POPs and telomere length in a free-living bird and highlights sex-related susceptibility to banned pesticides.

A 'slow pace of life' in Australian old-endemic passerine birds is not accompanied by low basal metabolic rates.

Life history theory suggests that species experiencing high extrinsic mortality rates allocate more resources toward reproduction relative to self-maintenance and reach maturity earlier ('fast pace of life') than those having greater life expectancy and reproducing at a lower rate ('slow pace of life'). Among birds, many studies have shown that tropical species have a slower pace of life than temperate-breeding species. The pace of life has been hypothesized to affect metabolism and, as predicted, tropical birds have lower basal metabolic rates (BMR) than temperate-breeding birds. However, many temperate-breeding Australian passerines belong to lineages that evolved in Australia and share 'slow' life-history traits that are typical of tropical birds. We obtained BMR from 30 of these 'old-endemics' and ten sympatric species of more recently arrived passerine lineages (derived from Afro-Asian origins or introduced by Europeans) with 'faster' life histories. The BMR of 'slow' temperate-breeding old-endemics was indistinguishable from that of new-arrivals and was not lower than the BMR of 'fast' temperate-breeding non-Australian passerines. Old-endemics had substantially smaller clutches and longer maximal life spans in the wild than new arrivals, but neither clutch size nor maximum life span was correlated with BMR. Our results suggest that low BMR in tropical birds is not functionally linked to their 'slow pace of life' and instead may be a consequence of differences in annual thermal conditions experienced by tropical versus temperate species.

Survival rate and breeding outputs in a high Arctic seabird exposed to legacy persistent organic pollutants and mercury.

Chronic exposure to pollutants may represent a threat for wildlife. We tested whether adult survival rate, breeding probability and breeding success the year of sampling and the following year were affected by blood levels of mercury or persistent organic pollutants in Svalbard black-legged kittiwake Rissa tridactyla, by using capture-mark-recapture models over a five-year period. Survival rate was negatively linked to HCB levels in females, to chlordane mixture and oxychlordane, tended to decrease with increasing PCBs or DDE levels, but was unrelated to mercury. Breeding probability decreased with increasing mercury levels during the sampling year and with increasing CHL or HCB levels during the following year, especially in males observed as breeders. Surprisingly, the probability of raising two chicks increased with increasing HCB levels. Although levels of these legacy pollutants are expected to decline, they represent a potential threat for adult survival rate and breeding probability, possibly affecting kittiwake population dynamics.

Is the rate of metabolic ageing and survival determined by Basal metabolic rate in the zebra finch?

The relationship between energy metabolism and ageing is of great interest because aerobic metabolism is the primary source of reactive oxygen species which is believed to be of major importance in the ageing process. We conducted a longitudinal study on captive zebra finches where we tested the effect of age on basal metabolic rate (BMR), as well as the effect of BMR on the rate of metabolic ageing (decline in BMR with age) and survival. Basal metabolic rate declined with age in both sexes after controlling for the effect of body mass, indicating a loss of functionality with age. This loss of functionality could be due to accumulated oxidative damage, believed to increase with increasing metabolic rate, c.f. the free radical theory of ageing. If so, we would expect the rate of metabolic ageing to increase and survival to decrease with increasing BMR. However, we found no effect of BMR on the rate of metabolic ageing. Furthermore, survival was not affected by BMR in the males. In female zebra finches there was a tendency for survival to decrease with increasing BMR, but the effect did not reach significance (P<0.1). Thus, the effect of BMR on the rate of functional deterioration with age, if any, was not strong enough to influence neither the rate of metabolic ageing nor survival in the zebra finches.

The stress of being contaminated? Adrenocortical function and reproduction in relation to persistent organic pollutants in female black legged kittiwakes.

High levels of environmental pollutants such as persistent organic pollutants (POPs) including PCB and DDT have been found in the Arctic and many of those pollutants may impair reproduction through endocrine disruption. Nevertheless, their effects on stress hormones remain poorly understood, especially in free-ranging birds. Corticosterone, the principal glucocorticoid in birds, can indirectly impair reproduction. The aim of the present study was to examine the relationships between POPs and reproduction through their potential consequences on different reproductive traits (breeding decision, egg-laying date, breeding success) and corticosterone secretion (baseline and stress-induced levels). We addressed those questions in an Arctic population of female black-legged kittiwakes during the pre-breeding stage and measured several legacy POPs (PCBs and pesticides: HCB, p,p'-DDE, CHL) in whole blood. POP levels were not related to breeding decision neither to breeding success, whereas females with high levels of pesticides laid their eggs earlier in the season. We found a negative relationship between POP levels and body condition index in non-breeding females. Black-legged kittiwakes with higher levels of PCB showed stronger adrenocortical response when subjected to a capture-handling stress protocol. We suggest that PCBs may disrupt corticosterone secretion whereas the positive relationship between pesticides and egg-laying date could either originate from a direct effect of pesticides or may be related to other confounding factors such as age or individual's quality. Although no direct negative reproduction output of POPs was found in this study, it is possible that the most contaminated individuals would be more sensitive to environmental stress and would be less able to maintain parental investment than less polluted individuals.

Migration and stress during reproduction govern telomere dynamics in a seabird.

Changes in telomere length are believed to reflect changes in physiological state and life expectancy in animals. However, much remains unknown about the determinants of telomere dynamics in wild populations, and specifically the influence of conditions during highly mobile life-history stages, for example migration. We tested whether telomere dynamics were associated with migratory behaviour and/or with stress during reproduction in free-living seabirds. We induced short-term stress during reproduction in chick-rearing, black-legged kittiwakes (Rissa tridactyla), tracked winter migration with geolocators and measured telomere length before and after winter migration. We found that time spent at wintering grounds correlated with reduced telomere loss, while stress during reproduction accelerated telomere shortening. Our results suggest that different life-history stages interact to influence telomere length, and that migratory patterns may be important determinants of variation in an individual's telomere dynamics.