Under pressure-exploring partner changes, physiological responses and telomere dynamics in northern gannets across varying breeding conditions.

Background: Life history theory predicts trade-offs between reproduction and survival in species like the northern gannet (Morus bassanus). During breeding, demanding foraging conditions lead them to expand their foraging range and diversify their diet, increasing the risk of reproductive failure. Changing partners may enhance breeding success but lead to more physiological costs. Methods: To investigate the physiological costs of reproduction upon partner changes, we measured and compared 21 biomarkers related to telomere dynamics, oxidative stress, inflammation, hematology, nutritional status, and muscle damage. We used a longitudinal approach with gannets (n = 38) over three contrasting years (2017, 2018 and 2019). Results: Our results suggest that annual breeding conditions exert a greater influence on physiological changes than partnership status. Individuals that changed partner experienced greater short-term stress than retained partners. This transient increase in stress was marked by short-term increases in oxidative lipid damage, lower antioxidant capacity, signs of inflammation, and greater weight loss than individuals that retained partners. During favorable conditions, individuals that changed mates had stabilized telomere length, decreased antioxidant capacity, glucose concentration, and muscle damage, along with increased oxygen transport capacity. Conversely, unfavorable breeding conditions led to increased telomere attrition, stabilized antioxidant capacity, decreased inflammation susceptibility, diminished oxygen transport capacity, and increased muscle damage. In the cases where partners were retained, distinct physiological changes were observed depending on the year's conditions, yet the telomere dynamics remained consistent across both partnership status categories. During the favorable year, there was an increase in unsaturated fatty acids and oxygen transport capacity in the blood, coupled with a reduction in inflammation potential and protein catabolism. In contrast, during the unfavorable year in the retained mates, we observed an increase in oxidative DNA damage, antioxidant capacity, weight loss, but a decrease in inflammation susceptibility as observed in changed mates. Discussion: Our study shows that behavioral flexibility such as mate switching can help seabirds cope with the challenges of food scarcity during reproduction, but these coping strategies may have a negative impact on physiological status at the individual level. In addition, the marked reduction in telomere length observed during harsh conditions, coupled with the stabilization of telomere length in favorable conditions, highlights the long-term physiological impact of annual breeding conditions on seabirds. These findings underscore the effect on their potential survival and fitness, emphasizing that the influence of annual breeding conditions is greater than that of partnership status.

Times and partners are a-changin': relationships between declining food abundance, breeding success, and divorce in a monogamous seabird species.

Seabirds exhibit considerable adjustment capacity to cope with environmental changes during the breeding season and to maximize lifetime reproductive output. For example, divorce has been proposed to be an adaptive behavioral strategy in social monogamous species, as a response to poor conditions and low breeding success. Here, we studied divorce at the population and individual levels in northern gannets (Morus bassanus, hereafter gannets) nesting on Bonaventure island (Quebec, Canada). At the population level, we used Granger's method for detecting and quantifying temporal causality between time series (from 2009 to 2019) of divorce rate and breeding success of gannets (n = 809) and we evaluated the relationship between breeding success and biomass of their two principal prey (Atlantic mackerel, Scomber scombrus, and Atlantic herring, Clupea harengus). Our results indicated that breeding success is mainly influenced by the spawning-stock biomass of Atlantic mackerel, and a decrease in breeding success is followed by an increase in divorce rate with a 1-year lag. However, the effect of the interaction between breeding success and year on the proportion of individuals that divorced showed significant inter-annual variation. At the individual level, our results support the adaptive strategy hypothesis of divorce. Indeed, gannets that changed partners did so following a reproductive failure, and there was an increase in breeding success 1 year following the divorce. Being central place foragers, opportunities for dispersal and adaptation are often limited for breeding seabirds in a context of low food abundance. We suggest that behavioral flexibility expressed as divorce would be an efficient short-term strategy for maintaining reproductive performance.

Egg-laying increases body temperature to an annual maximum in a wild bird.

Most birds, unlike reptiles, lay eggs successively to form a full clutch. During egg-laying, birds are highly secretive and prone to disturbance and predation. Using multisensor data loggers, we show that average daily body temperature during egg-laying is significantly increased (1 °C) in wild eider ducks (Somateria mollissima). Strikingly, this increase corresponds to the annual maximum body temperature (40.7 °C), representing a severe annual thermogenic challenge. This egg-laying-induced rise in body temperature may prove to be a common feature of wild birds and could be caused by habitat-related thermoregulatory adjustments and hormonal modulation of reproduction. We conclude our findings with new perspectives of the benefits of high body temperature associated with egg-laying of birds and the potential effect of heat stress that may occur with the future advent of heatwaves.

An investigation of physiological effects of the Deepwater Horizon oil spill on a long-distance migratory seabird, the northern gannet.

Exposure to oil can have long-term impacts on migratory birds. Following the 2010 Deepwater Horizon blowout in the Gulf of Mexico (GOM), we investigated potential impacts of oil exposure on a population of northern gannets (Morus bassanus) that breed on Bonaventure Island (Québec, Canada) and winter in GOM and along the U.S. Atlantic coast (AC). Blood and feather samples were collected from adults previously equipped with geolocators to determine wintering locations. Parent and alkylated polycyclic aromatic hydrocarbons (PAHs); trace metals; stable isotopes of carbon, nitrogen, and hydrogen; and immune, thyroid, steroid, retinoid, and genetic endpoints were measured. PAH and trace metal concentrations did not differ between gannets using different wintering sites. Feather stable isotope values varied significantly between birds from different wintering locations. Gannets wintering in GOM showed higher feather corticosterone and plasma thyroid hormone levels, which may indicate increased energetic demands and/or greater exposure to environmental stressors.

So far, so good Similar fitness consequences and overall energetic costs for short and long-distance migrants in a seabird.

Although there is a consensus about the evolutionary drivers of animal migration, considerable work is necessary to identify the mechanisms that underlie the great variety of strategies observed in nature. The study of differential migration offers unique opportunities to identify such mechanisms and allows comparisons of the costs and benefits of migration. The purpose of this study was to compare the characteristics of short and long-distance migrations, and fitness consequences, in a long-lived seabird species. We combined demographic monitoring (survival, phenology, hatching success) of 58 Northern Gannets (Morus bassanus) breeding on Bonaventure Island (Canada) and biologging technology (Global Location Sensor or GLS loggers) to estimate activity and energy budgets during the non-breeding period for three different migration strategies: to the Gulf of Mexico (GM), southeast (SE) or northeast (NE) Atlantic coast of the U.S. Survival, timing of arrival at the colony and hatching success are similar for short (NE, SE) and long-distance (GM) migrants. Despite similar fitness consequences, we found, as expected, that the overall energetic cost of migration is higher for long-distance migrants, although the daily cost during migration was similar between strategies. In contrast, daily maintenance and thermoregulation costs were lower for GM migrants in winter, where sea-surface temperature of the GM is 4-7o C warmer than SE and NE. In addition, GM migrants tend to fly 30 min less per day in their wintering area than other migrants. Considering lower foraging effort and lower thermoregulation costs during winter for long-distance migrants, this suggests that the energetic benefits during the winter of foraging in the GM outweigh any negative consequences of the longer-distance migration. These results support the notion that the costs and benefits of short and long-distance migration is broadly equal on an annual basis, i.e. there are no apparent carry-over effects in this long-lived bird species, probably because of the favourable conditions in the furthest wintering area.

It Takes Time to Be Cool: On the Relationship between Hyperthermia and Body Cooling in a Migrating Seaduck.

The large amount of energy expended during flapping flight is associated with heat generated through the increased work of the flight muscles. This increased muscle work rate can manifest itself in core body temperature (Tb) increase of 1-2°C in birds during flight. Therefore, episodic body cooling may be mandatory in migratory birds. To elucidate the thermoregulatory strategy of a short-distance migrant, common eiders (Somateria mollissima), we implanted data loggers in the body cavity of wild birds for 1 year, and report information on Tb during their entire migration for 19 individuals. We show that the mean body temperature during flight (TbMean) in the eiders was associated with rises in Tb ranging from 0.2 to 1.5°C, largely depending on flight duration. To understand how eiders are dealing with hyperthermia during migration, we first compare, at a daily scale, how Tb differs during migration using a before-after approach. Only a slight difference was found (0.05°C) between the after (40.30°C), the before (40.41°C) and the migration (40.36°C) periods, indicating that hyperthermia during flight had minimal impact at this time scale. Analyses at the scale of a flight cycle (flight plus stops on the water), however, clearly shows that eiders were closely regulating Tb during migration, as the relationship between the storage of heat during flight was highly correlated (slope = 1) with the level of heat dumping during stops, at both inter-individual and intra-individual levels. Because Tb at the start of a flight (TbStart) was significantly and positively related to Tb at the end of a flight (TbEnd), and the maximal attained Tb during a flight (TbMax), we conclude that in absence of sufficient body cooling during stopovers, eiders are likely to become increasingly hyperthermic during migration. Finally, we quantified the time spent cooling down during migration to be 36% of their daily (24 h) time budget, and conclude that behavioral body cooling in relation to hyperthermia represents an important time cost.

Feather corticosterone during non-breeding correlates with multiple measures of physiology during subsequent breeding in a migratory seabird.

Carry-over effects in migratory birds are likely mediated by physiological processes that are activated in response to environmental variation. Such processes affect body condition and/or reproductive success, and can include corticosterone (CORT) because this hormone responds to environmental stressors and influences energy balance. Few studies have considered how CORT levels during non-breeding relate to a broader physiological profile during subsequent breeding, and fewer still have considered measures other than body condition. To explore CORT's potential role in carry-over effects, we investigated the relationship between CORT and foraging ecology of northern gannets (Morus bassanus) during the non-breeding period, and tested for associations between these factors and variation in a suite of physiological and biochemical metrics during subsequent breeding. Northern gannets are the largest seabird top predator in the North Atlantic and were among the hardest hit by the Deepwater Horizon oil blowout in the Gulf of Mexico in 2010. We used light-level geolocators to confirm winter origins of individuals in our study. No interrelationships were found among levels of CORT from feathers grown during non-breeding (CORTf) and variation in foraging ecology, measured by stable isotopes of carbon (δ13C) and nitrogen (δ15N) from the same feathers. CORTf was correlated negatively with hematocrit and positively with triglyceride measured during subsequent incubation, and explained more variation in these variables than did body mass during incubation. These findings provide support for the hypothesis that energy management, measured using CORTf, during non-breeding carries over to influence physiological measures other than body condition. Gannets that previously wintered within the Gulf of Mexico in the years following the Deepwater Horizon oil blowout had higher levels of CORTf compared to birds that wintered along the Atlantic coast, suggesting an increased energetic cost associated with visiting the Gulf of Mexico. Our results indicate that CORT during non-breeding is associated with a broader physiological profile during subsequent breeding than previously reported in birds.

Does hyperthermia constrain flight duration in a short-distance migrant?

While some migratory birds perform non-stop flights of over 11 000 km, many species only spend around 15% of the day in flight during migration, posing a question as to why flight times for many species are so short. Here, we test the idea that hyperthermia might constrain flight duration (FD) in a short-distance migrant using remote biologging technology to measure heart rate, hydrostatic pressure and body temperature in 19 migrating eider ducks (Somateria mollissima), a short-distance migrant. Our results reveal a stop-and-go migration strategy where migratory flights were frequent (14 flights day(-1)) and short (15.7 min), together with the fact that body temperature increases by 1°C, on average, during such flights, which equates to a rate of heat storage index (HSI) of 4°C h(-1) Furthermore, we could not find any evidence that short flights were limited by heart rate, together with the fact that the numerous stops could not be explained by the need to feed, as the frequency of dives and the time spent feeding were comparatively small during the migratory period. We thus conclude that hyperthermia appears to be the predominant determinant of the observed migration strategy, and suggest that such a physiological limitation to FD may also occur in other species.This article is part of the themed issue 'Moving in a moving medium: new perspectives on flight'.

Does Water Temperature Affect the Timing and Duration of Remigial Moult in Sea Ducks? An Experimental Approach.

Aquatic birds have high cost of thermoregulation, especially during the moulting period, yet the effect of water temperature on the moulting strategy of aquatic birds has rarely been studied. Our general hypothesis is that energy savings associated with lower thermoregulation costs would be allocated to moulting processes. We predicted that aquatic birds moulting in warm water would have a higher level of body reserves, a faster growth rate of feathers, and an earlier remigial moult onset compared with birds moulting in cold water. We used the common eider (Somateria mollissima dresseri), a large sea duck, as the model species. Captive individuals were experimentally exposed to warm (18°C) and cold (8°C) water treatments during a three year period with individuals swapped between treatments. We found a similar feather growth rate for the two water temperature treatments and in contrast to our predictions, eiders exposed to warm water had a lower body mass and showed a delayed onset of remigial moult of approximately 7 days compared with those exposed to cold water. Our data indicate that body mass variations influence the timing of moult in unexpected ways and we suggest that it likely controls the occurrence of wing moult through a hormonal cascade. This study emphasizes the importance of improving our knowledge of the effects of water temperature on remigial moult of aquatic birds, to better assert the potential effects of global warming on their survival.

Associations between Resting, Activity, and Daily Metabolic Rate in Free-Living Endotherms: No Universal Rule in Birds and Mammals.

Energy management models provide theories and predictions for how animals manage their energy budgets within their energetic constraints, in terms of their resting metabolic rate (RMR) and daily energy expenditure (DEE). Thus, uncovering what associations exist between DEE and RMR is key to testing these models. Accordingly, there is considerable interest in the relationship between DEE and RMR at both inter- and intraspecific levels. Interpretation of the evidence for particular energy management models is enhanced by also considering the energy spent specifically on costly activities (activity energy expenditure [AEE] = DEE - RMR). However, to date there have been few intraspecific studies investigating such patterns. Our aim was to determine whether there is a generality of intraspecific relationships among RMR, DEE, and AEE using long-term data sets for bird and mammal species. For mammals, we use minimum heart rate (fH), mean fH, and activity fH as qualitative proxies for RMR, DEE, and AEE, respectively. For the birds, we take advantage of calibration equations to convert fH into rate of oxygen consumption in order to provide quantitative proxies for RMR, DEE, and AEE. For all 11 species, the DEE proxy was significantly positively correlated with the RMR proxy. There was also evidence of a significant positive correlation between AEE and RMR in all four mammal species but only in some of the bird species. Our results indicate there is no universal rule for birds and mammals governing the relationships among RMR, AEE, and DEE. Furthermore, they suggest that birds tend to have a different strategy for managing their energy budgets from those of mammals and that there are also differences in strategy between bird species. Future work in laboratory settings or highly controlled field settings can tease out the environmental and physiological processes contributing to variation in energy management strategies exhibited by different species.

Do seaducks minimise the flightless period? Inter- and intra-specific comparisons of remigial moult.

Remigial moult is one of the crucial events in the annual life cycle of waterfowl as it is energetically costly, lasts several weeks, and is a period of high vulnerability due to flightlessness. In waterfowl, remigial moult can be considered as an energy-predation trade-off, meaning that heavier individuals would minimise the flightless period by increasing feather growth rate and energy expenditure. Alternatively, they could reduce body mass at the end of this period, thereby reducing wing-loading to increase flight capability. We studied timing of remigial moult, primary growth rates, flightlessness duration, and the pattern of body mass variation in 5 species of captive seaducks (Melanitta fusca, M. perspicillata, Clangula hyemalis, Histrionicus histrionicus, and Somateria mollissima) ranging in size from 0.5 to 2.0 kg. Their feather growth rates weakly increased with body mass (M0.059) and no correlation was found at the intra-specific level. Consequently, heavier seaduck species and especially heavier individuals had a longer flightless period. Although birds had access to food ad libidum, body mass first increased then decreased, the latter coinciding with maximum feather growth rate. Level of body mass when birds regained flight ability was similar to level observed at the beginning of remigial moult, suggesting they were not using a strategic reduction of body mass to reduce the flightlessness duration. We suggest that the moulting strategy of seaducks may be the result of a compromise between using an intense moult strategy (simultaneous moult) and a low feather growth rate without prejudice to feather quality. Despite the controlled captive status of the studied seaducks, all five species as well as both sexes within each species showed timing of moult reflecting that of wild birds, suggesting there is a genetic component acting to shape moult timing within wild birds.

Endocrine status of a migratory bird potentially exposed to the Deepwater Horizon oil spill: a case study of northern gannets breeding on Bonaventure Island, Eastern Canada.

The Deepwater Horizon oil spill caused the death of a large number of seabirds in the Gulf of Mexico in 2010. However, the long term consequences of oil exposure on migratory birds overwintering in this area have received limited attention. The present study aimed to investigate the impact of oil contamination (e.g., polycyclic aromatic hydrocarbons (PAHs)) on the circulating status of prolactin and corticosterone, two hormones that influence reproductive success in birds, in Northern gannets (Morus bassanus) breeding on Bonaventure Island, Eastern Canada. Using light-based geolocators, it was found that 23.5% of Northern gannets from Bonaventure Island overwintered in the Gulf of Mexico in 2010-2011; the remainder of this population overwintered along the Atlantic Coast of the United States. PAH concentrations (eight compounds) in gannet blood cells were all found to be under the method limits of quantification, which could be the result of the ability of seabirds to metabolize these compounds and the time elapsed between oil exposure and blood sampling. Corticosterone and prolactin levels as well as body mass did not differ between the two major birds' wintering sites. Moreover, levels of both these hormones did not vary from early to late incubation period. Present results suggest that if Bonaventure Island-breeding Northern gannets had been exposed to oil in the Gulf of Mexico in the aftermath of this historical spill, this exposure could not be associated with changes in hormonal status and body mass in breeding individuals.

The body size-dependent diet composition of north american sea ducks in winter.

Daily food requirements scale with body mass and activity in animals. While small species of birds have higher mass-specific field metabolic rates than larger species, larger species have higher absolute energy costs. Under energy balance, we thus expect the small species to have a higher energy value diet. Also the weight and time constraints for flighted and diurnal foragers should set a maximum to the amount of prey items taken in one meal and to the daily number of meals, respectively. Further, avoidance of competition causes the species to reduce the amount of shared prey in their diet. Some diet segregation is therefore to be expected between species. We tested these hypotheses and investigated the role of body mass in the diet composition of 12 sea duck species (Somateria mollissima, Somateria spectabilis, Somateria fischeri, Polysticta stelleri, Bucephala clangula, Bucephala islandica, Bucephala albeola, Melanitta nigra, Melanitta perspicillata, Melanitta deglandi, Histrionicus histrionicus and Clangula hyemalis) wintering in North America. This study was based on a literature survey with special emphasis given to the diet data from the former US Bureau of Biological Survey. The data supported our hypothesis that the energy value of winter diet of sea ducks scales negatively with body mass. Diet diversity also scaled negatively with body mass. Our results suggest the existence of a minimum for the energy value of avian diets.

Seasonal variation in energy expenditure is not related to activity level or water temperature in a large diving bird.

There is considerable interest in understanding how the energy budget of an endotherm is modulated from a physiological and ecological point of view. In this paper, we used daily (24 h) heart rate (f(H24)), as a proxy of daily energy expenditure (DEE) across seasons, to test the effect of locomotion activity and water temperature on the energy budget of a large diving bird. f(H24) was monitored continuously in common eiders (Somateria mollissima) during 7 months together with measures of time spent flying and time spent feeding. f(H24) varied substantially during the recording period, with numerous increases and decreases that occurred across seasons, although we did not find any relationship between f(H24) and the time spent active (feeding and flying). However, inactive heart rate (f(H,inactive)) decreased as locomotion activity increased, suggesting that common eiders were using some form of compensation when under a high work load. We were also unable to detect a negative relationship between water temperature and resting heart rate, a proxy of resting metabolic rate. This was unexpected, based on the assumption that high thermoregulation costs would be associated with cold waters. We showed instead that a high level of energy expenditure coincided with feather moult and warm waters, which suggests that the observed variable pattern of seasonal DEE was driven by these two factors. Nevertheless, our results indicate that compensation and possibly the timing of moult may be used as mechanisms to reduce seasonal variation in energy expenditure.

Wild geese do not increase flight behaviour prior to migration.

Hypertrophy of the flight muscles is regularly observed in birds prior to long-distance migrations. We tested the hypothesis that a large migratory bird would increase flight behaviour prior to migration, in order to cause hypertrophy of the flight muscles, and upregulate key components of the aerobic metabolic pathways. Implantable data loggers were used to record year-round heart rate in six wild barnacle geese (Branta leucopsis), and the amount of time spent in flight each day was identified. Time in flight per day did not significantly increase prior to either the spring or the autumn migration, both between time periods prior to migration (5, 10 and 15 days), or when compared with a control period of low activity during winter. The lack of significant increase in flight prior to migration suggests that approximately 22 min per day is sufficient to maintain the flight muscles in condition for prolonged long-distance flight. This apparent lack of a requirement for increased flight activity prior to migration may be attributable to pre-migratory mass gains in the geese increasing workload during short flights, potentially prompting hypertrophy of the flight muscles.

The use of body mass loss to estimate metabolic rate in birds.

During starvation, energy production occurs at the expense of body reserve utilisation which results in body mass loss. Knowing the role of the fuels involved in this body mass loss, along with their energy density, can allow an energy equivalent of mass loss to be calculated. Therefore, it is possible to determine daily energy expenditure (DEE) if two body mass loss measurements at an interval of a few days are obtained. The technique can be cheap, minimally stressful for the animals involved, and the data relatively simple to gather. Here we review the use of body mass loss to estimate DEE in birds through critiquing the strengths and weaknesses of the technique, and detail the methodology and considerations that must be adhered to for accurate measures of DEE to be obtained. Owing to the biology of the species, the use of the technique has been used predominantly in Antarctic seabirds, particularly penguins and albatrosses. We demonstrate how reliable the technique can be in predicting DEE in a non-Antarctic species, common eiders (Somateria mollissima), the female of which undergoes a fasting period during incubation. We conclude that using daily body mass loss to estimate DEE can be a useful and effective approach provided that (1) the substrate being consumed during mass loss is known, (2) the kinetics of body mass loss are understood for the species in question and (3) only species that enter a full phase II of a fast (where substrate catabolism reaches a steady state) and are not feeding for a period of time are appropriate for this method.

To fly or not to fly: high flight costs in a large sea duck do not imply an expensive lifestyle.

A perennial question in ornithology is whether flight has evolved mostly to facilitate access to food or as an anti-predator strategy. However, flight is an expensive mode of locomotion and species using flight regularly are associated with an expensive lifestyle. Using heart rate (HR) data loggers implanted in 13 female common eiders (Somateria mollissima), our objective was to test the hypothesis that a high level of flight activity increases their energy budget. We used the long-term recording (seven months) of HR as an index of energy expenditure and the HR flight signature to compile all flight events. Our results indicate that the eider is one of the thriftiest volant birds with only 10 minutes of flight time per day. Consequently, we were not able to detect any effect of flight activity on their energy budget despite very high flight costs (123-149 W), suggesting that flight was controlled by energy budget limitations. However, the low flight activity of that species may also be related to their prey landscape requiring few or no large-scale movements. Nevertheless, we suggest that the (fitness) benefits of keeping flight ability in this species exceed the costs by allowing a higher survival in relation to predation and environmental harshness.

Flightlessness and the energetic cost of wing molt in a large sea duck.

Although the replacement of feathers apparently represents the major event of somatic production in the annual cycle of wild birds, knowledge about the energetics of molt has always been hampered by logistical and technical difficulties, which are exacerbated by the fact that birds are able to compensate behaviorally to buffer any variation in energy demand. During wing molt, sea ducks (Mergini) and other diving birds lose all of their wing feathers at once, leading to a period of temporary flightlessness of variable duration, a condition that considerably restricts their movements and increases the probability of predation. In the present study, we present the first results aimed at quantifying the duration of flightlessness, energy expenditure, and foraging effort during molt of a wing-propelled diving bird, the Common Eider (Somateria mollissima). Data loggers were implanted in the body cavity of 13 females to record heart rate and hydrostatic pressure (depth) every two seconds for a period of 220 days. Flight frequency and duration were assessed from elevated and constant heart rate, and the absence of flight was used to quantify the duration of flightlessness, which lasted, on average, 36 +/- 8 days (mean +/- SD). Using a period of four weeks before and four weeks after the flightless period, we found that dive depth (ranging from 1 to 2 m, on average) and daily diving time did not vary during the course of the study. Daily metabolic rate increased by 9%, and resting metabolic rate by 12% from the pre-molt period to the flightless period and remained high during the post-molt period. This study indicates that the energetic costs of replacing flight remiges in female eiders are substantial, although this is not associated with any change in foraging effort, which suggests that female Common Eiders lose mass during wing molt. Finally, estimates of energy savings associated with the total absence of flights during wing molt represent 6% of daily metabolic rate or 14% of resting metabolic rate. This finding contrasts with the classical view that little or no benefit is associated with a flightless condition. We suggest that such energy savings may have favored the evolution of temporary flightlessness in diving birds.

It is time to move: linking flight and foraging behaviour in a diving bird.

Although the adaptive value of flight may seem obvious, it is the most difficult behaviour of birds to monitor. Here, we describe a technique to quantify the frequency and the duration of flights over several months by implanting a data logger that records heart rate (fH), hydrostatic pressure (diving depth) and the body angle of a large sea duck species, the common eider (Somateria mollissima). According to the mean fH recorded during flight and the parameters recorded to identify the fH flight signature, we were able to identify all flights performed by 13 individuals during eight months. We cumulated local flight time (outside migrations) and found that activity occurs primarily during dawn and morning and that flying activities are strongly related to diving activities (Pearson's r=0.88, permutation test p<0.001). This relationship was interpreted as a consequence of living in a dynamic environment where sea currents move the ducks away from the food patches. We believe that the technique described here will open new avenues of investigation in the adaptive value of flight.