Seasonal sex–specific energy expenditure in breeding and non-breeding Palestine sunbirds Nectarinia osea

The timing of the chick‐rearing phase is known to have a profound effect on the reproductive success of birds. However, little is known about the energetic costs faced by the parents during different periods of the breeding season. These costs may have vital consequences for both their survival and future reproduction. In most studies, daily energy expenditure (DEE) of breeding and non‐breeding birds has been compared, without controlling for the effect of season. In the present study, we examined the energy demands of breeding compared to non‐breeding Palestine sunbirds Nectarinia osea and whether there were sex‐specific differences in DEE within and between different seasons. We predicted that DEE would be elevated when birds rear chicks, especially at cooler ambient temperatures. Time‐energy budgets were constructed for pairs of sunbirds, rearing chicks, or not breeding, in spring and summer. There were significant seasonal differences in estimates of DEE in non‐breeders that were 21% higher in spring than in summer. We attributed these to increases in non‐flight metabolic rate rather than changes in time spent on different activities. Our estimates of DEE for the birds that were rearing chicks were higher than non‐breeding adults. In females the increase in DEE when breeding, compared to when not breeding, was similar in both spring and summer, while males increased their DEE much less when breeding in spring. The differences in estimated DEE, however, were not significant between male and female birds in any season. Between seasons, female breeders had 17.1% higher DEE in spring than in summer, while male breeders showed no difference in DEE when rearing chicks in different seasons. Accordingly, our initial prediction was supported, as DEE in chick‐rearing adults was higher than in non‐breeding adults. In addition, although temperatures are lower in spring, breeding in the spring is only more costly than breeding in summer for females. Apparently, males are more flexible in reallocating their time and energy spent on different activities.     

Optimal body size and energy expenditure during winter: why are voles smaller in declining populations?

Winter is energetically challenging for small herbivores because of greater energy requirements for thermogenesis at a time when little energy is available. We formulated a model predicting optimal wintering body size, accounting for the scaling of both energy expenditure and assimilation to body size, and the trade-off between survival benefits of a large size and avoiding survival costs of foraging. The model predicts that if the energy cost of maintaining a given body mass differs between environments, animals should be smaller in the more demanding environments, and there should be a negative correlation between body mass and daily energy expenditure (DEE) across environments. In contrast, if animals adjust their energy intake according to variation in survival costs of foraging, there should be a positive correlation between body mass and DEE. Decreasing temperature always increases equilibrium DEE, but optimal body mass may either increase or decrease in colder climates depending on the exact effects of temperature on mass-specific survival and energy demands. Measuring DEE with doubly labeled water on wintering Microtus agrestis at four field sites, we found that DEE was highest at the sites where voles were smallest despite a positive correlation between DEE and body mass within sites. This suggests that variation in wintering body mass between sites was due to variation in food quality/availability and not adjustments in foraging activity to varying risks of predation.     

Daily energy expenditure of singing great reed warblers Acrocephalus arundinaceus

According to honest signalling theory, signals must be costly to produce to retain information about the signaller's quality. The song produced by male birds during breeding is a vocal "ornament" used for intra- and inter-sexual purposes. The energetic cost of this vocal signal remains a contentious issue. We used the doubly labelled water method to measure field metabolic rate by estimating CO₂ production and then convert this to daily energy expenditure (DEE) in great reed warbler males singing under natural conditions (10 at low to moderate intensity and 7 at very high intensity from dawn to dusk). There was a significant positive relationship between singing intensity and DEE. From this relationship we extrapolated the average DEE for intensely singing males (i.e., males producing song sounds 50% of the time and hence sitting at their elevated song post in the top of a reed stem more or less continuously throughout the ∼20 h of daylight) to 3.3×BMR (basal metabolic rate) and for non-singing males to 2.2×BMR. The mean DEE measured for the seven males singing with very high intensity was 3.1×BMR. The maximum measured DEE for a single male was 3.9×BMR, i.e. close to the maximum sustainable DEE (4×BMR), and the minimum DEE was 2.1×BMR for a male singing at very low intensity. These results imply that producing intensive advertising song in birds may incur a substantial cost in terms of increased energy expenditure.     

Differential reproductive responses to stress reveal the role of life-history strategies within a species

Life-history strategies describe that ‘slow’- in contrast to ‘fast’-living species allocate resources cautiously towards reproduction to enhance survival. Recent evidence suggests that variation in strategies exists not only among species but also among populations of the same species. Here, we examined the effect of experimentally induced stress on resource allocation of breeding seabirds in two populations with contrasting life-history strategies: slow-living Pacific and fast-living Atlantic black-legged kittiwakes. We tested the hypothesis that reproductive responses in kittiwakes under stress reflect their life-history strategies. We predicted that in response to stress, Pacific kittiwakes reduce investment in reproduction compared with Atlantic kittiwakes. We exposed chick-rearing kittiwakes to a short-term (3-day) period of increased exogenous corticosterone (CORT), a hormone that is released during food shortages. We examined changes in baseline CORT levels, parental care and effects on offspring. We found that kittiwakes from the two populations invested differently in offspring when facing stress. In response to elevated CORT, Pacific kittiwakes reduced nest attendance and deserted offspring more readily than Atlantic kittiwakes. We observed lower chick growth, a higher stress response in offspring and lower reproductive success in response to CORT implantation in Pacific kittiwakes, whereas the opposite occurred in the Atlantic. Our findings support the hypothesis that life-history strategies predict short-term responses of individuals to stress within a species. We conclude that behaviour and physiology under stress are consistent with trade-off priorities as predicted by life-history theory. We encourage future studies to consider the pivotal role of life-history strategies when interpreting inter-population differences of animal responses to stressful environmental events.     

Cost of living in free-ranging degus (Octodon degus): seasonal dynamics of energy expenditure

Animals process and allocate energy at different seasons at variable rates, depending on their breeding season and changes in environmental conditions and resulting physiological demands. Overall total energy expenditure, in turn, should either increase in some seasons due to special added demands (e.g. reproduction) or it could simply remain at about the same level, in which case the animals must show compensatory rebalancing of other expenditures that can be reduced. To test for the alternative hypotheses of seasonal variability or compensation, we measured total daily energy expenditure (DEE) in free-living degus (Octodon degus) at four seasons and followed this with determinations of basal metabolic rate (BMR) in the laboratory in the same individuals. DEE varied seasonally but was only significantly different (lower) in summer (non-breeding season), with a DEE:BMR ratio of only 1.6, whereas autumn, winter and spring DEE values were statistically indistinguishable from one another and showed DEE:BMR ratios ranging from 1.9 to 2.2. Our values of DEE in the field fall within the broad range of allometric expectation for herbivorous mammals in general, but the ratios of DEE:BMR are lower than expected. This, together with the lack of strong major shifts in total levels of DEE, suggests that degus are showing compensatory shifts among various categories of energy expenditure that allow them to manage their overall energy balance by minimizing total expenditure.     

Testosterone increases activity but not daily energy expenditure in captive male dark-eyed juncos, Junco hyemalis

Plasma testosterone (T) levels in male dark-eyed juncos peak early in the breeding season, then decline. If T enhances opportunities for reproductive success, as suggested by previous experiments, why does elevated T not occur naturally? To address this question, we prolonged the early peak level throughout the breeding season and explored potential energetic costs of maintaining elevated T. We measured daily energy expenditure (DEE) of treated males (T-males) and controls (C-males) using doubly labelled water (DLW). We also conducted behaviour scans of T- and C-males housed in outdoor aviaries. DEE was not higher in T-males than in C-males. However, T-males did increase locomotion and foraging and decrease rest and self-maintenance. These results suggest that elevated T may increase the contribution of some components of DEE and lower the contribution of others. Furthermore, the T-induced decrease in allocation of time to rest and maintenance may represent a long-term cost that has led to selection against the maintenance of elevated T beyond the natural early spring peak. Copyright 2000 The Association for the Study of Animal Behaviour.     

Energy expenditure during egg laying is equal for early and late breeding free-living female great tits

In many bird populations, variation in the timing of reproduction exists but it is not obvious how this variation is maintained as timing has substantial fitness consequences. Daily energy expenditure (DEE) during the egg laying period increases with decreasing temperatures and thus perhaps only females that can produce eggs at low energetic cost will lay early in the season, at low temperatures. We tested whether late laying females have a higher daily energy expenditure during egg laying than early laying females in 43 great tits (Parus major), by comparing on the same day the DEE of early females late in their laying sequence with DEE of late females early in their egg laying sequence. We also validated the assumption that there are no within female differences in DEE within the egg laying sequence. We found a negative effect of temperature and a positive effect of female body mass on DEE but no evidence for differences in DEE between early and late laying females. However, costs incurred during egg laying may have carry-over effects later in the breeding cycle and if such carry-over effects differ for early and late laying females this could contribute to the maintenance of phenotypic variation in laying dates.     

Male Weasels Decrease Activity and Energy Expenditure in Response to High Ambient Temperatures

The heat dissipation limit (HDL) hypothesis suggests that the capacity of endotherms to dissipate body heat may impose constraints on their energy expenditure. Specifically, this hypothesis predicts that endotherms should avoid the detrimental consequences of hyperthermia by lowering their energy expenditure and reducing their activity in response to high ambient temperatures (T_a). We used an extensive data set on the daily energy expenditure (DEE, n = 27) and the daily activity time (AT, n = 48) of male weasels (Mustela nivalis) during the spring and summer breeding season to test these predictions. We found that T_a was related in a “hump-shaped” (i.e. convex) manner to AT, DEE, resting metabolic rate (RMR) and metabolic scope (the ratio of DEE to RMR). These results support the HDL hypothesis because in response to warm T_as male weasels reduced their AT, DEE, and RMR. Although the activity and energy expenditure of large endotherms are most likely to be constrained in response to warm T_as because they are less able to dissipate heat, our results suggest that small endotherms may also experience constraints consistent with the HDL hypothesis.     

Hormone and metabolite changes associated with extended breeding fasts in male northern elephant seals (Mirounga angustirostris)

We measured metabolic hormones and several key metabolites in breeding adult male northern elephant seals to examine the regulation of fuel metabolism during extended natural fasts of over 3 months associated with high levels of energy expenditure. Males were sampled twice, early and late in the fast, losing an average of 23% of body mass and 47% of adipose stores between measurements. Males exhibited metabolic homeostasis over the breeding fast with no changes in glucose, non-esterified fatty acids, or blood urea nitrogen. Ketoacids increased over the fast but were very low when compared to other fasting species. Changes within individuals in total triiodothyronine (tT₃) were positively related to daily energy expenditure (DEE) and protein catabolism. Differences in levels of thyroid hormones relative to that observed in weaned pups and females suggest a greater deiodination of T₄ to support the high DEE of breeding males. Relative levels of leptin and ghrelin were consistent with the suppression of appetite but a significant reduction in growth hormone across the fast was contrary to expectation in fasting mammals. The lack of the increase in cortisol during fasting found in conspecific weaned pups and lactating females may contribute to the ability of breeding males to spare protein despite high levels of energy expenditure. Together these findings reveal significant differences with conspecifics under varying nutrient demands, suggesting metabolic adaptation to extended high energy fasts.     

The energetics of lactation in cooperatively breeding meerkats Suricata suricatta

Species may become obligate cooperative breeders when parents are unable to raise their offspring unassisted. We measured the daily energy expenditure of mothers, helpers and offspring during peak lactation in cooperatively breeding meerkats Suricata suricatta using the doubly labelled water technique. Lactating mothers expended more energy per day than allo-lactating subordinate females, non-lactating females or suckling offspring. Metabolizable energy intakes of lactating mothers were calculated from isotope-based estimates of offspring milk energy intake, and were not significantly different from the previously suggested maximal limit for mammals. Allo-lactating females were the only category of animals that lost weight during the period of study, probably because they spent more time babysitting than non-lactating females. Daily energy expenditure (DEE) of lactating mothers increased with litter size but decreased with the number of helpers. Calculations show that for every 10 helpers, even in the absence of allo-lactators, mothers are able to reduce their DEE during peak lactation by an amount equivalent to the energy cost of one pup. These results indicate that helpers have beneficial energetic consequences for lactating mothers in an obligate cooperatively breeding mammal.     

Reproductive phenology of a food-hoarding mast-seed consumer: resource- and density-dependent benefits of early breeding in red squirrels

The production of offspring by vertebrates is often timed to coincide with the annual peak in resource availability. However, capital breeders can extend the energetic benefits of a resource pulse by storing food or fat, thus relaxing the need for synchrony between energy supply and demand. Food-hoarding red squirrels (Tamiasciurus hudsonicus) breeding in the boreal forest are reliant on cones from a masting conifer for their nutrition, yet lactation is typically completed before the annual crop of cones is available for consumption such that peaks in energy supply and demand are not synchronized. We investigated the phenological response of red squirrels to annual variation in environmental conditions over a 20-year span and examined how intra- and inter-annual variation in the timing of reproduction affected offspring recruitment. Reproductive phenology was strongly affected by past resource availability with offspring born earlier in years following large cone crops, presumably because this affected the amount of capital available for reproduction. Early breeders had higher offspring survival and were more likely to renest following early litter loss when population density was high, perhaps because late-born offspring are less competitive in obtaining a territory when vacancies are limited. Early breeders were also more likely to renest after successfully weaning their first litter, but renesting predominantly occurred during mast years. Because of their increased propensity to renest and the higher survival rates of their offspring, early breeders contribute more recruits to the population but the advantage of early breeding depends on population density and resource availability.     

Energy expenditure and testosterone in free-living male yellow-pine chipmunks

The onset of mating in yellow-pine chipmunks (Tamias amoenus) follows emergence from a prolonged period of energy conservation during hibernation. Energy expenditures are greatly accelerated to meet the demands of the reproductive season. When emerging from hibernation, typical male chipmunks (breeders) have enlarged testes and a high level of plasma testosterone (T). However, certain males that do not participate in reproduction (nonbreeders) maintain small testes and low plasma T levels and emerge several weeks later than the breeders. The timing of the terminal arousal from hibernation and onset of mating are associated with increased plasma T levels. Experimental elevation of T levels in T. amoenus outside the mating season has been associated with a decrease in body mass, further suggesting an effect of T on energy balance. To test this hypothesis, we measured daily energy expenditure (DEE) in free-living, nonbreeding male chipmunks in the presence and absence of a T-implant. We also measured DEE in breeding males when endogenous T levels were high. DEE of the nonbreeders was not affected by our manipulation of plasma T, and the DEE of breeding males did not differ from that of nonbreeders. We conclude that energy expenditure on a daily basis in male yellow-pine chipmunks is not influenced by levels of T. However, on a seasonal basis, the earlier emergence from hibernation by breeding males, which appears to be influenced by T, represents an overall seasonal energy expenditure that exceeds that of nonbreeding males.     

Ultimate and proximate factors affecting the breeding performance of a marine top-predator

Variability in ecosystems affects the life history of organisms. In marine ecosystems where interannual variability is high, relationships between fluctuations in oceanographic parameters and top-predator breeding performance are increasingly documented but it is less clear why such relationships exist. In this study, we examined the connections between marine environment fluctuations and breeding performance of a long-lived top-predator, the black-browed albatross Diomedea melanophris at Kerguelen, through study of resource acquisition and allocation processes. Our results show that this population used the same foraging zones and spent similar time foraging year after year, but adult body condition varied between years. Foraging trips are regulated mainly by changes in body condition. During years of low resource availability, birds return to their nest with lower body condition and adults in low body condition were more frequent and therefore were more likely to stop breeding. Poor breeding success was related to the presence of colder waters in the foraging zones of breeding albatrosses as measured by the positive correlation between sea surface temperatures and breeding success measured over 18 years. Lower breeding success was mainly due to failure by inexperienced birds. The results of this study demonstrate how oceanographic conditions affect breeding performance through allocation processes. We compared these results to those at South Georgia where the breeding success is lower and more variable. This population relies mainly on krill, a resource that shows a very variable year-to-year availability compared to fish prey consumed by Kerguelen birds. This study shows that, in the same species, differences in resource variability and availability affect the demographic strategies probably through differences in allocation strategies.     

Evaluating the prudence of parents: daily energy expenditure throughout the annual cycle of a free-ranging bird, the macaroni penguin Eudyptes chrysolophus

We measured daily energy expenditure (DEE) continuously for a whole year in a free ranging bird, the macaroni penguin Eudyptes chrysolophus . We combined these measurements with concurrently recorded foraging behaviour, and literature information on body mass and dietary factors to estimate prey consumption rates and foraging success. DEE was at a maximum during late chick-rearing but was equally high during all other active phases of the breeding season. DEE was approximately 4×resting metabolic rate, which accords with established theory and suggests a common 'energetic ceiling' throughout the summer period. However, whether this represents a maximum in physiological capacity, or a rate which optimises fitness is still unclear. Rates of prey consumption and foraging success followed different patterns from daily energy expenditure. Daily prey consumption was high as the penguins prepared for long fasts associated with moulting and incubation but relatively low during chick-rearing, when foraging areas were restricted and foraging success lower. It appears that the energy intake of macaroni penguins is subject to extrinisic or environmental constraints rather than to intrinsic physiological limits.     

Variable energetic responses to clutch size manipulations in white-throated dippers Cinclus cinclus

We used the doubly-labelled water technique to measure daily energy expenditure (DEE) of a free-living uniparental incubator, the white-throated dipper Cinclus cinclus , in Scotland. DEE was 205±8 (s.e.m.) kJ d⁻¹ for 17 females incubating their natural clutch sizes, equivalent to 3.2±0.1×basal metabolic rate (BMR). To investigate the influence of clutch size on the energy budget, we measured the DEE of 14 females with clutches increased or reduced by a single egg. Birds with reduced clutch sizes had an energy expenditure with a mean and variance that did not differ from those of birds with unmanipulated clutches. Enlarging the clutch led to an increase in energy expenditure to over 4×BMR for some individuals but not for others, resulting in greater variance in energy expenditure for birds with enlarged clutches. Individual variation in energy expenditure could not be fully explained by environmental conditions, by patterns of behaviour or clutch size. Incubating females received a maximum of only 4 kJ d⁻¹ (2% of DEE) from provisioning by the male, and mobilised up to 6 kJ d⁻¹ (3% of DEE) from reserves. Females spent 2.9±0.2 h (n=20) away from the nest each day, so a foraging rate of 95 kJ h⁻¹ was required during incubation recesses to balance DEE. This 'required foraging rate' is double previous estimates of the maximum rates of energy acquisition for birds of this size. We suggest that the greater likelihood of a raised energy expenditure associated with larger clutches, combined with the difficulties in maintaining energy supplies, may constitute a constraint on avian clutch size.     

Age-related reproductive variation in a wild marine mammal population

Life history theory predicts a change in reproduction success with age as energy resources are limited and must be allocated effectively to maximize reproduction and survival. In this study, we use three reproductive performance measures, maternal expenditure, offspring weaning mass, and first-year survival, to investigate the role that maternal age plays in successful reproduction. Long-term uninterrupted life history data available for Marion Island’s southern elephant seals and mass change estimates from photogrammetry data allow for assessment of age-related reproduction performance and trade-offs. Known-aged adult females were photographed for photogrammetric mass estimation (n = 29) and their pups weighed at weaning during the 2009 breeding season. Maternal age and proportional mass loss positively influenced pup weaning mass. In turn, first-year pup return rates (as a proxy for survival) were assessed through the intensive mark–recapture program. Pup survival increased with female age and weaning mass. Pups of young females aged 3–6 years have a lower first-year survival probability compared with pups of older and larger females.     

Spying on your neighbours? Social information affects timing of breeding and stress hormone levels in a colonial seabird

A good overlap between offspring energetic requirements and availability of resources is required for successful reproduction. Accordingly, individuals from numerous species fine-tune their timing of breeding by integrating cues that predict environmental conditions during the offspring period. Besides acquiring information from their direct interaction with the environment (personal information), individuals can integrate information by observing the behaviours or performance of others (social information). The use of social information is often beneficial because the accumulated knowledge of conspecifics may represent a source of information more reliable than the intrinsically more limited personal information. However, although social information constitutes the major source of information in a wide range of contexts, studies investigating its use in the context of timing of breeding are scarce. We investigated whether black-legged kittiwakes (Rissa tridactyla) used social information to adjust the timing of egg-laying. We manipulated social information using a food-supplementation experiment, known to advance kittiwakes' reproductive phenology. We expected food-supplemented and unsupplemented pairs to delay and advance, respectively, their timing of laying when surrounded by a majority of neighbours from the opposite food-treatment. However, both unsupplemented and food-supplemented kittiwakes delayed egg-laying when surrounded by a higher proportion of neighbours from the opposite food-treatment. This result shows that kittiwakes use social information to time egg-laying, but that it is not used to match the seasonal peak of food availability. We suggest that when social and personal cues give contradictory environmental information, individuals may benefit from delaying laying to gather more information to make better decisions about investment into eggs. Further, we explored a potential proximate mechanism for the pattern we report. We show that baseline corticosterone, known to mediate reproductive decisions, was lower in unsupplemented females facing a higher proportion of food-supplemented neighbours. Altogether, our results suggest that to fine-tune their timing of laying, kittiwakes use complex decision-making processes in which social and personal information interplay.

     

The energetic and survival costs of growth in free-ranging chipmunks

The growth/survival trade-off is a fundamental aspect of life-history evolution that is often explained by the direct energetic requirement for growth that cannot be allocated into maintenance. However, there is currently no empirical consensus on whether fast-growing individuals have higher resting metabolic rates at thermoneutrality (RMRt) than slow growers. Moreover, the link between growth rate and daily energy expenditure (DEE) has never been tested in a wild endotherm. We assessed the energetic and survival costs of growth in juvenile eastern chipmunks (Tamias striatus) during a year of low food abundance by quantifying post-emergent growth rate (n = 88), RMRt (n = 66), DEE (n = 20), and overwinter survival. Both RMRt and DEE were significantly and positively related to growth rate. The effect size was stronger for DEE than RMRt, suggesting that the energy cost of growth in wild animals is more likely to be related to the maintenance of a higher foraging rate (included in DEE) than to tissue accretion (included in RMRt). Fast growers were significantly less likely to survive the following winter compared to slow growers. Juveniles with high or low RMRt were less likely to survive winter than juveniles with intermediate RMRt. In contrast, DEE was unrelated to survival. In addition, botfly parasitism simultaneously decreased growth rate and survival, suggesting that the energetic budget of juveniles was restricted by the simultaneous costs of growth and parasitism. Although the biology of the species (seed-storing hibernator) and the context of our study (constraining environmental conditions) were ideally combined to reveal a direct relationship between current use of energy and future availability, it remains unclear whether the energetic cost of growth was directly responsible for reduced survival.     

Energetics of the annual cycle of Dippers Cinclus cinclus

Time-activity budgets and energy expenditure of Dippers Cinclus cinclus were studied in all months of the year and for every stage of the annual cycle. The commonest daytime activity was feeding (54%), then resting (43%) and flying (4%). On a 24-hr day basis the most marked changes in activity followed from changing daylengths. DEE (Daily Energy Expenditure), derived from time-activity budgets through the year and laboratory estimates of metabolism, averaged 201 kj d ^-1 in females and 228 kj d ^-1 in the larger males. Over a more restricted range of circumstances, direct estimates of DEE obtained from 77 Dippers using the doubly labelled water technique averaged 205 ± 43 kj d^-1 and 251 ± 55 kj d^-1 in females and males, respectively. Overall, the correspondence between these largely independent estimates of energy expenditure was reasonably close. DEE was highest during breeding (laying-females; rearing-males) and in late winter for both sexes. The lowest energy expenditures occurred during moult, amongst juveniles and in early winter. Incubating females and mate-guarding males also had low energy costs. Across all stages of the annual cycle body size, activity patterns, ambient temperature and river flow had significant effects on energy expenditure. The rate at which food was gathered to meet these changing energy demands varied widely. While some of this variation was imposed by a seasonal environment, it was also likely to reflect adaptive shifts in rates of food gathering, in some cases consequent upon the changing fitness benefits of various non-feeding activities.     

Seasonal variation in energy expenditure and body composition in captive White Storks (Ciconia ciconia)

North Western European populations of White Storks (Ciconia ciconia) appear to have been saved from extinction by settling, i.e. stopping migration. Settled storks exposed to winter conditions must cope with periods of potentially high energy demands that would otherwise be avoided by the migration process. Doubly labeled water (DLW) was therefore used to examine the seasonal variation (summer vs winter) in daily energy expenditure (DEE) and the body composition of adult and immature storks of both sexes. Male White Storks showed a higher DEE over the winter period than in summer compared with females; in particular, immature males exhibited greater energy expenditure in winter than adult males. Thus, the DEE did not significantly differ between summer and winter (except for immature males), reflecting an absence of thermoregulation cost in winter. For both age classes, total body mass increased in winter, which was mainly due to an increase in fat mass. Adult storks were 5% heavier than immature storks. The sexes differed in body mass, with males weighing significantly more than females by 11%. Mean LBM (lean body mass) was 8.5% higher in adults than in immatures, and was 11.5% higher in males compared with females. Between their first and second summers, immatures accumulated a lean body mass to finally reach the same values as adults, indicating a phase of muscle development. The mean fat mass of the storks did not differ between age classes or between sexes. Based on physiological parameters, this study shows that settled White Storks are able to cope with mild winter periods when they are artificially provided with food. In a view to preserve favourable habitats for this species, it is therefore necessary to decide on a plan of action for breeding areas.