Sleep Posture Influences Metabolic Rate And Vigilance in The Common Whitethroat (Curruca Communis).

Migration is an important life-history strategy that is adopted by a significant proportion of bird species from temperate areas. Birds initiate migration after accumulating considerable energy reserves, primarily in the form of fat and muscle. Sustained exercise, such as during the crossing of ecological barriers, leads to the depletion of energy reserves and increased physiological stress. Stopover sites, where birds rest and restore energy, play a fundamental role in mitigating these challenges. The duration of resting at stopover sites is influenced by environmental and physiological conditions upon arrival, and the amount of body fat reserves plays an important role. While sleep is recognized as essential for all organisms, its importance is accentuated during migration, where energy management becomes a survival constraint. Previous research indicated that individuals with larger fat reserves tend to sleep less and favor an untucked sleep posture, influencing energy recovery and anti-predatory vigilance. We explored the relationship between sleep behavior and posture, metabolic state, and energy conservation strategies during migration in the common whitethroat (Curruca communis). We were able to confirm that sleeping in a tucked position results in metabolic energy savings, at the cost of reduced vigilance. However, whitethroats did not show alterations of their sleep patterns as a response to the amount of stored reserves. This suggests that they may not be taking full advantage of the metabolic gains of sleeping in a tucked posture, at least at this stage of their migratory journey. We suggest that, to achieve optimal fuel accumulation and maximize stopover efficiency, whitethroats prioritize increased foraging over modulating their sleep patterns.

Controlled expression of avian pre-migratory fattening influences indices of innate immunity.

While immunity is frequently dampened when birds engage in strenuous migratory flights, whether and how immunity changes during the rapid accumulation of energy stores in preparation for migration remains largely unknown. Here we induced pre-migratory fattening through controlled changes of daylight in common quails (Coturnix coturnix) and regularly assessed changes in three markers of constitutive innate immunity (leukocyte coping capacity or LCC, hemagglutination and hemolysis titres) and measures of body composition (lean and fat mass). All the three markers showed similar changes over the pre-migratory fattening process. LCC responses, hemagglutination titres, and hemolysis titres, were on average higher in the mid-fattening phase compared to the peak-fattening phase, when values were similar to those observed prior the start of pre-migratory fattening. At mid-fattening, we found that the birds that showed a larger accumulation of fat mass (as % of body mass) had lower LCC peak responses and hemolysis titres. Reversibly, at mid-fattening, we also found that the birds that kept a higher proportion of lean mass (as % of body mass) had the highest LCC peaks. Our results indicate that migratory birds undergo changes in immune indices (over 8 weeks) as they accumulate energy stores for migration and propose that this could be due to competing or trade-off processes between metabolic remodelling and innate immune system function.

Co-migration fidelity at a stopover site increases over time in African-European migratory landbirds.

Migratory species are changing their timing of departure from wintering areas and arrival to breeding sites (i.e. migration phenology) in response to climate change to exploit maximum food availability at higher latitudes and improve their fitness. Despite the impact of changing migration phenology at population and community level, the extent to which individual and species-specific response affects associations among co-migrating species has been seldom explored. By applying temporal co-occurrence network models on 15 years of standardized bird ringing data at a spring stopover site, we show that African-European migratory landbirds tend to migrate in well-defined groups of species with high temporal overlap. Such 'co-migration fidelity' significantly increased over the years and was higher in long-distance (trans-Saharan) than in short-distance (North African) migrants. Our findings suggest non-random patterns of associations in co-migrating species, possibly related to the existence of regulatory mechanisms associated with changing climate conditions and different uses of stopover sites, ultimately influencing the global economy of migration of landbirds in the Palearctic-African migration system.

Physiologically vulnerable or resilient? Tropical birds, global warming, and redistributions.

Tropical species are considered to be more threatened by climate change than those of other world regions. This increased sensitivity to warming is thought to stem from the assumptions of low physiological capacity to withstand temperature fluctuations and already living near their limits of heat tolerance under current climatic conditions. For birds, despite thorough documentation of community-level rearrangements, such as biotic attrition and elevational shifts, there is no consistent evidence of direct physiological sensitivity to warming. In this review, we provide an integrative outlook into the physiological response of tropical birds to thermal variation and their capacity to cope with warming. In short, evidence from the literature suggests that the assumed physiological sensitivity to warming attributed to tropical biotas does not seem to be a fundamental characteristic of tropical birds. Tropical birds do possess the physiological capacities to deal with fluctuating temperatures, including high-elevation species, and are prepared to withstand elevated levels of heat, even those living in hot and arid environments. However, there are still many unaddressed points that hinder a more complete understanding of the response of tropical birds to warming, such as cooling capacities when exposed to combined gradients of heat and humidity, the response of montane species to heat, and thermoregulation under increased levels of microclimatic stress in disturbed ecosystems. Further research into how populations and species from different ecological contexts handle warming will increase our understanding of current and future community rearrangements in tropical birds.

Comparing population trend estimates of migratory birds from breeding censuses and capture data at a spring migration bottleneck.

Europe has a well-established network of breeding bird monitoring that is used to produce supranational indices of population trends for many species. However, a comparison of breeding bird censuses with other methods may be beneficial to confirm the validity of such indices. The aim of this study was to assess the value of standardized capture data of migratory birds at migration bottlenecks as an indicator of the effective breeding populations. One limitation to this method is that several populations are co-occurring at these bottlenecks and their catchment areas need to be clearly identified to allow extrapolation of population indices. Here, we used standardized trends in capture numbers of 30 species on the island of Ponza, a migration bottleneck in the central Mediterranean, and compared them to population trends estimated in the putative catchment breeding areas between 2005 and 2016. The catchment areas were identified through the analysis of ring recoveries during the breeding season of birds passing through Ponza. Our results show an agreement between the population trends observed on Ponza and those in the breeding areas in 15 out of 30 species. The correlations were strongest in species with a more robust definition of the catchment areas, that is, species with more than 10 recoveries, and for which the recoveries were most likely of breeding birds. The main reason for disagreement between the two indices in the remaining species might be related to different intensity of sampling in different areas. This issue can be solved by further developing monitoring projects in underrepresented countries, as well as by intensifying monitoring through ringing, both in the breeding grounds and at migration bottlenecks. These results show that spring migration monitoring at bottlenecks has the potential to provide a valuable complement and an independent control of breeding bird surveys, allowing raising early warnings of population declines and contributing to their conservation.

Heat loss in sleeping garden warblers (Sylvia borin) during migration.

For small songbirds, energy is often a limiting factor during migration and, for this reason, they are forced to alternate nocturnal flights with stopovers to rest and replenish energy stores. Stopover duration has a key role for a successful migration and may have an important impact on fitness. Thus, migrants need to optimize their energy consumption at this stage to reduce their permanence at the site. A recent study has shown that lean individuals reduce their metabolic rate when tucking the head in the feathers during sleep. The underlying mechanism is very likely a reduction in conductance, but the thermoregulatory benefit of the increased insulation has never been quantified yet. Here, we compared heat loss in individual migratory birds while sleeping in different postures. Using a thermal camera and a within-individual approach, we estimated that Garden Warblers can reduce their rate of heat loss by 54% by sleeping with the head tucked in the feathers. This energy saving has a relevant impact on the individual's energy balance because it can account for up to 8.69% of daily energy expenditure during stopover. Our study provides novel and important information to understand the fundamental role of thermoregulatory strategies on bird's energy management.

Individual variability and versatility in an eco-evolutionary model of avian migration.

Seasonal migration is a complex and variable behaviour with the potential to promote reproductive isolation. In Eurasian blackcaps (Sylvia atricapilla), a migratory divide in central Europe separating populations with southwest (SW) and southeast (SE) autumn routes may facilitate isolation, and individuals using new wintering areas in Britain show divergence from Mediterranean winterers. We tracked 100 blackcaps in the wild to characterize these strategies. Blackcaps to the west and east of the divide used predominantly SW and SE directions, respectively, but close to the contact zone many individuals took intermediate (S) routes. At 14.0° E, we documented a sharp transition from SW to SE migratory directions across only 27 (10-86) km, implying a strong selection gradient across the divide. Blackcaps wintering in Britain took northwesterly migration routes from continental European breeding grounds. They originated from a surprisingly extensive area, spanning 2000 km of the breeding range. British winterers bred in sympatry with SW-bound migrants but arrived 9.8 days earlier on the breeding grounds, suggesting some potential for assortative mating by timing. Overall, our data reveal complex variation in songbird migration and suggest that selection can maintain variation in migration direction across short distances while enabling the spread of a novel strategy across a wide range.

Recent phenological shifts of migratory birds at a Mediterranean spring stopover site: Species wintering in the Sahel advance passage more than tropical winterers.

Spring migration phenology is shifting towards earlier dates as a response to climate change in many bird species. However, the patterns of change might not be the same for all species, populations, sex and age classes. In particular, patterns of change could differ between species with different ecology. We analyzed 18 years of standardized bird capture data at a spring stopover site on the island of Ponza, Italy, to determine species-specific rates of phenological change for 30 species following the crossing of the Mediterranean Sea. The advancement of spring passage was more pronounced in species wintering in Northern Africa (i.e. short-distance migrants) and in the Sahel zone. Only males from species wintering further South in the forests of central Africa advanced their passage, with no effect on the overall peak date of passage of the species. The migration window on Ponza broadened in many species, suggesting that early migrants within a species are advancing their migration more than late migrants. These data suggest that the cues available to the birds to adjust departure might be changing at different rates depending on wintering location and habitat, or that early migrants of different species might be responding differently to changing conditions along the route. However, more data on departure time from the wintering areas are required to understand the mechanisms underlying such phenological changes.

Energy Stores, Oxidative Balance, and Sleep in Migratory Garden Warblers (Sylvia borin) and Whitethroats (Sylvia communis) at a Spring Stopover Site.

Little is known about how songbirds modulate sleep during migratory periods. Due to the alternation of nocturnal endurance flights and diurnal refueling stopovers, sleep is likely to be a major constraint for many migratory passerine species. Sleep may help to increase the endogenous antioxidant capacity that counteracts free radicals produced during endurance flight and reduces energy expenditure. Here, we investigated the relationship between sleep behavior, food intake, and two markers of physiological condition-the amount of energy reserves and oxidative status-in two migratory songbird species, the garden warbler (Sylvia borin) and the whitethroat (Sylvia communis). In garden warblers, birds with high energy stores were more prone to sleep during the day, while this condition-dependent sleep pattern was not present in whitethroats. In both species, birds with low energy stores were more likely to sleep with their head tucked in the feathers during nocturnal sleep. Moreover, we found a positive correlation between food intake and the extent of energy reserves in garden warblers, but not in whitethroats. Finally, we did not find significant correlations between oxidative status and sleep, or oxidative status and energy stores. Despite our study was not comparative, it suggests that different species might use different strategies to manage their energy during stopover and, additionally, it raises the possibility that migrants have evolved physiological adaptations to deal with oxidative damage produced during migration.

Il pattern del sonno nei passeriformi migratori è praticamente sconosciuto. Durante la migrazione, il sonno può diventare una importante limitazione fisiologica per molte specie che alternano estenuanti voli notturni con intensi foraggiamenti diurni per recuperare le energie. Oltre a ridurre il dispendio energetico, il sonno potrebbe promuovere le capacità antiossidanti endogene che contrastano i radicali liberi che si accumulano durante i voli prolungati. In questo studio, abbiamo indagato la relazione tra sonno, assunzione di cibo, e due marcatori della condizione fisiologica­la quantità di riserve di energia e lo stato ossidativo­in due specie di passeriformi migratori, il Beccafico (Sylvia borin) e la Sterpazzola (Sylvia communis). Nei Beccafichi, gli individui con abbondanti riserve energetiche erano più inclini a dormire durante il giorno che durante la notte, mentre non è stata trovata una associazione significativa tra il pattern di sonno e le riserve energetiche nelle Sterpazzole. In entrambe le specie, gli uccelli con scarse riserve energetiche erano più propensi a dormire con la testa inserita tra le penne durante la notte. Inoltre, abbiamo trovato una correlazione positiva tra la quantità di cibo ingerito e le riserve energetiche alla cattura nei Beccafichi, ma non nelle Sterpazzole. Infine, non abbiamo trovato correlazioni significative dello stato ossidativo con il pattern del sonno e le riserve energetiche. Questi risultati suggeriscono che specie diverse potrebbero utilizzare strategie differenti per gestire la loro energia durante la sosta, e indicano che i passeriformi migratori posseggono adattamenti fisiologici per limitare i danni ossidativi prodotti durante la migrazione. Translated to Italian by Leonida Fusani and Andrea Ferretti.

The amount of available food affects diurnal locomotor activity in migratory songbirds during stopover.

Migratory passerine birds fly long distances twice a year alternating nocturnal flights with stopovers to rest and replenish energy stores. The duration of each stopover depends on several factors including internal clocks, meteorological conditions, and environmental factors such as availability of food. Foraging entails energetic costs, and if birds need to refuel efficiently, they should modulate their activity in relation to food availability. We investigated how food availability influences locomotor activity in migrating birds of six passerine species at a spring stopover site in the central Mediterranean Sea. We selected birds with low fat scores which we expected to be strongly motivated to refuel. We simulated stopover sites of different quality by providing temporarily caged birds with different amounts of food to simulate scarce to abundant food. We analysed the diurnal locomotory activity as a proxy for food searching effort. Low food availability resulted in an increased diurnal locomotor activity in almost all species, while all birds showed low intensity of nocturnal migratory restlessness. In conclusion, our study shows that food availability in an important determinant of behaviour of migratory birds at stopover sites.

Endogenous control of fuelling in a migratory songbird.

The Northern wheatear (Oenanthe oenanthe) is a small long-distance migratory songbird that breeds throughout the Northern hemisphere and winters in sub-Saharan Africa. The main components of its migratory behaviour, i.e. seasonal migratory restlessness and body mass changes, have been shown to be under endogenous control. However, it is still unknown whether the disposition to accumulate fuel reserves is an inherited trait. We cross-bred Northern wheatears from two populations known to accumulate different amounts of fuel in a common-garden setup, and measured their maximum fuelling as the difference between the lightest and heaviest body mass recorded over each year for 4 years or longer. We used the largest value as a measure of maximum voluntary fuelling potential. F1-generation "hybrids" showed intermediate values to those of the parent populations. It was previously shown that in the wheatear the amount of fuel accumulated is closely linked to the presence of large ecological barriers to cross. This study shows that this adaptation has been fixed at the genetic level, and that intermediate traits are transferred to the next generation, with possible implications on the viability of such individuals in nature.

Light oiling of feathers increases flight energy expenditure in a migratory shorebird.

Flying birds depend on their feathers to undertake most activities, and maintain them in peak condition through periodic molt and frequent preening. Even small exposures to crude oil reduce the integrity of feathers, and could impair flight performance. We trained wild western sandpipers (Calidris mauri) to perform endurance flights in a wind tunnel, and used magnetic resonance body composition analysis to measure energy expenditure after birds were exposed to weathered MC252 crude oil from the Deepwater Horizon oil spill. The cost of transport was 0.26±0.04 kJ km-1 in controls, and increased by 22% when the trailing edges of the wing and tail were oiled (<20% of body surface; considered light oiling). Additional crude oil on breast and back feathers (∼30% total surface; moderate oiling) increased the cost of transport by 45% above controls. Oiling tended to decrease flight control, and only half of moderately oiled birds completed the flight test. We then flew birds at a range of speeds to estimate basic kinematic parameters. At low speeds, light and moderately oiled birds had larger wingbeat amplitudes than controls, while moderately oiled birds showed greater wingbeat frequencies across all speeds, and a shift in optimal flight speed towards higher wind speeds. We suggest these changes reflect poorer lift production and increased drag on the wings and body. Oiling will increase the difficulty and energy costs of locomotion for daily and seasonal activities such as foraging, predator evasion, territory defense, courtship, chick provisioning, commuting and long-distance migration. These sub-lethal effects must be considered in oil spill impact assessments.

Reprint of: Trouble on takeoff: Crude oil on feathers reduces escape performance of shorebirds.

The ability to takeoff quickly and accelerate away from predators is crucial to bird survival. Crude oil can disrupt the fine structure and function of feathers, and here we tested for the first time how small amounts of oil on the trailing edges of the wings and tail of Western sandpipers (Calidris mauri) affected takeoff flight performance. In oiled birds, the distance travelled during the first 0.4s after takeoff was reduced by 29%, and takeoff angle was decreased by 10° compared to unoiled birds. Three-axis accelerometry indicated that oiled sandpipers produced less mechanical power output per wingbeat during the initial phase of flight. Slower and lower takeoff would make oiled birds more likely to be targeted and captured by predators, reducing survival and facilitating the exposure of predators to oil. Whereas the direct mortality of heavily-oiled birds is often obvious and can be quantified, our results show that there are significant sub-lethal effects of small amounts crude oil on feathers, which must be considered in natural resource injury assessments for birds.

Toxicological and thermoregulatory effects of feather contamination with artificially weathered MC 252 oil in western sandpipers (Calidris mauri).

The external contamination of bird feathers with crude oil might have effects on feather structure and thus on thermoregulation. We tested the thermoregulatory ability of western sandpipers (Calidris mauri) in a respirometry chamber with oil applied either immediately prior, or three days before the experiment. The birds were then exposed to a sliding cold temperature challenge between 27°C and -3°C to calculate thermal conductance. After the experiment, a large blood sample was taken and the liver extracted to measure a range of parameters linked to toxicology and oxidative stress. No differences in thermal conductance were observed among groups, but birds exposed to oil for three days had reduced body temperatures and lost more body mass during that period. At necropsy, oiled birds showed a decrease in plasma albumin and sodium, and an increase in urea. This is reflective of dysfunction in the kidney at the loop of Henle. Birds, especially when exposed to the oil for three days, showed signs of oxidative stress and oxidative damage. These results show that the ingestion of externally applied oil through preening or drinking can cause toxic effects even in low doses, while we did not detect a direct effect of the external oil on thermoregulation over the temperature range tested.

Effect of oral exposure to artificially weathered Deepwater Horizon crude oil on blood chemistries, hepatic antioxidant enzyme activities, organ weights and histopathology in western sandpipers (Calidris mauri).

Shorebirds were among birds exposed to Mississippi Canyon 252 (MC252) crude oil during the 2010 Deep Water Horizon (DWH) oil spill in the Gulf of Mexico. The western sandpiper (Calidris mauri) was chosen as one of four species for initial oral dosing studies conducted under Phase 2 of the avian toxicity studies for the DWH Natural Resource Damage Assessment (NRDA). Thirty western sandpipers were assigned to one of three treatment groups, 10 birds per group. The control group was sham gavaged and the treatment groups were gavaged with 1 or 5mL oil kg bw-1 daily for 20 days. Periodic blood samples for hemoglobin measurements were collected during the trial. A final blood sample used to determine hemoglobin concentration in addition to complete blood counts, plasma clinical chemistries, haptoglobin concentration and plasma electrophoresis was collected when birds were euthanized and necropsied on day 21. Tissues were removed, weighed and processed for subsequent histopathological evaluation. There were numerical decreases in hemoglobin concentrations in oil-dosed birds over the 21-day trial, but values were not significantly different compared to controls on day 21. There were no significant differences between controls and oiled birds in complete blood counts, plasma chemistries, haptoglobin concentration, and plasma electrophoresis endpoints. Of the hepatic oxidative stress endpoints assessed, the total antioxidant capacity assessment (Trolox equivalents) for the control group was lower compared to the 1mL oil kg bw-1 group. Absolute liver weights in the 5mL oil kg bw-1 group were significantly greater compared to controls. While not conclusive, the numerical decrease in hemoglobin concentration and significant increase in absolute liver weight are consistent with exposure to oil. Histological changes in the adrenal gland could be considered a non-specific indicator of stress resulting from exposure to oil. It is possible that the quantity of oil absorbed was not sufficient to induce clearly evident hemolytic anemia or that the western sandpiper is relatively insensitive to ingested oil.

Trouble on takeoff: Crude oil on feathers reduces escape performance of shorebirds.

The ability to takeoff quickly and accelerate away from predators is crucial to bird survival. Crude oil can disrupt the fine structure and function of feathers, and here we tested for the first time how small amounts of oil on the trailing edges of the wings and tail of Western sandpipers (Calidris mauri) affected takeoff flight performance. In oiled birds, the distance travelled during the first 0.4s after takeoff was reduced by 29%, and takeoff angle was decreased by 10° compared to unoiled birds. Three-axis accelerometry indicated that oiled sandpipers produced less mechanical power output per wingbeat during the initial phase of flight. Slower and lower takeoff would make oiled birds more likely to be targeted and captured by predators, reducing survival and facilitating the exposure of predators to oil. Whereas the direct mortality of heavily-oiled birds is often obvious and can be quantified, our results show that there are significant sub-lethal effects of small amounts crude oil on feathers, which must be considered in natural resource injury assessments for birds.

Morphometrics and stable isotopes differentiate wintering populations of a migratory bird.

BACKGROUND: Describing migratory connectivity in mobile animals is crucial for understanding the selective pressures acting on different populations throughout their life cycle. Tracking single individuals has provided valuable data, but for most species the data available are still spurious and usually limited to a few individuals. Since different populations of migratory birds can be distinguished by a combination of morphometric measurements and the isotopic composition of their feathers, it is possible to measure these parameters on a large sample to differentiate populations. METHODS: We studied northern wheatears, Oenanthe oenanthe, captured in their African wintering range and applied discriminant analyses on morphometric measurements and stable isotope signatures to determine whether birds found in different areas were distinguishable from each other. RESULTS: Morphometric and isotopic measurements alone were not sufficient to discriminate between the birds of ssp. oenanthe from different areas in Africa. When combining the two measurements, however, assignment to the different groups became substantially more accurate. Following the discriminant analysis of morphometrics and δ(2)H, δ(13)C, and δ(15)N isotopes signatures, 19 of 20 oenanthe from Kenya, 15 of 20 oenanthe from Mali/Mauritania, and 19 of 20 oenanthe from Niger were assigned correctly to their wintering area. CONCLUSIONS: Our results show that birds at different wintering sites can be distinguished from each other when using a combination of markers. We discuss the possible breeding origins of these wintering birds.

Stopover optimization in a long-distance migrant: the role of fuel load and nocturnal take-off time in Alaskan northern wheatears (Oenanthe oenanthe).

INTRODUCTION: In long-distance migrants, a considerably higher proportion of time and energy is allocated to stopovers rather than to flights. Stopover duration and departure decisions affect consequently subsequent flight stages and overall speed of migration. In Arctic nocturnal songbird migrants the trade-off between a relatively long migration distance and short nights available for travelling may impose a significant time pressure on migrants. Therefore, we hypothesize that Alaskan northern wheatears (Oenanthe oenanthe) use a time-minimizing migration strategy to reach their African wintering area 15,000 km away. RESULTS: We estimated the factors influencing the birds' daily departure probability from an Arctic stopover before crossing the Bering Strait by using a Cormack-Jolly-Seber model. To identify in which direction and when migration was resumed departing birds were radio-tracked. Here we show that Alaskan northern wheatears did not behave as strict time minimizers, because their departure fuel load was unrelated to fuel deposition rate. All birds departed with more fuel load than necessary for the sea crossing. Departure probability increased with stopover duration, evening fuel load and decreasing temperature. Birds took-off towards southwest and hence, followed in general the constant magnetic and geographic course but not the alternative great circle route. Nocturnal departure times were concentrated immediately after sunset. CONCLUSION: Although birds did not behave like time-minimizers in respect of the optimal migration strategies their surplus of fuel load clearly contradicted an energy saving strategy in terms of the minimization of overall energy cost of transport. The observed low variation in nocturnal take-off time in relation to local night length compared to similar studies in the temperate zone revealed that migrants have an innate ability to respond to changes in the external cue of night length. Likely, birds maximized their potential nightly flight range by taking off early in the night which in turn maximizes their overall migration speed. Hence, nocturnal departure time may be a crucial parameter shaping the speed of migration indicating the significance of its integration in future migration models.

Innate sex differences in the timing of spring migration in a songbird.

In migrating animals protandry is the phenomenon whereby males of a species arrive at the breeding grounds earlier than females. In the present study we investigated the proximate causes of protandry in a migratory songbird, the northern wheatear Oenanthe oenanthe. Previous experiments with caged birds revealed that males and females show differentiated photoperiod-induced migratory habits. However, it remained open whether protandry would still occur without photoperiodic cues. In this study we kept captive first-year birds under constant photoperiod and environmental conditions in a "common garden" experiment. Male northern wheatears started their spring migratory activity earlier than females, even in the absence of environmental cues. This indicates that protandry in the northern wheatear has an endogenous basis with an innate earlier spring departure of males than females.

Endogenous rhythms of seasonal migratory body mass changes and nocturnal restlessness in different populations of Northern Wheatear Oenanthe oenanthe.

The Northern Wheatear (Oenanthe oenanthe) is a migratory bird species that shows different strategies of migration between populations, adapted to cope with different ecological barriers. This raises the question whether and to which extent these adaptations are endogenously determined. We studied seasonal patterns of body mass change and nocturnal restlessness in wheatears from Iceland, which face an initial sea crossing of at least 800 km; from Norway, which fly a similar distance as Icelandic birds but without a long sea crossing; and from Morocco, which fly a shorter distance to reach their wintering grounds. To isolate the endogenous component of the regulation of these migratory traits, we kept the wheatears in a "common garden," all 3 populations experiencing the same environmental conditions and a constant photoperiod during their first year of life. Icelandic birds showed a greater increase of their body mass in autumn than the other 2 populations, indicating preparation for the initial barrier crossing. The autumnal timing of nocturnal restlessness and the total activity during autumn were related to the distance to be covered, although the differences between populations were smaller than expected. In all 3 populations, body mass increased to a greater extent in autumn than in spring, whereas nocturnal activity was higher in spring than in autumn. This suggests that the endogenous program responds to specific seasonal needs, with more time invested in storing fuel for a safe journey in autumn and more time invested in flying to reach the breeding grounds early in spring. Contrary to expectations, the timing of onset of body mass increase and nocturnal restlessness in spring did not differ between populations. This might be explained by the lack of external cues, most likely photoperiod, which are responsible for the fine tuning of the expression of migratory behavior.