Energy reserves stored by migrating Gray‐cheeked Thrushes Catharus minimus at a spring stopover site in northern Colombia are sufficient for a long‐distance flight to North America

Stopover sites used to accumulate the energy that fuels migration, especially those used prior to crossing ecological barriers, are regarded as critically important for the survival of Nearctic?Neotropical migratory birds. To assess whether South American stopover sites are used to store the energy required to cross the Caribbean Sea and the Gulf of Mexico to North America by a Neotropical migratory landbird, we studied Gray‐cheeked Thrushes in northern Colombia through constant effort mist‐netting during spring migration in 2010 and 2011. We combined stopover duration estimates and models of body mass change based on recaptures to estimate departure body mass and potential flight range from our study site. We recaptured 62 birds, the majority of which gained mass. Models indicated significant differences in rates of mass gain between years and age groups and with arrival date. Estimated total stopover durations varied between 15.4 (2010) and 12.5 days (2011). Predicted departure mass ranged between 41.3 and 44.9 g, and potential flight range was estimated at between 2727 and 4270 km. Gray‐cheeked Thrushes therefore departed our study site with sufficient energy reserves to cross the Caribbean Sea and the Gulf of Mexico (2550 km). As the first demonstration that birds departing from South American stopover sites can reach North America without refuelling, this has important implications for stopover site protection. Strategic conservation measures in the Sierra Nevada de Santa Marta could protect habitats in which up to 40% of the energy required to complete spring migration is stored by a Neotropical migratory land bird.     

Wide ranging stopover movements and substantial fuelling in first year garden warblers at a northern stopover site

Migratory birds use stopovers to replenish their fuel reserves and they generally spend more time at stopover sites than they do in actual flight. When arriving at a new stopover site birds may need to search extensively to find a suitable feeding area and this search and settling period may affect the duration of stopover. Stopover behaviour can thus have profound effects on the migratory programme and studies on stopover behaviour are important to understand migratory strategies. We followed 51 first‐year garden warblers Sylvia borin with radio‐transmitters at an autumn stopover site on the island of Gotland in southern Sweden. Our aim was to determine the distance birds relocated from the coastal capture site when searching for an area to settle in, and also to establish the duration of stopover and put it in relation to refuelling rate by recapturing a subset of the radio‐tracked individuals. Sixteen birds made an extended stopover (> 2 d), relocated inland from the capture site and settled on average 5.6 km from the capture site, with the longest recorded relocation being fourteen kilometres. Birds that relocated nocturnally settled in areas further away than birds that relocated diurnally. Thirteen birds that continued migration after a short stop carried larger fuel stores than birds that stopped over longer and they remained close to the capture site until departure. Three birds were re‐trapped and showed high fuelling rates, between 0.3 and 1.1 g d^–1. They left the stopover site with fuel loads between 40–56 percent of lean body mass, which possibly would have allowed them to reach the Mediterranean area without additional refuelling stops.     

Bats on a Budget: Torpor-Assisted Migration Saves Time and Energy

Bats and birds must balance time and energy budgets during migration. Migrating bats face similar physiological challenges to birds, but nocturnality creates special challenges for bats, such as a conflict between travelling and refueling, which many birds avoid by feeding in daylight and flying at night. As endothermic animals, bats and birds alike must expend substantial amounts of energy to maintain high body temperatures. For migratory birds refueling at stopovers, remaining euthermic during inactive periods reduces the net refuelling rate, thereby prolonging stopover duration and delaying subsequent movement. We hypothesized that bats could mitigate similar ambient-temperature dependent costs by using a torpor-assisted migration strategy. We studied silver-haired bats Lasionycteris noctivagans during autumn migration using a combination of respirometry and temperature-sensitive radiotelemetry to estimate energy costs incurred under ambient temperature conditions, and the energy that bats saved by using torpor during daytime roosting periods. All bats, regardless of sex, age, or body condition used torpor at stopover and saved up to 91% of the energy they would have expended to remain euthermic. Furthermore, bats modulated use of torpor depending on ambient temperature. By adjusting the time spent torpid, bats achieved a rate of energy expenditure independent of the ambient temperature encountered at stopover. By lowering body temperature during inactive periods, fuel stores are spared, reducing the need for refuelling. Optimal migration models consider trade-offs between time and energy. Heterothermy provides a physiological strategy that allows bats to conserve energy without paying a time penalty as they migrate. Although uncommon, some avian lineages are known to use heterothermy, and current theoretical models of migration may not be appropriate for these groups. We propose that thermoregulatory strategies should be an important consideration of future migration studies of both bats and birds.     

Population‐wide body mass increment at stopover sites is an unreliable indicator of refuelling rates in migrating waders

Estimates of refuelling rates in migrating waders are best based on intra‐seasonal recaptures of individually marked birds. This method, however, has methodological problems associated with capture effects and difficulties in attaining sufficient sample sizes. An alternative method had been proposed whereby refuelling rates are approximated by the body mass increment from the slope of the regression between body masses of all birds caught at a site and date. We investigated the appropriateness of this indicator with a simulation study in non‐synchronized migratory species (i.e. arrivals and departures of individuals at the stopover site are not synchronized). Simulation results indicated that the mass increment across the population may be used as an approximation of refuelling rate only in populations with low turnover rates (percentage of birds arriving at/departing from stopover site per day <2%) and ideally with constant numbers of staging birds. The mass increment of non‐synchronized populations with moderate or high turnover rates (higher than 5%) depends mainly on body masses of arriving birds and is not indicative of the individual rate of refuelling. The results of the simulation study were confirmed with empirical data gathered from Wood Sandpiper Tringa glareola and Common Snipe Gallinago gallinago during their autumn migration at a stopover site in Poland. The population mass increment methods considerably underestimated refuelling rate obtained from the recapture‐based approach of individual birds in both species. As a consequence, we suggest that population mass increment should not be used as an indicator of refuelling rates in non‐synchronized stopover populations of migrating waders.     

Landscape movements of migratory birds and bats reveal an expanded scale of stopover

Many species of birds and bats undertake seasonal migrations between breeding and over-wintering sites. En-route, migrants alternate periods of flight with time spent at stopover--the time and space where individuals rest and refuel for subsequent flights. We assessed the spatial scale of movements made by migrants during stopover by using an array of automated telemetry receivers with multiple antennae to track the daily location of individuals over a geographic area ~20 × 40 km. We tracked the movements of 322 individuals of seven migratory vertebrate species (5 passerines, 1 owl and 1 bat) during spring and fall migratory stopover on and adjacent to a large lake peninsula. Our results show that many individuals leaving their capture site relocate within the same landscape at some point during stopover, moving as much as 30 km distant from their site of initial capture. We show that many apparent nocturnal departures from stopover sites are not a resumption of migration in the strictest sense, but are instead relocations that represent continued stopover at a broader spatial scale.     

Long-distance travellers stopover for longer: a case study with spoonbills staying in North Iberia

Long-distance migration is widespread among birds, connecting breeding and wintering areas through a set of stopover localities where individuals refuel and/or rest. The extent of the stopover is critical in determining the migratory strategy of a bird. Here, we examined the relationship between minimum length of stay of PVC-ringed birds in a major stopover site and the remaining flight distance to the overwintering area in the Eurasian spoonbill (Platalea l. leucorodia) during four consecutive autumn migrations. We also analysed the potential effect of timing (arrival date), as well as the role of experience in explaining stopover duration of spoonbills. Overall, birds wintering in Africa, and facing long-distance travel from the stopover site (ca. 3,000 km) stay for longer (2.7 ± 0.4 days) than Iberian winterers (1.5 ± 0.2 days) that perform a much shorter migration (ca. 800 km). These differences were consistent between years. Stopover duration was not significantly affected by the age of the bird. However, there was a significant reduction as migration advanced. Our results suggest that spoonbills develop different stopover strategies depending on the expected distance to the wintering grounds. Adults, especially long-distance migratory ones, could reduce the potential negative effects of density-dependence processes by avoiding stopover at the end of the migration period. These findings are of significant relevance for understanding differences in migratory behaviour within single populations, especially for declining waterbirds, as well as stress the relevance of preserving stopover localities for the conservation of intraspecific diversity in migratory species.     

Metabolic profile of long-distance migratory flight and stopover in a shorebird

Migrating birds often complete long non-stop flights during which body energy stores exclusively support energetic demands. The metabolic correlates of such long-distance travel in free-living migrants are as yet poorly studied. Bar-tailed godwits, Limosa lapponica taymyrensis, undertake a 4500 km flight to their single spring stopover site and thus provide an excellent model in which to determine the energy fuels associated with endurance travel. To this end, we evaluated plasma concentrations of six key metabolites in arriving godwits caught immediately upon landing near their stopover site. Initial metabolite levels were compared with levels after 5 h of inactive rest to determine how flight per se affects energy metabolism. Birds refuelling on the stopover site were also examined. Arriving godwits displayed elevated plasma free fatty acids, glycerol and butyrate, confirming the importance of lipid fuel in the support of extended migratory activity. Further-more, elevated plasma triglycerides in these birds suggest that fatty acid provisioning is facilitated through hepatic synthesis and release of neutral lipids, as previously hypothesized for small migrants with high mass-specific metabolic rates. Finally, elevations in plasma uric acid suggest that protein breakdown contributes to the support of long-distance movement, to possibly maintain citric acid cycle intermediates, gluconeogenesis and/or water balance.     

基于卫星跟踪大天鹅春季迁徙时间及其影响因素

候鸟春季迁徙时间与其能否顺利完成迁徙过程,以及对繁殖地的成功选择和繁殖成效密切相关,通过对大天鹅越冬地和停歇地的春季迁徙时间选择原因及其影响因素进行分析,有助于深入理解候鸟春季迁徙时间策略和栖息地保护状况。2015年2月和12月,在河南三门峡湿地捕捉了60只越冬大天鹅并佩戴卫星跟踪器,获取了详细的大天鹅在越冬地和停歇地的春季迁徙时间等信息,并进一步分析了春季迁徙时间与气候因子的相关性。结果表明,大天鹅主要在夜间22：00-0：00和0：00-1：59迁离越冬地。大天鹅迁离越冬地的日期与温度呈显著性正相关,迁离时多选择顺风且风速较小的天气。大天鹅迁离越冬地后主要是在夜间飞行,而迁离停歇地后主要是在日间飞行。越冬地的温度越高,越有利于植物的生长,大天鹅可以快速地积累能量,提前开始春季迁徙。野外调查表明,内蒙古黄河中上游作为迁徙的重要停歇地,春季两岸捕鱼、农耕和放牧等为活动较多,因此大天鹅多选择在夜间觅食补充能量,在日间迁离。最后,针对黄河流域大天鹅栖息地的保护现状,提出了禁止经济开发项目、建立保护区和开展宣传教育等保护建议。     

Body fat influences departure from stopover sites in migratory birds: evidence from whole-island telemetry

Migration remains one of the great mysteries of animal life. Small migratory birds rely on refuelling stopovers after crossing ecological barriers such as deserts or seas. Previous studies have suggested that fuel reserves may determine stopover duration but this hypothesis could not be tested because of methodological limitations. Here, we provide evidence that subcutaneous fat stores determine stopover duration by measuring the permanence of migratory garden warblers (Sylvia borin) on a small Mediterranean island during spring migration with telemetry methods. Garden warblers with large amounts of fat stores departed the island significantly sooner than lean birds. All except one fat bird left the island on the same evening after capture, with a mean total stopover estimate of 8.8 hours. In contrast, the mean estimated total stopover duration of lean birds was 41.3 hours. To our knowledge, this is the first study that measures the true minimum stopover duration of a songbird during migration.     

Landscape‐scale habitat availability,and not local geography,predicts migratory landbird stopover across the Gulf of Maine

While it is clear that many migratory behaviors are shared across taxa, generalizable models that predict the distribution and abundance of migrating taxa at the landscape scale are rare. In migratory landbirds, ephemeral concentrations of refueling birds indicate that individual behaviors sometimes produce large epiphenomena in particular geographic locations. Identifying landscape factors that predict the distribution and abundance of birds during migratory stopover will both improve our understanding of the migratory process and assist in broad, regionally relevant conservation. In this study we used autumnal passerine stopover data from a five‐year period and eleven stopover sites across coastal Maine, USA, to test four broad hypotheses of migrant distribution and abundance that have been supported in other regions: a) the community characteristics of the pool of potential migrants, b) a site's local geography, c) landscape composition and configuration measured at different spatial scales, and d) interactions between these factors. Our final model revealed that birds concentrate at ‘habitat islands’, sites that possess a disproportionate percentage of the vegetated habitat in the 4‐km surrounding landscape. The strength of this pattern, however, was inversely proportional to a species' remaining migratory distance. Our results corroborate several studies that emphasize the importance of land cover composition at finer spatial scales (< 80 km²) for predicting the stopover distribution and abundances of migratory birds. This suggests that different migrants likely assess stopover sites with similar mechanisms along their migratory route, and these commonalities may be broadly applied to identify stopover locations of conservation importance across the continent.     

Inland flights of young red‐eyed vireos Vireo olivaceus in relation to survival and habitat in a coastal stopover landscape

Inland dispersal of migrating land birds away from the coast, often opposite to the direction of migration, occurs frequently. Many of these movements may involve migrants seeking improved stopover conditions farther inland, but direct study of inland flights and of the ecological factors influencing their occurrence is limited. We used an automated telemetry array and ground‐tracking to assess flight behaviours, survival, and habitat use of young red‐eyed vireos Vireo olivaceus during fall migration at a coastal island and an inland stopover site in southwest Nova Scotia. We recorded inland flights for 41% (11/27) of individuals that departed the island. At least 25% of 16 individuals tagged at the inland site also relocated within the landscape prior to continuing migration, but due to the higher proportion of ambiguous flights at the inland site (44%) compared to the island (15%), we could not be sure if actual proportions of relocations differed between sites. Mortality on the island (at least 10 of 39 individuals) was significantly higher than at the inland site (0 of 16 individuals). At mainland sites near the coast where we found 6 of 11 individuals after they relocated away from the island, mortality remained high (2/6). Lack of deciduous canopy cover may have contributed to the high mortality on the island, but coastal mainland sites had a relatively high amount of deciduous canopy cover, similar to at the inland site where there was no mortality. Although coastal stopover sites may be important for migrating songbirds, especially before or after making a large water crossing, our results show that mortality can be much higher, and habitat poorer, at the coast compared to farther inland. Therefore, relocating inland may be an adaptive strategy for individuals that initially settle at the coast and that need to rest and refuel before they continue migration.     

African Odyssey project – satellite tracking of black storks Ciconia nigra breeding at a migratory divide

The African Odyssey project focuses on studying the migration of the black stork Ciconia nigra breeding at a migratory divide. In 1995–2001, a total of 18 black storks breeding in the Czech Republic were equipped with satellite (PTT) and VHF transmitters. Of them, 11 birds were tracked during at least one migration season and three birds were tracked repeatedly. The birds migrated either across western or eastern Europe to spend the winter in tropical west or east Africa, respectively. One of the juveniles made an intermediate route through Italy where it was shot during the first autumn migration. The mean distance of autumn migration was 6,227 km. The eastern route was significantly longer than the western one (7,000 km and 5,667 km respectively). Important stopover sites were discovered in Africa and Israel. Wintering areas were found from Mauritania and Sierra Leone in the west to Ethiopia and Central African Republic in the east and south. One of the storks migrating by the eastern migration route surprisingly reached western Africa. Birds that arrived early in the wintering areas stayed longer than those arriving later. On the average, birds migrating via the western route spent 37 d on migration compared to 80 d for birds migrating via the eastern route. The mean migration speed in the autumn was 126 km/d and the fastest stork flew 488 km/d when crossing the Sahara. The repeatedly tracked storks showed high winter site fidelity.     

Foraging behavior of three species of songbirds during stopover in southeastern Morocco during spring migration

Investigators studying the stopover ecology of migrating birds typically use the capture–recapture method to examine important parameters such as fuel deposition rates (FDR) and stopover duration. However, such studies can be constrained by the number of recaptures. An alternative method is to calculate a regression of mass over time of day, but this method may not be reliable because patterns of mass change of individual birds through the day may not reflect that of the whole population. Given the potential constraints of these methods, using them in combination with other methods, such as behavioral observations of foraging birds, may improve our understanding of the patterns of fuelling in birds at stopover sites. We observed the foraging behavior of three songbird species, including Western Bonelli's (Phylloscopus bonelli), Subalpine (Sylvia cantillans), and Willow (Phylloscopus trochilus) warblers, from 15 March to 30 April 2011 at a small oasis at the northern border of the Sahara desert in southeast Morocco. Given the location of our study site at the northern edge of the Sahara desert, birds migrating north likely needed to replenish their energy reserves at this stage of their journey. We assessed foraging effort by determining the rate (number per unit time) at which birds pecked at substrates or made aerial forays after flying insects. Peck rates were higher for Western Bonelli's Warblers than for Subalpine and Willow warblers, suggesting either species‐specific adaptations to feeding in arid environments or differences in the motivation to feed. In addition, Western Bonelli's Warblers had FDRs that were negative or close to zero and, therefore, were apparently unable to refuel successfully (i.e., increase their fuel stores) despite greater effort, possibly indicating less efficiency in obtaining food (i.e., more unsuccessful pecks). The lower peck rates of Subalpine and Willow warblers suggest either that they were less efficient at finding prey or were simply foraging at lower rates. For all three species, peck rates were lower at higher wind speeds, suggesting that wind may alter prey availability and detectability, especially of flying insects. Interactions among species‐specific migration strategies, environmental conditions, and habitat quality ultimately define the success of migration. Our results suggest that using observational data in combination with capture data may improve our understanding of these interactions at migration stopover sites.     

Genetically‐based behavioural morph affects stopover refuelling performance in white‐throated sparrows Zonotrichia albicollis

Intraspecific competition can influence refuelling at migration stopover sites. White‐throated sparrows Zonotrichia albicollis have genetically‐determined plumage morphs that differ in dominance behaviour and competitive abilities. This study examines the effects of plumage morph, sex and age, three likely indicators of competitive ability, on fall migration timing, body composition, and refuelling rates during stopover at Long Point, Ontario. We used quantitative magnetic resonance analysis and plasma metabolite profiling to determine body composition and refuelling rates, respectively. We determined sex and plumage morph genetically. Competitive ability did not influence migration timing. Controlling for structural size, males had larger lean mass than females, but we found no differences in body fat or lean mass between plumage morphs. Plasma metabolite concentrations indicated that the aggressive white‐stripe morph refuelled faster than the less aggressive tan‐stripe morph, though there were no differences among sex and age groups. We suggest that increased refuelling rates did not result in increased fat or lean mass because 1) individuals categorised as less competitive had longer stopover durations to compensate for slower refuelling rates; or 2) costs of behaving more competitively offset gains from faster refuelling rates.     

Seasonal Survival Probabilities Suggest Low Migration Mortality in Migrating Bats

Migration is adaptive if survival benefits are larger than costs of residency. Many aspects of bat migration ecology such as migratory costs, stopover site use and fidelity are largely unknown. Since many migrating bats are endangered, such information is urgently needed to promote conservation. We selected the migrating Leisler''s bat (Nyctalus leisleri) as model species and collected capture-recapture data in southern Switzerland year round during 6 years. We estimated seasonal survival and site fidelity with Cormack-Jolly-Seber models that accounted for the presence of transients fitted with Bayesian methods and assessed differences between sexes and seasons. Activity peaked in autumn and spring, whereas very few individuals were caught during summer. We hypothesize that the study site is a migratory stopover site used during fall and spring migration for most individuals, but there is also evidence for wintering. Additionally, we found strong clues for mating during fall. Summer survival that included two major migratory journeys was identical to winter survival in males and slightly higher in females, suggesting that the migratory journeys did not bear significant costs in terms of survival. Transience probability was in both seasons higher in males than in females. Our results suggest that, similarly to birds, Leisler''s bat also use stopover sites during migration with high site fidelity. In contrast to most birds, the stopover site was also used for mating and migratory costs in terms of survival seemed to be low. Transients'' analyses highlighted strong individual variation in site use which makes particularly challenging the study and modelling of their populations as well as their conservation.     

Expensive Traditions: Energy Expenditure of Aleutian Geese in Traditional and Recently Colonized Habitats

ABSTRACT In an effort to reduce goose depredation at a traditional spring migratory stopover site, private landowners implemented a coordinated hazing plan to scare Aleutian cackling geese (Branta hutchinsii leucopareia) from private lands to adjacent public pastures that were cultivated and set aside specifically for geese. Coincidentally, some Aleutian geese began using a new stopover site 150 km farther south in their spring migratory range; numbers at the new site continue to increase. We tested the idea that when their ability to acquire resources deteriorates geese are likely to seek improved foraging conditions, especially during spring migration when individuals strive to maximize nutrient stores and minimize energy expenditure. We quantified measures of goose foraging performance in traditional and new spring staging sites by calculating foraging opportunity, foraging effort, body condition, and daily energy expenditure. Geese staging at the site with higher levels of human disturbance had less foraging opportunity and, despite increased foraging effort and more nutritious food-plants at the site, birds there experienced an elevated energy expenditure and poorer body condition than birds at the new stopover site. Reduced foraging time and increased energy expenditure at the traditional spring staging site may have triggered the colonization process. Suitability assessment of habitat for migratory geese should include measures of foraging opportunity, disturbance risks, and daily energy expenditure in addition to quantity and quality of foods.     

Stopover strategies in passerine bird migration: a simulation study

Long-distance bird migration consists of a series of stopovers (for refuelling) and flights, with flights taking little time compared to stopovers. Therefore, it has been hypothesized that birds minimize the total time taken for migration through efficient stopover behaviour. Current optimality models for stopover include (1) the fixed expectation rule and (2) the global update rule. These rules maximize the speed of migration by determining the optimal departure fuel load for a given fuel deposition rate. We were interested in simple behavioural rules approaching the stopover behaviour of real birds and how these rules compare to the time minimizing models above with respect to the total time taken for migration. The simple strategies were to stay at a site (1) until a fixed fuel load was reached or (2) for a constant number of days. We simulated migration of small nocturnal passerine birds across an environment of continuously distributed but variable fuel deposition rates, and investigated the influence of different stopover strategies on the duration of migration. Staying for a constant number of days at each stopover site, irrespective of the fuel deposition rate, resulted in only slightly longer than minimum values for migration duration. Additionally, the constant stopover duration, e.g. 10 days, may change by a day or two (per stopover) without having a large effect on total migration duration. There is therefore a possibility that real birds may be close to optimal migration speed without the need for very complex behaviour. When assessing the sensitivity of migration duration to factors other than stopover duration, we found that flight costs, search and settling time, mean fuel deposition rate and the accuracy in the choice of flight direction were the factors with the largest influence. Our results suggest that migrating birds can approximate optimal stopover duration relatively easy with a simple rule, and that other factors, e.g. those above, are more relevant for travel time.     

Skipping‐type migration in a small Arctic wader,the Temminck's stint Calidris temminckii

By using morphometric data and geolocator tracking we investigated fuel loads and spatio‐temporal patterns of migration and non‐breeding in Temminck's stints Calidris temminckii. Body masses in stints captured at autumn stopover sites from Scandinavia to northern Africa were generally not much higher than during breeding and did not vary geographically. Thus, we expected migrating stints to make several stopovers and either circumventing the Sahara desert with low fuel loads or fuelling at north African stopover sites before desert crossing. Geolocation revealed that birds (n = 6) departed their Norwegian breeding site in the last part of July and all but one migrated south‐west over continental western Europe. A single bird headed south‐east to the Balkan Peninsula where the geolocator died. As predicted, southbound migration proceeded in a typical skipping manner with 1–4 relatively short stopovers (median 4 d) during 10–27 d of migration before reaching north‐west Africa. Here birds spent 11–20 d before crossing the Sahara. The non‐breeding sites were located at or near the Niger River in Mali and were occupied continuously for more than 215 d with no indications of itinerancy. Spring migration commenced in late April/early May when birds crossed the desert and used stopover sites in the western Mediterranean basin in a similar manner as during autumn. The lowest body masses were recorded in spring at islands in the central Mediterranean basin, indicating that crossing the Sahara and Mediterranean barriers is exhausting to these birds. Hence, the skipping‐type pattern of migration revealed by geolocators is likely to be natural in this species and not an effect of instrumentation.     

Migrating songbirds on stopover prepare for,and recover from,oxidative challenges posed by long‐distance flight

Managing oxidative stress is an important physiological function for all aerobic organisms, particularly during periods of prolonged high metabolic activity, such as long‐distance migration across ecological barriers. However, no previous study has investigated the oxidative status of birds at different stages of migration and whether that oxidative status depends on the condition of the birds. In this study, we compared (1) energy stores and circulating oxidative status measures in (a) two species of Neotropical migrants with differing migration strategies that were sampled at an autumn stopover site before an ecological barrier; and (b) a species of trans‐Saharan migrant sampled at a spring stopover site after crossing an ecological barrier; and (2) circulating oxidative measures and indicators of fat metabolism in a trans‐Saharan migrant after stopovers of varying duration (0–8 nights), based on recapture records. We found fat stores to be positively correlated with circulating antioxidant capacity in Blackpoll Warblers and Red‐eyed Vireos preparing for fall migration on Block Island, USA, but uncorrelated in Garden Warblers on the island of Ponza, Italy, after a spring crossing of the Sahara Desert and Mediterranean Sea. In all circumstances, fat stores were positively correlated with circulating lipid oxidation levels. Among Garden Warblers on the island of Ponza, fat anabolism increased with stopover duration while oxidative damage levels decreased. Our study provides evidence that birds build antioxidant capacity as they build fat stores at stopover sites before long flights, but does not support the idea that antioxidant stores remain elevated in birds with high fuel levels after an ecological barrier. Our results further suggest that lipid oxidation may be an inescapable hazard of using fats as the primary fuel for flight. Yet, we also show that birds on stopover are capable of recovering from the oxidative damage they have accrued during migration, as lipid oxidation levels decrease with time on stopover. Thus, the physiological strategy of migrating songbirds may be to build prophylactic antioxidant capacity in concert with fuel stores at stopover sites before a long‐distance flight, and then repair oxidative damage while refueling at stopover sites after long‐distance flight.     

Connecting the dots: Stopover strategies of an intercontinental migratory songbird in the context of the annual cycle

The phases of the annual cycle for migratory species are inextricably linked. Yet, less than five percent of ecological studies examine seasonal interactions. In this study, we utilized stable hydrogen isotopes to geographically link individual black‐and‐white warblers (Mniotilta varia) captured during spring migration with breeding destinations to understand a migrant's stopover strategy in the context of other phases of the annual cycle. We found that stopover strategy is not only a function of a bird's current energetic state, but also the distance remaining to breeding destination and a bird's time‐schedule, which has previously been linked to habitat conditions experienced in the preceding phase of the annual cycle. Birds in close proximity to their breeding destination accumulate additional energy reserves prior to arrival on the breeding grounds, as reflected by higher migratory condition upon arrival, higher refueling rates measured via blood plasma metabolites, and longer stopover durations compared to birds migrating to breeding destinations farther from the stopover site. However, late birds near their breeding destination were more likely to depart on the day of arrival (i.e., transients), and among birds that stopped over at the site, the average duration of stopover was almost half the time of early conspecifics, suggesting late birds are trying to catch‐up with the overall time‐schedule of migration for optimal arrival time on the breeding grounds. In contrast, birds with long distances remaining to breeding destinations were more likely to depart on the day of arrival and primarily used stopover to rest before quickly resuming migration, adopting similar strategies regardless of a bird's time‐schedule. Our study demonstrates that migrants adjust their en route strategies in relation to their time‐schedule and distance remaining to their breeding destination, highlighting that strategies of migration should be examined in the context of other phases of the annual cycle.