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.     

Stopover decision during migration: physiological conditions predict nocturnal restlessness in wild passerines

During migration, a number of bird species rely on stopover sites for resting and feeding before and after crossing ecological barriers such as deserts or seas. The duration of a stopover depends on the combined effects of environmental factors, endogenous programmes and physiological conditions. Previous studies indicated that lean birds prolong their refuelling stopover compared with fat birds; however, the quantitative relationship between physiological conditions and stopover behaviour has not been studied yet. Here, we tested in a large sample of free-living birds of three European passerines (whinchats, Saxicola rubetra, garden warblers, Sylvia borin and whitethroats, Sylvia communis) whether the amount of migratory restlessness (Zugunruhe) shown at a stopover site depends on physiological conditions. An integrated measure of condition based on body mass, amount of subcutaneous fat and thickness of pectoral muscles strongly predicted the intensity of Zugunruhe shown in recording cages in the night following capture. These results provide novel and robust quantitative evidence in support of the hypothesis that the amount of energy reserves plays a major role in determining the stopover duration in migratory birds.     

Fat reserves of migrating passerines at arrival on the breeding grounds in Swedish Lapland

Mist-net capture data taken during 6 years (1988–1990 and 1992–1994) of field work were used to describe the arrival sequences and fat loads of nine species of migratory passerines which breed in a near-Arctic environment in Swedish Lapland. Long-distance migrants arrived with significantly larger mean fat reserves than did medium- and short-distance migrants. Long-distance migrants carried fat loads at arrival which corresponded to potential remaining flight distances between 242 and 500 km. When females and males arrived on the breeding grounds simultaneously, females carried significantly larger energy reserves than did males in seven out of nine species. In contrast, when the sexes showed a significant difference in median arrival date (two out of nine species), there was no difference in mean fat load carried into the breeding area. A relationship was found between the migratory habits and foraging ecology of each species and the amount of fat reserves at arrival, suggesting that species-specific migratory distances and feeding habits determine the amount of fat that is needed during the transition period between migration and onset of breeding. The short growing season in the study area restricts the time available for breeding and moult, and large energy reserves at arrival may speed up the breeding schedule to counteract possible time constraints. Overloading at the last stopover site during spring migration may be an adaptation allowing birds to cope with a restricted time frame for breeding and moult at high latitudes.     

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.     

Migratory stopover conditions affect the developmental state of male gonads in garden warblers (Sylvia borin)

Long-distance migrants face the challenge of a short window for reproduction that requires optimal timing and full functional gonads. Male garden warblers (Sylvia borin) meet these demands by initiating testicular recrudescence during spring migration, enabling them to reproduce immediately after arrival at the breeding grounds. In a combined field and laboratory study, we investigated testicular size, plasma levels of luteinizing hormone (LH), androstenedione (AE), 5α-dihydrotestosterone (DHT), testosterone and nocturnal migratory restlessness (Zugunruhe) under different stopover conditions. We manipulated food availability, the duration of stopover and simulated migration by food deprivation. Garden warblers showed significantly retarded testicular development after nine days of stopover under limited food conditions compared to birds that had ad libitum access to food. However, there was no significant difference in Zugunruhe between the two groups. Thus, the degree of Zugunruhe was unaffected by the quality of the stopover site and migration continued independent of the developmental state of the testis. We suggest that male garden warblers face the necessity to either compensate for slowed testicular recrudescence during the subsequent leg of migration and delay arrival at the breeding grounds, or arrive with less developed testes. Either of these may reduce annual reproductive success.     

During stopover, migrating blackcaps adjust behavior and intake of food depending on the content of protein in their diets

During migration, birds undergo alternating periods of fasting and re-feeding that are associated with dynamic changes in body mass (m(b)) and in organ size, including that of the digestive tract. After arrival at a migratory stopover site, following a long flight, a bird must restore the tissues of its digestive tract before it can refuel. In the present study we examined how the availability of dietary protein influences refueling of migrating blackcaps (Sylvia atricapilla) during a migratory stopover. We tested the following predictions in blackcaps deprived of food and water for 1-2 days to induce stopover behavior: (1) birds provided with a low-protein diet will gain m(b), lean mass and fat mass, and increase in pectoral muscle size slower than do birds fed a high-protein diet; (2) since stopover time is shorter in spring, birds will gain m(b) and build up fat tissue and lean tissue faster than in autumn; and (3) if low dietary protein limits a bird's ability to gain m(b) and fat reserves, then birds that do not obtain enough protein will initiate migratory restlessness (Zugunruhe) earlier than will birds with adequate dietary protein. These predictions were tested by providing captured migrating blackcaps with semisynthetic isocaloric diets differing only in their protein content. Each day, we measured m(b), and food intake; also lean mass and fat mass were measured using dual energy X-ray absorptiometry. In addition, we monitored nocturnal activity with a video recording system. In both spring and autumn, birds fed diets containing either 3 or 20% protein increased in m(b), lean mass and fat mass at similar rates during the experiment. However, the group receiving 3% protein ate more than did the group receiving 20% protein. In support of our predictions, m(b), lean mass, fat mass, and intake of food all were higher in spring than in autumn. We also found that in spring all birds had higher levels of migratory restlessness, but birds fed 3% protein were less active at night than were birds fed 20% protein, possibly an adaptation conserving energy and protein. We conclude that protein requirements of migrating blackcaps during stopover are lower than expected, and that birds can compensate for low dietary protein by behavioral responses, i.e. hyperphagia and decreased migratory restlessness, that ensure rapid refueling.     

Faster migration in autumn than in spring: seasonal migration patterns and non‐breeding distribution of Icelandic whimbrels Numenius phaeopus islandicus

Migration is fundamental in the life of many birds and entails significant energetic and time investments. Given the importance of arrival time in the breeding area and the relatively short period available to reproduce (particularly at high latitudes), it is expected that birds reduce spring migration duration to a greater extent than autumn migration, assuming that pressure to arrive into the wintering area might be relaxed. This has previously been shown for several avian groups, but recent evidence from four tracked Icelandic whimbrels Numenius phaeopus islandicus, a long distance migratory wader, suggests that this subspecies tends to migrate faster in autumn than in spring. Here, we 1) investigate differences in seasonal migration duration, migration speed and ground speed of whimbrels using 56 migrations from 19 individuals tracked with geolocators and 2) map the migration routes, wintering and stopover areas for this population. Tracking methods only provide temporal information on the migration period between departure and arrival. However, migration starts with the fuelling that takes place ahead of departure. Here we estimate the period of first fuelling using published fuel deposition rates and thus explore migration speed using tracking data. We found that migration duration was shorter in autumn than in spring. Migration speed was higher in autumn, with all individuals undertaking a direct flight to the wintering areas, while in spring most made a stopover. Wind patterns could drive whimbrels to stop in spring, but be more favourable during autumn migration and allow a direct flight. Additionally, the stopover might allow the appraisal of weather conditions closer to the breeding areas and/or improve body condition in order to arrive at the breeding sites with reserves.     

迁徙鸟类中途停歇期的生理生态学研究

大多数候鸟的迁徙活动由迁徙飞行和中途停歇两个部分组成。在迁徙过程中,鸟类要多次交替经历消耗能量的飞行阶段和积累能量的中途停歇阶段。从鸟类在中途停歇时期的能量积累速度、体重变化模式以及迁徙飞行中的禁食或食物限制、食物种类的改变、中途停歇的能量快速积累过程对消化器官的影响等方面,对目前迁徙鸟类的生理生态学研究成果进行回顾,并提出有待解决的问题及今后的研究方向。     

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.     

Can conditions experienced during migration limit the population levels of birds?

Populations of migratory birds are usually considered to be limited by conditions in breeding or wintering areas, but some might be limited by conditions encountered on migration. This could occur at stopover sites where competition for restricted food supplies can reduce subsequent survival or breeding success, or during the flights themselves, when adverse weather can occasionally kill large numbers of individuals. Competition for food could act in a density-dependent manner and help to regulate populations, whereas weather effects are more likely to act in a density-independent manner. The evidence for these views is explored in this paper. When preparing for migration, birds must normally obtain more food per day than usual, in order to accumulate the body reserves that fuel their flights. Birds often concentrate in large numbers at particular stopover sites, where food can become scarce, thus affecting migratory performance. Rates of weight gain, departure weights, and stopover durations often correlate with food supplies at stopover sites, sometimes influencing the subsequent survival and reproductive success of individuals, which can in turn affect subsequent breeding numbers. Many studies have provided evidence for interference and depletion competition at stopover sites, relatively few for migration conditions influencing the subsequent breeding or survival of individuals, and even fewer for effects on subsequent breeding numbers. Migrants in flight occasionally suffer substantial mortality in storms, especially over water, sometimes involving many thousands of birds at a time. Other mass mortalities have resulted from atypical ‘winter-like’ weather, occurring soon after the arrival of summer migrants in their breeding areas or just before their departure in autumn. Again, many thousands of birds at a time have been killed in such incidents, causing reductions of 30–90% in local breeding densities. In some bird species, migration-related events can at times have substantial effects on the year-to-year changes in breeding population levels. Nonetheless, the difficulties involved in investigating migrating birds at different points on their migration routes have so far limited the number of studies on the influence of events during migration periods on population levels.     

Simulation of migratory flight and stopover affects night levels of melatonin in a nocturnal migrant

Several species of diurnal birds are nocturnal migrants. The activation of nocturnal activity requires major physiological changes, which are essentially unknown. Previous work has shown that during migratory periods nocturnal migrants have reduced night-time levels of melatonin. Since this hormone is involved in the modulation of day-night rhythms, it is a good candidate regulator of nocturnal migratory activity. We studied whether melatonin levels change when nocturnally active blackcaps (Sylvia atricapilla) are experimentally transferred from a migratory to a non-migratory state. We simulated a long migratory flight by depriving birds of food for 2 days, and a refuelling stopover by subsequently re-administering food. Such a regimen is known to induce a reduction in migratory restlessness ('Zugunruhe') in the night following food reintroduction. The experiments were performed in both autumn and spring using blackcaps taken from their breeding grounds (Sweden) and their wintering areas (Kenya). In autumn, the food regimen induced a suppression of Zugunruhe and an increase in melatonin in the night following food reintroduction. In spring, the effects of the treatment were qualitatively similar but their extent depended on the amount of body-fat reserves. This work shows that the reduction of night-time melatonin during migratory periods is functionally related to nocturnal migration, and that fat reserves influence the response of the migratory programme to food deprivation.     

Seasonal migration strategies of Common Ringed Plovers Charadrius hiaticula

How individual birds schedule their movements and use different sites during the non‐breeding season are fundamental issues in avian migration ecology, and studies have often revealed strong seasonal variation in such strategies. Using geolocators we tracked Common Ringed Plovers Charadrius hiaticula from northern Norway to West Africa and back to assess whether there were differences in migratory speed, duration and stopover use between autumn and spring migration and whether birds used multiple sites during the non‐breeding season. Although the pace of migration was similar between autumn and spring, the length of flight bouts and duration of the preceding stopovers were positively correlated only in autumn. Four of five birds showed a marked southward movement in mid‐winter.     

Food availability but not melatonin affects nocturnal restlessness in a wild migrating passerine

A large number of passerine species migrate at night, although most of them are diurnal outside the migratory seasons. This diurnal-to-nocturnal transition is a major life-history event, yet little is known about its physiological control. Previous work showed that during the migratory periods captive birds showing nocturnal migratory restlessness (Zugunruhe) have reduced concentrations of circulating melatonin at night compared to non-migratory periods. This suggested that the hormone melatonin, a main component of the avian circadian system, is involved in the expression of Zugunruhe. Other studies demonstrated that the relationship between low melatonin levels and Zugunruhe is not a seasonal correlation. When Zugunruhe was interrupted by exposing birds to a fasting-and-refeeding protocol, melatonin levels increased. Here we studied whether melatonin and food availability influence the intensity of Zugunruhe in wild migrating garden warblers (Sylvia borin) at a stopover site. Birds were held in recording cages overnight, with or without food available, and either bled to determine melatonin concentrations or treated transdermally with melatonin. We found that melatonin levels at night were correlated with the intensity of diurnal locomotor activity and with condition, but were not correlated with Zugunruhe. Similarly, the melatonin treatment did not have effects on Zugunruhe, whereas food availability increased it. Our study shows that the nocturnal melatonin levels in migrating warblers depend on food availability and are correlated with condition. In addition, it suggests that melatonin does not control Zugunruhe and might rather be involved in energy conservation and/or clock synchronization during migration.     

Stopover duration of Palearctic passerine migrants in the western Sahara – independent of fat stores?

Birds on migration spend much more time on stopover sites to refuel for the next migration step than aloft, but empirical data on stopover duration are rare, especially for Palearctic trans-Sahara migrants whilst crossing the desert. Previous studies suggest that stopover duration of fat birds in oases is much shorter than that of lean birds. During 2003 and 2004 capture–recapture data of migrating passerines from two inland oases in spring and from one coastal site in autumn in Mauritania, West Africa, were analysed to test whether the probability of being a transient and the stopover duration depend on fuel stores at first capture. The application of capture–recapture models revealed that during autumn migration at the coast the proportion of transients (individuals that stop over only for 1 day) was relatively high (77–90%) in three out of four species investigated and stopover duration was short (1.9–4.6 days). In the inland oases in spring, transients were detected in only four out of 12 analyses. Stopover duration was longer than at the coast in autumn and surprisingly long in some species with durations of up to 30 days. Models taking into account the initial fat load of birds on the first capture occasion were, with one exception, never the most parsimonious ones. This indicates that the time spent after and before capture at the stopover site did not depend on the fat stores at first capture. Therefore, we cannot confirm the assumption that birds arriving at stopover sites in the desert with low fat loads stay longer than birds that arrive with high fat loads.     

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.     

Light‐level geolocation reveals wintering distribution,migration routes,and primary stopover locations of an endangered long‐distance migratory songbird

The importance of understanding the geographic distribution of the full annual cycle of migratory birds has been increasingly highlighted over the past several decades. However, the difficulty of tracking small birds between breeding and wintering areas has hindered progress in this area. To learn more about Kirtland's warbler Setophaga kirtlandii movement patterns throughout the annual cycle, we deployed archival light‐level geolocators across their breeding range in Michigan. We recovered devices from 27 males and analyzed light‐level data within a Bayesian framework. We found that most males wintered in the central Bahamas and exhibited a loop migration pattern. In both fall and spring, departure date was the strongest predictor of arrival date, but in spring, stopover duration and migration distance were also important. Though stopover strategies varied, males spent the majority of their spring migration at stopover sites, several of which were located just before or after large ecological barriers. We argue that loop migration is likely a response to seasonal variation in prevailing winds. By documenting a tight link between spring departure and arrival dates, we provide a plausible mechanism for previously documented carry‐over effects of winter rainfall on reproductive success in this species. The migratory periods remain the least understood periods for all birds, but by describing Kirtland's warbler migration routes and timing, and identifying locations of stopover sites, we have begun the process of better understanding the dynamics of their full annual cycle. Moreover, we have provided managers with valuable information on which to base future conservation and research priorities.     

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.     

Body composition of north and southbound migratory blackcaps is influenced by the lay‐of‐the‐land ahead

The annual migration of small birds depends on the optimal management of time and energy. Since refueling at stopovers between flights consumes most of the birds’ time and energy, selection of food‐rich sites, and timely departure therefrom are likely crucial to success. We examined this concept quantifying body composition of 200 migrating blackcaps, Sylvia atricapilla, in Eilat, Israel, using dual‐energy x‐ray absorptiometry and generated a model to predict body composition as it changes with body mass (m_b). We then back‐calculated body composition of > 20 000 blackcaps ringed between 1984 and 2005, and tested the hypothesis that the amount of fuel that a bird stores determines the length of its stopover. We predicted that 1) if time‐constrained in spring, birds at the stopover site carry less than a maximum fuel load, but 2) if not time‐constrained, as in autumn, their fuel load is much higher than in spring. We found the change in body composition of blackcaps to be biphasic and correlated with increasing m_b. At m_b < ? 17.8 g, increasing m_b is due to increasing lean mass (m_l), while at m_b > ? 17.8 g increasing m_b results from increasing fat mass (m_f), which is accompanied by decreasing m_l. Body composition of blackcaps at a spring stopover site indicates that blackcaps leave stopovers as soon as they regain functionality of their digestive systems, but before laying down much m_f. In autumn blackcaps arrive with fuel stores much larger than in spring. For these birds, the Eilat stopover apparently serves to complete fat accumulation before crossing the deserts ahead. We conclude that in spring, the decision to depart is not determined by the bird's fuel stores, especially when early arrival at the breeding site, and therefore time, is of the essence. In autumn, accumulating enough fuel to ensure successful crossing of the deserts ahead probably dictates stopover time.     

Simultaneous tetragyny in Northern Lapwing Vanellus vanellus

Capsule Fuel load is correlated with fuel deposition rate; stopover duration is affected by arrival fuel load.

Aims To determine the stopover duration, fuel management and flight ranges at departure of Blackcaps stopping over in northern Spain.

Methods Systematic mist-netting and ringing allowed the use of mark–recapture Cormack–Jolly–Seber models for the estimation of stopover duration. Trapped birds were measured and weighed in order to estimate mass gain. FLIGHT software was used to estimate flight ranges.

Results Stopover duration ranged from 3.6 to 13.6 days, and was negatively correlated with arrival body mass (assessed by body mass at the first capture event). On average, arrival body mass was 18.4 g, whilst body mass at departure was 19.8 g. No significant differences in arrival body mass and departure body mass were observed between age or sex classes. Mass deposition rate did not differ between age or sex classes (mean = 0.20 g/day). Birds recaptured one day after the first capture event lost mass, whilst recaptures from the second day onwards had a mean gain of mass; mass was observed to increase linearly with the stopover duration. Mass deposition rate was positively correlated with departure body mass. Finally, with a mean departure body mass of 19.8 g, a Blackcap stopping over in northern Spain should be able to fly up to 1100 km.

Conclusions Stopover duration assessed by Cormack–Jolly–Seber models was longer than that observed in birds recaptured more than once (‘minimum stopover duration’). Stopover was longer for birds arriving with less fuel. The positive relationship between departure body mass and mass deposition rate suggests a time-minimizing strategy. The lack of difference in fuel deposition rate between age and sex classes suggests a relatively abundant food supply at the study site, but other explanations might also account for the lack of age and sex differences, for example if competition for food was not determined by social hierarchies but by scramble competition. Departing fuel load would allow these birds to arrive at their wintering areas in southern Spain under still-air conditions, without needing to refuel.     

长距离迁徙鸟类对应于能量积累状态的取食行为调整

迁徙鸟类能够预计到迁徙过程中对能量需求的增加和迁徙途中获得能量的不确定性。最佳迁徙理论指出：迁徙停留期的一系列决策受到体内能量状态、取食机会和迁徙时间的影响。利用刚完成春季跨越墨西哥湾迁徙的鸫类,我们研究了取食行为、体内能量状态和能量积累速度的相互关系。我们用雾网捕获了停留的鸫,然后给每只鸟进行了环志和称重,并估测了表皮下积累的脂肪。为了检测体内能量积累大小对取食行为的影响,我们把在野外观察到的鸟分成肥、瘦两组。当一天里被捕获鸟的平均体重低于相关种的瘦体重时,这一天被观察到的取食鸟被归到瘦组,反之就属于肥组。我们同时对一部分鸟用有色环进行了标记,以便能在野外观察到它们时能准确地知道每一只鸟的初始能量积累状态。在春天完成跨越墨西哥湾迁徙以后的鸫大约有50％在到达停留地时就已经消耗了所有的表皮下脂肪。与肥组鸟相比,瘦组的个体在停留期扩展了取食方法和取食基底,增加了取食速度。与此相关的是,瘦鸟表现出体重积累更多、速度更快。我们的数据表明迁徙鸟在到达迁徙中途停留地时的能量状态会影响到它们停留期间的取食行为和继续迁徙时的生理机能,从而影响停留期的长短和在迁徙途中停留与否的决定[动物学报51(1)：12—23,2005]。