Size and accumulation of fuel reserves at stopover predict nocturnal restlessness in a migratory bird

Early arrival at the breeding site positively affects the breeding success of migratory birds. During migration, birds spend most of their time at stopovers. Therefore, determining which factors shape stopover duration is essential to our understanding of avian migration. Because the main purpose of stopover is to accumulate fat as fuel for the next flight bout, fuel reserves at arrival and the accumulation of fuel are both expected to affect stopover departure decisions. Here, we determined whether fuel reserves and fuel accumulation predict a bird''s motivation to depart, as quantified by nocturnal migratory restlessness (Zugunruhe), using northern wheatears (Oenanthe oenanthe) that were captured and temporarily contained at spring stopover. We found that fuel reserves at capture were positively correlated with Zugunruhe, and negatively correlated with fuel accumulation. This indicates that fat birds were motivated to depart, whereas lean birds were set on staying and accumulating fuel. Moreover, the change in fuel reserves was positively correlated with the concurrent change in Zugunruhe, providing the first empirical evidence for a direct link between fuel accumulation and Zugunruhe during stopover. Our study indicates that, together with innate rhythms and weather, the size and accumulation of fuel reserves shape stopover duration, and hence overall migration time.     

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.     

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.     

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.     

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.     

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.     

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.     

A separate circadian oscillator controls nocturnal migratory restlessness in the songbird Sylvia borin

When confined to a cage, migratory songbirds exhibit nocturnal migratory restlessness (also called Zugunruhe) during the spring and autumn migratory periods, even though these birds are exclusively diurnal during the remainder of the year. Zugunruhe, which has been demonstrated to be under the direct control of a circannual timer, is characterized by a stereotypic "wing-whirring" behavior while the bird is perched. To elucidate the role played by the circadian system in the regulation of Zugunruhe, the authors studied the activity of garden warblers (Sylvia borin), long-distance nocturnal migrants, under skeleton photoperiods of different lengths and under constant dim light. In 11.5D:1L:10.5D:1L skeleton photoperiods, the authors found that Zugunruhe free-ran in a substantial proportion of birds, while their normal daily activities (e.g., feeding and preening) remained synchronized to 24 h. Some birds expressing Zugunruhe under constant dim light continued to show 2 distinct bouts of activity: one corresponding to daily activities, the other to wing-whirring. In some cases, these 2 bouts crossed while free-running with different periods. Birds expressing Zugunruhe also had significantly longer free-running periods than birds that did not. The study data suggest that the seasonal appearance of Zugunruhe is the result of the interactions of at least 2 circadian oscillators and that it is the phase relationship of these 2 oscillators that determines when nocturnal migratory restlessness is expressed. Furthermore, these data are consistent with the previously proposed internal coincidence hypothesis as a model for the ontogeny of circannual rhythms.     

Stopover ecology of Red-flanked Bush Robin (Tarsiger cyanurus) at Maoershan, Northeast China

The conservation of migratory songbirds in China faces challenges similar to those in other parts of the world such as Europe and North America. Migratory songbirds have a complex life history and are associated with large spatial scale. Events that occur during migration play a vital role in determining population status. Little is known about the stopover behavior of migratory songbirds in China during their passage, including stopover duration, energetic condition (e.g., the amount of fat stores), and the habitat of these birds. We investigated migration timing, stopover duration, change of energetic condition and sex-related variations among Red-flanked Bush Robins (Tarsiger cyanurus) at a stopover site in Northeast China. A total of 1 751 Red-flanked Bush Robins were captured in the fall of 2002, with further 684 captured in the spring of 2003. The body mass of the fall birds was higher than that of the spring ones. The condition index (i.e., body mass adjusted for body size) was lowest among spring females. Birds were more likely to stay longer in fall than in spring. We detected a positive relationship between the time of initial capture and condition index regardless of season or sex, suggesting that the birds were able to replenish energy stores. The net daily mass gain was the highest among males in the fall (3% of body mass). The net daily mass gain would sustain a flight of 0.6 hours for females and 3.1 hours for males in the fall. The stopover biology of Red-flanked Bush Robins is similar to that of some migratory songbirds in Europe and North America. For example, the spring passage time and fat store variation between the sexes concurs with the hypothesis that males are selected to arrive at their breeding grounds as early as food resources or climatic conditions are adequate in the spring. Further research on stopover ecology is urgently required in China to achieve a better understanding of migratory behavior and ensure the conservation of these migratory songbirds.     

Accounting for heterogeneity when estimating stopover duration,timing and population size of red knots along the Luannan Coast of Bohai Bay,China

相似文献   

Nocturnal Activity of Caged Resident and Migrant Silvereyes (Zosteropidae: Aves)

Night-activity patterns of individually-caged grey-breasted silvereyes Zosterops lateralis of resident and migrant status were investigated. True migratory restlessness or ‘Zugunruhe’ was exhibited by some individuals, but there was also considerable ‘noise’ in the data caused by cageing and neighbouring disturbance. In general, ‘Zugunruhe’ took place from about 2 h before, to about 4 h after ‘sunrise’, and both the daily timing and annual duration of Zugunruhe were consistent with the migration pattern observed in the field. A total of 42 % of resident birds from Tasmania, 77 % of migrant birds from Tasmania and 90 % of migrant birds from the mainland were considered to have exhibited true ‘Zugunruhe’, while none of the resident birds from Brisbane were considered to have displayed it. A few individuals spontaneously developed ‘Zugunruhe’ in the absence of other birds, but many birds (both residents and migrants) displayed no elevated night unrest when placed in a room on their own, although displaying unrest when placed among birds exhibiting ‘Zugunruhe’. The migration of the silvereye probably has a genetic basis, but whether an individual migrates or not is also influenced by social factors which differ in magnitude between individuals.     

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.     

红胁蓝尾鸲(Tarsiger cyanurus)在中国东北部帽儿山地区的迁徙中途停歇生态

中国迁徙鸣禽类的保护面对着与世界其他地区如欧洲和北美洲鸟类保护相似的挑战。迁徙鸣禽类具有复杂生活周期和很大的空间关联。迁徙过程中发生的事件对迁徙鸣禽类种群动态具有决定作用。对于鸣禽类迁徙中途停歇期的生态,比如停歇期的长短,能量的积累,生境的利用等,了解还非常有限。在中国东北部的一个鸟类迁徙停歇地对红胁蓝尾鸲(Tarsiger cyanurus)的中途停歇生态包括迁徙时间、停歇时间、能量状态和性比进行了研究。2002年秋和2003年春分别捕获了1751只和684只红胁蓝尾鸲。红胁蓝尾鸲的体重在秋季迁徙时要比在春季迁徙时重。春季雌性红胁蓝尾鸲停歇时的能量状态指数最低; 而秋季的红胁蓝尾鸲比春季的红胁蓝尾鸲停歇时间更长。无论季节和性别,红胁蓝尾鸲的能量状态指数和第1次捕获的时间早晚成正相关, 间接证明红胁蓝尾鸲在停歇期间能够比较快地积累能量。秋季雄性红胁蓝尾鸲日体重净增率最大。估测秋季停歇期的每日能量净增能维持红胁蓝尾鸲雌性0.6h和雄性3.1h的飞行。红胁蓝尾鸲的中途停歇生态与北美和欧洲一些迁徙鸣禽类很相似。比如,春季迁徙过境的时间和脂肪积累的变化与自然选择对雄性的要求：当食物和气候适宜时尽快到达繁殖地的假设是一致的。对迁徙中途的停歇生态研究有利于更好地了解鸟类的迁徙行为和更有效地保护迁徙鸣禽类。     

红胁蓝尾鸲(Tarsiger cyanurus)在中国东北部帽儿山地区的迁徙中途停歇生态

中国迁徙鸣禽类的保护面对着与世界其他地区如欧洲和北美洲鸟类保护相似的挑战.迁徙鸣禽类具有复杂生活周期和很大的空间关联.迁徙过程中发生的事件对迁徙鸣禽类种群动态具有决定作用.对于鸣禽类迁徙中途停歇期的生态,比如停歇期的长短,能量的积累,生境的利用等,了解还非常有限.在中国东北部的一个鸟类迁徙停歇地对红胁蓝尾鸲（Tarsiger cyanurus）的中途停歇生态包括迁徙时间、停歇时间、能量状态和性比进行了研究.2002年秋和2003年春分别捕获了1751只和684只红胁蓝尾鸲.红胁蓝尾鸲的体重在秋季迁徙时要比在春季迁徙时重.春季雌性红胁蓝尾鸲停歇时的能量状态指数最低;而秋季的红胁蓝尾鸲比春季的红胁蓝尾鸲停歇时间更长.无论季节和性别,红胁蓝尾鸲的能量状态指数和第1次捕获的时间早晚成正相关,间接证明红胁蓝尾鸲在停歇期间能够比较快地积累能量.秋季雄性红胁蓝尾鸲日体重净增率最大.估测秋季停歇期的每日能量净增能维持红胁蓝尾鸲雌性0.6h和雄性3.1h的飞行.红胁蓝尾鸲的中途停歇生态与北美和欧洲一些迁徙鸣禽类很相似.比如,春季迁徙过境的时间和脂肪积累的变化与自然选择对雄性的要求：当食物和气候适宜时尽快到达繁殖地的假设是一致的.对迁徙中途的停歇生态研究有利于更好地了解鸟类的迁徙行为和更有效地保护迁徙鸣禽类.     

STOPOVER AND FAT DEPOSITION BY BLACKCAPS SYLVIA ATRICAPILLA FOLLOWING SPRING MIGRATION OVER THE SAHARA

Maitav, A. & Izhaki, I. 1994. Stopover and fat deposition by Blackcaps Sylvia atricapilla following spring migration over the Sahara. Ostrich 65:160-166.

Length of stopover and rate of weight change were studied in Blackcaps that stopover in Eilat (southern Israel) in spring after a trans-Saharan flight. The birds studied had already completed 2000 km of desert crossing and faced just 200 km more. The potential predicted flight range for these transients suggests that the fat depleted Blackcaps (fat class=0) could not reach the east Mediterranean scrublands, which offer more extensive foraging opportunities than the desert area around Eilat, without refueling. However, 46% of the Blackcaps had sufficient fat (fat class > 0) to reach central or northern Israel in a single flight in still air. Recaptured birds stayed in Eilat for 2,4±1,9 days. Although the body mass of an average recaptured bird increased by 0,27±1,44 g/day, which was an increase of 0,8%±7,0% of its body mass per day during stopover, these results were not significantly different from zero. Stopover length, mass change during stopover, and the rate of change in mass did not differ between males and females and between those who stopped over for only one day to those which stayed > 1 day. However, there was a tendency of these lean birds to stay for a shorter period in Eilat than relatively fat birds. There was also some tendency for lean birds to gain more mass during their stopover than fat birds. It seems that in spring, when less time is allotted for the whole migration program, the migrants resume their journey before fat reserves have been replenished.     

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.     

Aminopeptidase-N modulation assists lean mass anabolism during refuelling in the white-throated sparrow

Songbirds meet the extreme metabolic demands of migration by burning both stored fat and protein. However, catabolizing these endogenous tissues for energy leads to organ atrophy, and reductions in gastrointestinal tissue can be as great as 50% of the pre-flight mass. Remarkably, during stopover refuelling birds quickly regain digestive mass and performance. Aminopeptidase-N (APN) is a brush-border enzyme responsible for late-stage protein digestion and may critically assist tissue reconstruction during the stopover, thus compensating for reduced gut size. We hypothesized that birds recovering from a fast would differentially upregulate APN activity relative to disaccharidases to rapidly process and assimilate dietary protein into lean mass. We fasted 23 wild-caught migratory white-throated sparrows (Zonotrichia albicollis) for 48 h to mimic mass reductions experienced during migratory flight and measured intestinal APN activity before the fast, immediately after the fast, and during recovery at 24 h and 48 h post-fast. Total fat mass, lean mass and basal metabolic rate were measured daily. We show that fasted birds maintain APN activity through the fast, despite a 30% reduction in intestine mass, but during refuelling, APN activity increases nearly twofold over pre-fasted individuals. This suggests that dynamically regulating APN may be necessary for rapid protein reconstruction during the stopover.     

Migratory restlessness in caged BramblingsFringilla montifringilla in northern Sweden

Summary The migratory restlessness was studied in caged Bramblings in northern Sweden under natural light conditions 17 September 1977 to 25 May 1978. The experimental birds showed a well-developed migratory restlessness during the natural migratory season of the free-living conspecifics. The migratory restlessness lasted longer than the natural migratory period in autumn, and the onset of migratory restlessness in spring was delayed compared with the natural migratory period. The discrepancies are probably due to the abnormal photoperiodic conditions to which the experimental birds were exposed. Different diurnal activity patterns during different times of the year reflect the migratory habit of the species, i.e. the Brambling is both a nocturnal and diurnal migrant. The quantity of the diurnal activity per hour was clearly positively correlated with the length of photoperiod. The photoperiodic and endogenous influence on the migration of the Brambling is discussed.

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Zugunruhe bei gekäfigten Bergfinken in Nordschweden

Zusammenfassung Die Zugunruhe gekäfigter Bergfinken wurde vom 17. Sept. 1977 bis 15. Mai 1978 unter natürlichen Lichtbedingungen in Nordschweden untersucht. Während der Zugperiode freilebender Bergfinken zeigten die gekäfigten Individuen deutliche Zugunruhe. Im Herbst dauerte die Periode der Zugunruhe länger als die Zugperiode freilebender Bergfinken, im Frühjahr war das Einsetzen der Zugunruhe gegenüber dem natürlichen Zugbeginn verzögert. Möglicherweise waren diese Unterschiede eine Folge der abnormen photoperiodischen Bedingungen, unter denen die Käfigvögel lebten. Unterschiedliche Muster der Tagesaktivität in den einzelnen Jahreszeiten deuten das Zugverhalten der Art an: Bergfinken sind sowohl Nacht- als auch Tagzieher. Die Menge der Tagesaktivität pro Stunde war deutlich positiv mit der Länge der Photoperiode korreliert. Der photoperiodische und endogene Einfluß auf das Zugverhalten des Bergfinken wird diskutiert.

     

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.