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。     

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.     

Stopover ecology of a migratory ungulate

1. Birds that migrate long distances use stopover sites to optimize fuel loads and complete migration as quickly as possible. Stopover use has been predicted to facilitate a time-minimization strategy in land migrants as well, but empirical tests have been lacking, and alternative migration strategies have not been considered. 2. We used fine-scale movement data to evaluate the ecological role of stopovers in migratory mule deer Odocoileus hemionus- a land migrant whose fitness is strongly influenced by energy intake rather than migration speed. 3. Although deer could easily complete migrations (range 18-144 km) in several days, they took an average of 3 weeks and spent 95% of that time in a series of stopover sites that had higher forage quality than movement corridors. Forage quality of stopovers increased with elevation and distance from winter range. Mule deer use of stopovers corresponded with a narrow phenological range, such that deer occupied stopovers 44 days prior to peak green-up, when forage quality was presumed to be highest. Mule deer used one stopover for every 5·3 and 6·7 km travelled during spring and autumn migrations, respectively, and used the same stopovers in consecutive years. 4. Study findings indicate that stopovers play a key role in the migration strategy of mule deer by allowing individuals to migrate in concert with plant phenology and maximize energy intake rather than speed. Our results suggest that stopover use may be more common among non-avian taxa than previously thought and, although the underlying migration strategies of temperate ungulates and birds are quite different, stopover use is important to both. 5. Exploring the role of stopovers in land migrants broadens the scope of stopover ecology and recognizes that the applied and theoretical benefits of stopover ecology need not be limited to avian taxa.     

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.     

Danger management and the seasonal adjustment of migratory speed by sandpipers

The behavior of migrating birds is governed by time‐, energy‐ and danger‐minimizing strategies. The adjustment of migration speed (i.e. the rate at which distance is covered during a migration) is a behavioral tactic that might contribute to these strategic goals. Shorter stopovers and greater fuel loads increase migration speed, but both require more intensive foraging at stopovers, making migrants more vulnerable to predators. A simple numerical model shows how seasonal alterations in migration speed can lower the exposure of western sandpipers to peregrine falcons, their most important predator. The ‘caution–speed–caution’ pattern of higher migration speed in the mid‐passage period, observed in earlier work, requires that the intensive foraging necessary heightens vulnerability, and that migrants are exposed to both migrant predators as well as predators resident at migratory stopovers.     

Spring migration strategies of Whinchat Saxicola rubetra when successfully crossing potential barriers of the Sahara and the Mediterranean Sea

The flexibility for migrant land birds to be able to travel long distances rapidly without stopovers, and thus to cross wide inhospitable areas such as deserts and oceans, is likely to be a major determinant of their survival during migration. We measured variation in flight distance, speed and duration of major stopovers (more than 2 days), using geolocator tracks of 35 Whinchats Saxicola rubetra that migrated successfully from central Nigeria to Eastern Europe in spring, and examined how these measures changed, or depended on age, when crossing the barriers of the Sahara or the Mediterranean Sea. In all, 31% of Whinchats crossed at least the Sahara and the Mediterranean before a major stopover and 17% travelled over 4751 km on average without any major stopovers. Flight distance and speed during, and duration of major stopovers after, crossing the Mediterranean Sea were indistinguishable from migration over Continental Europe. Speed during a migration leg was lowest crossing Continental Europe and fastest, with longer duration major stopovers afterwards, when crossing the Sahara, but there was much individual variation, and start date of migration was also a good predictor of stopover duration. As the distance travelled during a leg increased, so major stopover duration afterwards increased (1 day for every 1000 km), but the speed of travel during the leg had no effect. There were no differences in any migration characteristics with age, other than an earlier start date for adult birds. The results suggest that adaptive shortening or even dropping of daily stopovers may occur often, allowing rapid, long‐distance migration at the cost of major stopovers afterwards, but such behaviour is not restricted to or always found when crossing barriers, even for birds on their first spring migration. The results may highlight the importance of stopover sites rather than barrier width as the likely key component to successful migration. Individual variation in spring migration may indicate that small passerine migrants like Whinchats may be resilient to future changes in the extent of barriers they encounter, although this may not be true of first autumn migrations or if stopover sites are lost.     

Male post-breeding movements and stopover habitat selection of an endangered short-distance migrant,the Little Bustard Tetrax tetrax

Migratory decisions, such as the selection of stopover sites, are critical for the success of post-breeding migratory movements and subsequent survival. Recent advances in bio-logging have revealed the stopover strategies of many long-distance migrants, but far less attention has been given to short-distance migrants. We investigated the stopover ecology of an endangered grassland bird, the Little Bustard Tetrax tetrax, a short-distance migrant in Iberia. Using high-resolution spatial GPS/GSM data, 27 male Little Bustards breeding in southern Portugal were tracked between 2009 and 2011. We studied post-breeding movements using Dynamic Brownian Bridge models to identify the main stopover sites, and generalized linear mixed models to examine habitat selection in stopovers. During their post-breeding movements, males were essentially nocturnal migrants, making frequent stopovers while maintaining a relatively fast pace to reach more productive agricultural post-breeding areas. Stopovers occurred in most post-breeding movements (83%) regardless of the total distance covered (average 64.3 km), and most stopovers (84%) lasted less than 24 h. Birds used mostly agricultural non-irrigated and irrigated croplands as stopover sites and avoided other land uses and rugged terrain. There was a negative relationship between stopovers and the proximity to roads, but not to power lines. The high frequency of stopovers during post-breeding movements, despite the short distances travelled, together with the nocturnal migratory behaviour of bustards, may impose additional risks to a bird mainly threatened by collision with power lines in non-breeding areas. We also conclude that even for short-distance migrants, habitat connectivity between breeding and post-breeding areas is likely to be a key conservation concern.     

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.     

Effects of migration distance and sex on stopover timing and refueling by Wilson's Warblers

Migration distance and sex are integral to avian migration strategies, yet these intrinsic factors are understudied with respect to their effects on stopover ecology and behavior. We investigated how individual variation in migration distance and sex affected body condition, refueling performance, and stopover durations of Wilson's Warblers (Cardellina pusilla) during spring and fall 2006 at a stopover site in southern Arizona. Migration distance of individuals was inferred using the hydrogen stable isotope ratio of feathers (δ²H_f) as an index of breeding latitude, refueling rate was assessed using plasma triglyceride and β‐OH‐butyrate, and minimum length of stay was estimated by re‐sighting color‐banded birds. In the spring, migration distance and sex were strong determinants of the timing of migration by Wilson's Warblers, with males and shorter‐distance migrants passing through the site earlier than females and longer‐distance migrants. Later‐arriving migrants also had higher fat scores and refueling rates. However, neither migration distance nor sex independently affected body condition (fat score or size‐corrected mass), refueling rate, or minimum length of stay. In a smaller sample of fall migrants, we found that longer‐distance migrants and males had higher refueling rates than shorter‐distance migrants and females. Our results show that differences in migration distance can, under some circumstances, affect how birds use stopover sites. Stable isotope analysis and other methods can provide information on migration distance, and should be integrated with measurements of timing, fuel stores, refueling performance, and departure behavior to gain a deeper understanding of bird migration.     

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.     

Heterothermy, and the energetic consequences of huddling in small migrating passerine birds

The success of migration of small passerine birds depends largely on effective refueling at stopover sites. In our previous studies, we found that hypothermia facilitates accumulation of fuel at the beginning of a stopover. Later we found that blackcaps, Sylvia atricapilla, might further reduce their energy expenditure by huddling while at rest. Here, we report experimental results supporting our hypothesis that huddling is beneficial to small migrating passerines both from energetic and thermoregulatory points of view. To test this hypothesis we measured metabolic rates and body temperatures of seven blackcaps placed in respirometry chambers overnight, either solitarily or in groups of three or four at ambient temperatures of 5, 10, and 15°C. Concurring with our predictions, huddling blackcaps maintained higher body temperatures than did solitary birds, but had mass-specific metabolic rates lower by ～30% than those of solitary individuals. Based on our previous studies, we estimated energy savings through huddling to be comparable to energy savings through hypothermia in solitary blackcaps and suggest that huddling may be an important way of saving energy for small passerine birds resting at migratory stopovers. At the same time it might offer the additional benefit of lower risk of predation. In this light, we predict that huddling occurs frequently in nature, leading to significant savings of energy, faster accumulation of fuel, presumably lower risk of becoming a prey, more successful migration, and eventually increased fitness.     

Why fly the extra mile? Latitudinal trend in migratory fuel deposition rate as driver of trans‐equatorial long‐distance migration

Trans‐equatorial long‐distance migrations of high‐latitude breeding animals have been attributed to narrow ecological niche widths. We suggest an alternative hypothesis postulating that trans‐equatorial migrations result from a possible increase in the rate at which body stores to fuel migration are deposited with absolute latitude; that is, longer, migrations away from the breeding grounds surpassing the equator may actually enhance fueling rates on the nonbreeding grounds and therewith the chance of a successful, speedy and timely migration back to the breeding grounds. To this end, we first sought to confirm the existence of a latitudinal trend in fuel deposition rate in a global data set of free‐living migratory shorebirds and investigated the potential factors causing this trend. We next tested two predictions on how this trend is expected to impact the migratory itineraries on northward migration under the time‐minimization hypothesis, using 56 tracks of high‐latitude breeding shorebirds migrating along the East Asian‐Australasian Flyway. We found a strong positive effect of latitude on fuel deposition rate, which most likely relates to latitudinal variations in primary productivity and available daily foraging time. We next confirmed the resulting predictions that (1) when flying from a stopover site toward the equator, migrants use long jumps that will take them to an equivalent or higher latitude at the opposite hemisphere; and (2) that from here onward, migrants will use small steps, basically fueling only enough to make it to the next suitable staging site. These findings may explain why migrants migrate “the extra mile” across the equator during the nonbreeding season in search of better fueling conditions, ultimately providing secure and fast return migrations to the breeding grounds in the opposite hemisphere.     

Spring migration dynamics and sex-specific patterns in stopover strategy in the Wood Sandpiper Tringa glareola

Due to being a virtually monomorphic wader species, migration dynamics and sex-related migration patterns in the Wood Sandpiper (Tringa glareola) have rarely been investigated. We captured spring migrants at an important stopover site in northeastern Austria. Birds were individually color-marked, and sex was determined by an analysis of DNA from tail feather material. Among temporary residents (birds seen again after day of capture), males migrated on average 3 days earlier than females. However, since sexes did not differ in fat score, the length of stay and the proportion of transients (birds not seen again after day of capture) and temporary residents, we suggest that males and females adopt similar migration strategies in the spring. The large number of transients captured as well as shorter stopover durations in later temporary residents indicate that Wood Sandpipers minimize time at this stage of their northbound migration. Temporary residents earlier in the season exhibited lower fat stores than later ones. Nevertheless, since the fat stores of transients and temporary residents were similar even after the progress of the season had been accounted for, we assume that Wood Sandpipers may afford to exhibit individual flexibility in migration strategy and the use of stopover sites, especially early in the season. This variability may be a necessary adaptation to cope with possible varying environmental conditions at dynamic and unpredictable inland stopover sites. After having reached North Mediterranean regions, mean body mass of spring migrants gradually increases during successive stopovers, indicating that Wood Sandpipers follow a ‘hopping’ migration strategy. This emphasizes the high conservation value of even small artificial mudflat pools as important stepping stones in order to maintain a continuous network of wetland habitats for this continental migrant.     

A place to land: spatiotemporal drivers of stopover habitat use by migrating birds

Migrating birds require en route habitats to rest and refuel. Yet, habitat use has never been integrated with passage to understand the factors that determine where and when birds stopover during spring and autumn migration. Here, we introduce the stopover‐to‐passage ratio (SPR), the percentage of passage migrants that stop in an area, and use 8 years of data from 12 weather surveillance radars to estimate over 50% SPR during spring and autumn through the Gulf of Mexico and Atlantic coasts of the south‐eastern US, the most prominent corridor for North America’s migratory birds. During stopovers, birds concentrated close to the coast during spring and inland in forested landscapes during autumn, suggesting seasonal differences in habitat function and highlighting the vital role of stopover habitats in sustaining migratory communities. Beyond advancing understanding of migration ecology, SPR will facilitate conservation through identification of sites that are disproportionally selected for stopover by migrating birds.     

Experimental evidence for the interplay of exogenous and endogenous factors on the movement ecology of a migrating songbird

Movement patterns during songbird migration remain poorly understood despite their expected fitness consequences in terms of survival, energetic condition and timing of migration that will carry over to subsequent phases of the annual cycle. We took an experimental approach to test hypotheses regarding the influence of habitat, energetic condition, time of season and sex on the hour-by-hour, local movement decisions of a songbird during spring stopover. To simulate arrival of nocturnal migrants at unfamiliar stopover sites, we translocated and continuously tracked migratory red-eyed vireos (Vireo olivaceus) throughout spring stopover with and without energetic reserves that were released in two replicates of three forested habitat types. Migrants moved the most upon release, during which time they selected habitat characterized by greater food abundance and higher foraging attack rates. Presumably under pressure to replenish fuel stores necessary to continue migration in a timely fashion, migrants released in poorer energetic condition moved faster and further than migrants in better condition and the same pattern was true for migrants released late in spring relative to those released earlier. However, a migrant's energetic condition had less influence on their behavior when they were in poor quality habitat. Movement did not differ between sexes. Our study illustrates the importance of quickly finding suitable habitat at each stopover site, especially for energetically constrained migrants later in the season. If an initial period prior to foraging were necessary at each stop along a migrant's journey, non-foraging periods would cumulatively result in a significant energetic and time cost to migration. However, we suggest behavior during stopover is not solely a function of underlying resource distributions but is a complex response to a combination of endogenous and exogenous factors.     

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]。     

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.     

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.     

迁徙鸟类对中途停歇地的利用及迁徙对策

中途停歇地是迁徙鸟类在繁殖地和非繁殖地之间的联系枢纽,对于迁徙鸟类完成其完整的生活史过程具有重要作用。从鸟类的迁徙对策、中途停歇地的选择、鸟类在中途停歇地的停留时间、体重变化和种群特征以及中途停歇地的环境状况等方面,回顾了中途停歇生态学在近年来的研究进展,并提出了在迁徙对策理论的实验研究,小型鸟类在中途停歇地的停歇时间及体重变化的准确确定等目前有待解决的问题。     

Fuel deposition of three passerine bird species along the migration route

The rate at which migrant birds replenish their energy stores at intermittent stopovers largely determines overall migration speed, the manner in which migration proceeds and success of migration. In this study, data on the fuel deposition rate (FDR) of three long-distance migrants from 17 ringing sites along their autumn migration route were used to examine: (1) effects of endogenous factors on FDR, and (2) how relationships between exogenous factors and FDR affect the organisation of migration. We developed a model to estimate FDR from retrapped birds which takes into account time of day and various other factors which might influence FDR. The two endogenous factors, moult and current energy stores, generally reduced FDR. This may result in lower departure energy loads and more stopovers than expected from optimal migration theory. Differences between species with respect to seasonal, year-to-year and geographical patterns of FDR could be related to differences in availability and predictability of food resources, and help to explain differences in the organisation of migration. A low FDR in northern and central Europe could be related to low, but predictable, food resources and an early departure during moult of the reed warbler (Acrocephalus scirpaceus); FDRs varying between years were related to large spatial and year-to-year variation in the density of the main prey of the sedge warbler (Acrocephalus schoenobaenus); and a high FDR in the garden warbler (Sylvia borin) was related to abundant food resources, due to a switch from a purely invertebrate diet to a mixed diet including fruits which are abundant over large areas of Europe and north Africa. This study demonstrated that the organisation of migration is the outcome of a complex interplay of the seasonal timing of moult, food availability and predictability and a seasonal switch in diet, and can be modified by individual birds in response to a limited amount of time in which to migrate. Received: 26 April 1999 / Accepted: 24 September 1999