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.     

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.     

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.     

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

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

Comparison of autumn and spring migration strategies of Neotropical migratory landbirds in northeast Belize

ABSTRACT Migration represents one of the most vulnerable stages of a migrant's life cycle, but the strategies and stopover sites used by Neotropical migrants in Central America are not well known. We carried out constant‐effort mist netting and conducted censuses along transects during one autumn (2007) and one spring (2008) migration in northeast Belize. We recorded more landbird migrant species in autumn (63) than in spring (54), and spring abundance was >25% lower for 88% of transient species. These differences in presence and abundance indicate that routes and stopover strategies vary between seasons and species. In autumn, fuel loads, calculated as any increase in mass above lean body mass (LBM), were generally small (mean = 5.9% LBM and 10.1% LBM for wintering and transient species, respectively) and fuel deposition rates and minimum stopover durations suggest that some individuals replenished energy reserves in our study area. Variation in autumn fuel loads meant that some individuals had reserves sufficient for flights >1000 km. Fuel loads were larger in spring for 16 of 17 species, and the mean spring fuel load for transient species (32.5% LBM) was sufficient for a flight from northeast Belize to North America without refueling. The similarity in spring passage times between northeast Belize and the Gulf Coast of the United States also suggests that energy reserves were not replenished in northeast Belize prior to crossing the Gulf of Mexico. We hypothesize that sufficient energy reserves are accumulated during spring stopovers in northern South America or elsewhere in Mesoamerica to allow migrants to fly directly to North America without refueling.     

Faster spring migration in northern wheatears is not explained by an endogenous seasonal difference in refueling rates

A widespread phenomenon in migrant birds is that they travel faster in spring than in autumn. During migration birds spend most time at stopover sites and, correspondingly, the faster spring migration is mainly explained by shorter stopovers in spring than autumn. Because a main purpose of stopovers is to replenish the fuel used in flight, a higher rate of fuel deposition (FDR) in spring is thought to explain the shorter stopovers and hence shorter total duration of migration in spring. Critical migratory processes, including the onset and extent of pre‐migratory fueling, are endogenously regulated. It is therefore not unlikely that refueling at stopover sites is, at least partly, also under endogenous control. We here tested whether there is an endogenous seasonal difference in food intake and FDR, which could contribute to shorter stopovers and hence faster migration in spring. We measured daily food intake and daily FDR in two subspecies of the northern wheatear Oenanthe oenanthe, temporarily confined at stopover under identical constant indoor conditions in spring and autumn. The two wheatear subspecies differed markedly in absolute food intake and FDR. Within subspecies, however, food intake and FDR did not differ between spring and autumn, indicating that faster spring migration in northern wheatears is not explained by an endogenously controlled seasonal difference in birds’ motivation to refuel. To further substantiate this claim, similar measurements should be taken at other locations along northern wheatears’ migration routes. Comparable experiments in other species could test the generality of our results.     

Songbirds are resilient to hurricane disturbed habitats during spring migration

The Gulf of Mexico is a conspicuous feature of the Neotropical–Nearctic bird migration system. Traveling long distances across ecological barriers comes with considerable risks, and mortality associated with intercontinental migration may be substantial, including that caused by storms or other adverse weather events. However, little, if anything, is known about how migratory birds respond to disturbance‐induced changes in stopover habitat. Isolated, forested cheniere habitat along the northern coast of the Gulf of Mexico often concentrate migrants, during weather conditions unfavorable for northward movement or when birds are energetically stressed. We expected hurricane induced degradation of this habitat to negatively affect the abundance, propensity to stopover, and fueling trends of songbirds that stopover in coastal habitat. We used spring banding data collected in coastal Louisiana to compare migrant abundance and fueling trends before (1993–1996 and 1998–2005) and after hurricanes Rita (2006) and Ike (2009). We also characterized changes in vegetative structure before (1995) and after (2010) the hurricanes. The hurricanes caused dramatic changes to the vegetative structure, which likely decreased resources. Surprisingly, abundance, propensity to stopover, and fueling trends of most migrant species were not influenced by hurricane disturbance. Our results suggest that: 1) the function of chenieres as a refuge for migrants after completing a trans‐Gulf flight may not have changed despite significant changes to habitat and decreases in resource availability, and 2) that most migrants may be able to cope with habitat disturbance during stopover. The fact that migrants use disturbed habitat points to their conservation value along the northern coast of the Gulf of Mexico.     

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.     

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。     

Foraging behaviour and fuel accumulation of capital breeders during spring migration as derived from a combination of satellite‐ and ground‐based observations

The migration strategy of many capital breeders is to garner body stores along the flyway at distinct stopover sites. The rate at which they can fuel is likely to be strongly influenced by a range of factors, such as physiology, food availability, time available for foraging and perceived predation. We analysed the foraging behaviour and fuel accumulation of pink‐footed geese, an Arctic capital breeder, at their mid‐flyway spring stopover site and evaluated to what extent their behaviour and fuelling were related to physiological and external factors and how it differed from other stopovers along the flyway. We found that fuel accumulation rates of geese at the mid‐flyway site were limited by habitat availability rather than by digestive constraints. However, as the time available for foraging increased over the stopover season, geese were able to keep constant fuelling rate. Putting this in perspective, geese increased their daily net energy intake along the flyway corresponding to the increase in time available for foraging. The net energy intake per hour of foraging remained the same. Geese showed differences in their reaction to predators/disturbance between the sites, taking higher risks particularly at the final stopover site. Hence, perceived predation along the flyway may force birds to postpone the final fuel accumulation to the last stopover along the flyway. Flexibility in behaviour appears to be an important trait to ensure fitness in this capital breeder. Our findings are based on a new, improved method for estimating fuel accumulation of animals foraging in heterogeneous landscapes based on data obtained from satellite telemetry and habitat specific intake rates.     

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.     

Towards a new understanding of migration timing: slower spring than autumn migration in geese reflects different decision rules for stopover use and departure

According to migration theory and several empirical studies, long‐distance migrants are more time‐limited during spring migration and should therefore migrate faster in spring than in autumn. Competition for the best breeding sites is supposed to be the main driver, but timing of migration is often also influenced by environmental factors such as food availability and wind conditions. Using GPS tags, we tracked 65 greater white‐fronted geese Anser albifrons migrating between western Europe and the Russian Arctic during spring and autumn migration over six different years. Contrary to theory, our birds took considerably longer for spring migration (83 days) than autumn migration (42 days). This difference in duration was mainly determined by time spent at stopovers. Timing and space use during migration suggest that the birds were using different strategies in the two seasons: In spring they spread out in a wide front to acquire extra energy stores in many successive stopover sites (to fuel capital breeding), which is in accordance with previous results that white‐fronted geese follow the green wave of spring growth. In autumn they filled up their stores close to the breeding grounds and waited for supportive wind conditions to quickly move to their wintering grounds. Selection for supportive winds was stronger in autumn, when general wind conditions were less favourable than in spring, leading to similar flight speeds in the two seasons. In combination with less stopover time in autumn this led to faster autumn than spring migration. White‐fronted geese thus differ from theory that spring migration is faster than autumn migration. We expect our findings of different decision rules between the two migratory seasons to apply more generally, in particular in large birds in which capital breeding is common, and in birds that meet other environmental conditions along their migration route in autumn than in spring.     

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.     

Niche dynamics of shorebirds in Delaware Bay: Foraging behavior,habitat choice and migration timing

Niche differentiation through resource partitioning is seen as one of the most important mechanisms of diversity maintenance contributing to stable coexistence of different species within communities. In this study, I examined whether four species of migrating shorebirds, dunlins (Calidris alpina), semipalmated sandpipers (Calidris pusilla), least sandpipers (Calidris minutilla) and short-billed dowitchers (Limnodromus griseus), segregate by time of passage, habitat use and foraging behavior at their major stopover in Delaware Bay during spring migration. I tested the prediction that most of the separation between morphologically similar species will be achieved by differential migration timing. Despite the high level of overlap along observed niche dimensions, this study demonstrates a certain level of ecological separation between migrating shorebirds. The results of analyses suggest that differential timing of spring migration might be the most important dimension along which shorebird species segregate while at stopover in Delaware Bay. Besides differences in time of passage, species exhibited differences in habitat use, particularly least sandpipers that foraged in vegetated areas of tidal marshes more frequently than other species, as well as short-billed dowitchers that foraged in deeper water more often than small sandpipers did. Partitioning along foraging techniques was less prominent than segregation along temporal or microhabitat dimensions. Such ranking of niche dimensions emphasizes significance of temporal segregation of migratory species – separation of species by time of passage may reduce the opportunity for interspecific aggressive encounters, which in turn can have positive effects on birds' time and energy budget during stopover period.     

Do Birds Select Habitat or Food Resources? Nearctic-Neotropic Migrants in Northeastern Costa Rica

Nearctic-neotropic migrant birds need to replenish energy reserves during stopover periods to successfully complete their semiannual movements. In this study we used linear models to examine the habitat use of 11 migrant species in northeastern Costa Rica to better understand the influence of food and structural resources on the presence of birds during stopover periods. Our models indicated that frugivorous migrants primarily used food abundance, while insectivorous migrants chiefly used vegetation structure as cues for habitat use during stopover. In addition to habitat use models, we documented fruiting plant phenology and found a general relationship between migrant arrival and the timing of ripe fruit availability. Our results suggest that insectivorous migrants probably rely on structural features when using habitat because it may be inherently difficult to assess cryptic-arthropod availability during a short period of time in a novel habitat, such as stopover periods.     

Migration of two calidrid sandpiper species on the predator landscape: how stopover time and hence migration speed vary with geographical proximity to danger

The effects of relative fuel load on migration speed and on vulnerability have been investigated, but the effects of seasonal variation in predation danger on the amount of fuel and duration of stopover have not been considered. We analyzed seasonal patterns of stopover residence times for western and semipalmated sandpipers Calidris mauri and C. pusilla on southward migration in relation to the passage of migratory peregrine falcons Falco peregrinus. We predicted that individuals on stopover far in advance of the seasonal arrival of falcons would adjust stopover length and hence relative fuel load to migrate slowly and cautiously. We predicted that individuals on stopover later in the season would increase migratory speed as the arrival of migratory falcons came closer, while individuals on stopover under or behind the passage of falcons would migrate slowly. Adult and juvenile semipalmated and adult western sandpipers migrated prior to seasonal increases in peregrine abundance, and as predicted, the seasonal patterns of their stopover durations are consistent with an increase in the speed of migration as the date of peregrine arrival approached. Juvenile western sandpipers, in contrast, migrating concurrently with falcons, slowed their speed of migration as predator abundance increased. Stopover patterns differ between species due to different relative fuel loads. The results fit predictions made based on a ‘mortality‐minimizing’ migration strategy.     

Use of a Florida Gulf Coast Barrier Island by Spring Trans-Gulf Migrants and the Projected Effects of Sea Level Rise on Habitat Availability

Barrier islands on the north coast of the Gulf of Mexico are an internationally important coastal resource. Each spring hundreds of thousands of Nearctic-Neotropical songbirds crossing the Gulf of Mexico during spring migration use these islands because they provide the first landfall for individuals following a trans-Gulf migratory route. The effects of climate change, particularly sea level rise, may negatively impact habitat availability for migrants on barrier islands. Our objectives were (1) to confirm the use of St. George Island, Florida by trans-Gulf migrants and (2) to determine whether forested stopover habitat will be available for migrants on St. George Island following sea level rise. We used avian transect data, geographic information systems, remote sensing, and simulation modelling to investigate the potential effects of three different sea level rise scenarios (0.28 m, 0.82 m, and 2 m) on habitat availability for trans-Gulf migrants. We found considerable use of the island by spring trans-Gulf migrants. Migrants were most abundant in areas with low elevation, high canopy height, and high coverage of forests and scrub/shrub. A substantial percentage of forest (44%) will be lost by 2100 assuming moderate sea level rise (0.82 m). Thus, as sea level rise progresses, less forests will be available for migrants during stopover. Many migratory bird species’ populations are declining, and degradation of barrier island stopover habitat may further increase the cost of migration for many individuals. To preserve this coastal resource, conservation and wise management of migratory stopover areas, especially near ecological barriers like the Gulf of Mexico, will be essential as sea levels rise.     

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.     

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