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.     

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

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

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.     

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

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

Warming winter effects,fat store accumulation and timing of spring departure of Greenland White-fronted Geese <Emphasis Type="Italic">Anser albifrons flavirostris</Emphasis> from their winter quarters

Migratory geese accumulate energy and nutrient stores in winter to fly to refuelling spring staging areas before onward migration to breeding areas. Mean ground temperatures at two important Greenland White-fronted Geese wintering sites rose in winter and spring by 1.0–1.3°C during 1973–2007. Greenland White-fronted Geese departed the Wexford winter quarters on 3rd April 2007 for Icelandic spring staging areas, the earliest on record, representing a mean advancement of 15 days since 1973, mirrored amongst mean dates of departure amongst Scottish wintering birds that have advanced by 12 days during 1973–2007. Icelandic temperatures at critical midway staging areas en route to Greenland showed no significant change since 1973, suggesting that it is warming on the winter quarters that enable geese to depart earlier, rather than elevated temperatures at ultimate spring staging areas. However, Wexford departure date did not correlate with spring temperature. Data presented here show that Greenland White-fronted Geese have accumulated threshold body stores progressively earlier in spring migration, especially during 1995–2007. Although this did not correlate with ambient temperature, the mean degree of accumulated fat stored by 1st April in each year was a statistically significant predictor of departure date for the wintering population at Wexford. These data support the hypothesis that it is intrinsic factors (i.e. improvements in internal body state resulting from better feeding conditions) that has permitted progressively earlier departure of these geese from Wexford on spring migration, rather than amelioration of spring conditions in Iceland or solely the result of warming of the winter quarters.     

Eggs brought in from afar: Svalbard‐breeding pink‐footed geese can fly their eggs across the Barents Sea

Many Arctic‐breeding waterbirds are thought to bring nutrients for egg production from southern latitudes to allow early breeding. It has proved problematic to quantify the extent of such capital breeding and identify whether nutrients for egg production are brought in from nearby or from afar. Before reaching their breeding grounds on Svalbard, pink‐footed geese Anser brachyrhynchus fly ～ 1100 km across the Barents Sea from Norway. Using abdominal profile indexing (API) we scored body stores in individually marked geese just prior to migration from the northernmost staging area in Norway to Svalbard, followed by their breeding success on their non‐breeding grounds in autumn. In productive breeding years leading to a high (> 13.8%) proportion of juveniles in the autumn population, there was a positive relationship between female API and number of young produced, suggesting that the geese are at least partial capital breeders. Moreover, focusing on the geographic origin of proteins used in egg synthesis and measuring nitrogen stable isotope ratios in pink‐footed geese's eggs and food sources in Norway and Svalbard, we identified that capital breeding in this species is ～ 50% on average but may potentially amount to as much as 100%, notably in females laying early. About 60% of this protein capital is carried in well‐developed follicles across the Barents Sea, the remainder likely being stored in muscle tissues. Conditions on the wintering grounds and migratory stopover sites can have profound effects on an individual's fitness but the here presented link between the use of migratory stopover sites and breeding performance is particularly noteworthy. Apparently, some individuals accept the putative costs of carrying body stores over large distances to the breeding grounds. The data also highlights considerable variation in the reliance on capital for breeding, suggesting substantial individual scope to adjust breeding strategy to changing environmental conditions.     

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.     

Demographic response to environmental variation in breeding,stopover and non-breeding areas in a migratory passerine

Demographic rates of migratory species passing through several areas during their annual cycle may be affected by environmental conditions at each of these areas. Recent studies provide evidence that their impact is not necessarily immediate, but can be delayed. We studied survival, reproductive success and arrival date at the breeding grounds of red-backed shrikes Lanius collurio, a trans-Saharan migrant, in relation to weather and vegetation on the breeding grounds, the stopover sites during migration and in the wintering areas. These environmental factors are used as proxy of the shrike’s food supply. We analysed detailed demographic data of some 4,600 individuals from 25 years with multistate capture–recapture and mixed models. Survival probabilities of juveniles and breeders of both sexes varied in parallel across time, suggesting that all cohorts were sensitive to similar causes of mortality. Reproductive performance increased with temperature and decreased with rainfall on the breeding area. Moreover, it increased with vegetation cover in the Sahelian stopover area used on autumn migration suggesting a carry-over effect. Arrival date was negatively affected by spring temperatures in the breeding area. Hence, demographic rates were affected by environmental factors on the breeding grounds, but also outside and elsewhere. This suggests that the shrike’s population dynamics are driven by environmental factors operating at various scales of space and time. However, only a small amount of the temporal variation in demographic rates is explained by the environmental factors considered, suggesting that additional factors, such as those operating during migration, might be important.     

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.     

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。     

Ecological factors associated with the breeding and migratory phenology of high-latitude breeding western sandpipers

Environmental conditions influence the breeding and migratory patterns of many avian species and may have particularly dramatic effects on long-distance migrants that breed at northern latitudes. Environment, however, is only one of the ecological variables affecting avian phenology, and recent work shows that migration tactics may be strongly affected by changes in predator populations. We used long-term data from 1978 to 2000 to examine the interactions between snowmelt in western Alaska in relation to the breeding or migration phenologies of small shorebirds and their raptor predators. Although the sandpipers’ time of arrival at Alaskan breeding sites corresponded with mean snowmelt, late snowmelts did delay breeding. These delays, however, did not persist to southward migration through British Columbia, likely due to the birds’ ability to compensate for variance in the length of the breeding season. Raptor phenology at an early stopover site in British Columbia was strongly related to snowmelt, so that in years of early snowmelt falcons appeared earlier during the sandpipers’ southbound migration. These differential effects indicate that earlier snowmelt due to climate change may alter the ecological dynamics of the predator–prey system.     

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.     

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

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

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]。     

Extensive Use of Intertidal Habitat by Shorebirds Outside Protected Nesting Areas

Conservation of shorebirds throughout their breeding and migratory ranges has become a priority as shorebird populations decline globally. Along the North Atlantic Coast, management efforts have particularly focused on preserving nesting habitat for piping plovers (Charadrius melodus), which are protected under the Endangered Species Act. It is unclear whether these conservation measures suffice to protect foraging habitat for piping plovers and other shorebirds on stopover during migration along the Atlantic Flyway. To evaluate the extent to which conservation of piping plover nesting areas extends to all habitats used by plovers, and to determine whether these protections also benefited non-breeding migratory shorebirds in the region, we conducted weekly shorebird surveys, recording the number and locations of piping plovers and other species, during northward and southward migration on Fire Island and Westhampton Island, New York, USA, from 2014–2016. We used canonical correspondence analysis (CCA) to assess the degree of spatiotemporal overlap between breeding plovers, foraging plovers, and other migratory shorebirds that temporarily stage at the site. The spatiotemporal distribution of migratory shorebirds matched more closely with piping plovers seen during foraging than piping plovers observed tending nests and engaging in other breeding activities. Migratory shorebirds and foraging piping plovers were more abundant and frequent in wet intertidal zones outside of fenced-off nesting areas, which were not protected under current management regimes. Therefore, additional protection of piping plover foraging habitat could benefit plovers and migratory shorebirds that use similar feeding grounds during stopover on northward and southward migration. © 2020 The Wildlife Society.     

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

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