雌性藏羚迁徙对青藏高原降水时空分布的适应性分析

藏羚主要分布在青藏高原,雌性有季节性的生殖迁徙行为。产仔前,分布在各地的雌性藏羚沿着比较固定的路线向青藏高原西北部可可西里的太阳湖、卓乃湖、阿尔金山保护区的慕孜塔格峰等产羔地迁徙,产羔后一段时间又返回原栖息地。对这种特殊的迁徙现象,已有一些假说用来解释,但都没有得到普遍认可。青藏高原由于其海拔高,常年温度低,在藏羚分布区降水主要以固态形式为主,并且青藏高原降水量由东南部向西北部递减,结合青藏高原地质历史时期植被演变的特点和藏羚迁徙的方向,本文提出了一个新假设,以此来解释藏羚迁徙的原因。我们的假设是:藏羚迁往产羔地产羔是为了躲避固态降水相对丰富的地区,以保证新生羚羊有高的存活率;藏羚迁往的产羔地曾是它们的故乡。笔者主要分析了提出这个假设的根据。     

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.     

Physiological, behavioral, and ecological aspects of migration in reptiles

Seasonal movements between foraging, breeding, and overwintering sites occur in a wide variety of reptile species. Terrestrial snakes, lizards, and turtles migrate short distances (<20 km) between seasonal habitats, whereas fully aquatic marine turtles migrate hundreds to thousands of kilometers between foraging and breeding areas. The purpose of this article is to summarize aspects of migratory physiology and behavior in reptiles, particularly with regards to energetics and sensory mechanisms for navigation and orientation. We discuss the influence of aerobic scope, endurance, and cost of transport on migratory capacity, the effects of temperature and circulating hormones on activity and behavior, and mechanisms of detecting and transducing environmental cues to successfully navigate and orient toward a goal during migration. Topics worthy of further research are highlighted in the text, and we conclude with a discussion of how information on migration patterns of reptiles may be used to manage and conserve threatened populations.     

Range-wide migration corridors and non-breeding areas of a northward expanding Afro-Palaearctic migrant,the European Bee-eater Merops apiaster

Across their ranges, different populations of migratory species often use separate routes to migrate between breeding and non-breeding grounds. Recent changes in climate and land-use have led to breeding range expansions in many species but it is unclear whether these populations also establish new migratory routes, non-breeding sites and migration phenology. Thus, we compared the migration patterns of European Bee-eaters Merops apiaster from two established western (n = 5) and eastern (n = 6) breeding populations in Europe, with those from a newly founded northern population (n = 19). We aimed to relate the breeding populations to the two known non-breeding clusters in Africa, and to test for similarities of migration routes and timing between the old and new populations. Western Bee-eaters used the western flyway to destinations in West Africa; the eastern birds uniformly headed south to southern African non-breeding sites, confirming a complete separation in time and space between these long-established populations. The recently founded northern population, however, also used a western corridor, but crossed the Mediterranean further east than the western population and overwintered mainly in a new non-breeding area in southern Congo/northern Angola. The migration routes and the new non-breeding range overlapped only slightly with the western, but not with the eastern, population. In contrast, migration phenology appeared to differ between the western and both the northern and the eastern populations, with tracked birds from the western population migrating 2–4 weeks earlier. The northern population thus shares some spatial traits with western Bee-eaters, but similar phenology only with eastern population. This divergence highlights the adjustments in the timing of migration to local environmental conditions in newly founded populations, and a parallel establishment of new breeding and non-breeding sites.     

Weak migratory connectivity,loop migration and multiple non-breeding site use in British breeding Whinchats Saxicola rubetra

Determining the links between breeding populations and the pressures, threats and conditions they experience presents a challenge for the conservation of migratory birds which can use multiple sites separated by hundreds to thousands of kilometres. Furthermore, migratory connectivity – the connections made by migrating individuals between networks of breeding and non-breeding sites – has important implications for population dynamics. The Whinchat Saxicola rubetra is declining across its range, and tracking data from a single African non-breeding site implies high migratory spread. We used geolocators to describe the migration routes and non-breeding areas of 20 Whinchats from three British breeding populations. As expected, migratory spread was high, with birds from the three populations overlapping across a wide area of West Africa. On average, in non-breeding areas, British breeding Whinchats were located 652 km apart from one another, with some likely to share non-breeding areas with individuals from breeding populations as far east as Russia. Four males made a direct non-breeding season movement to a second, more westerly, non-breeding location in January. Autumn migration was through Iberia and around the western edge of the Sahara Desert, whereas spring migration was more direct, indicating an anticlockwise loop migration. Weak migratory connectivity implies that Whinchat populations are somewhat buffered against local changes in non-breeding conditions. If non-breeding season processes have played a role in the species’ decline, then large-scale drivers are likely to be the cause, although processes operating on migration, or interactions between breeding and non-breeding processes, cannot be ruled out.     

Annual Cycle and Migration Strategies of a Trans-Saharan Migratory Songbird: A Geolocator Study in the Great Reed Warbler

Recent technological advancements now allow us to obtain geographical position data for a wide range of animal movements. Here we used light-level geolocators to study the annual migration cycle in great reed warblers (Acrocephalus arundinaceus), a passerine bird breeding in Eurasia and wintering in sub-Saharan Africa. We were specifically interested in seasonal strategies in routes and schedules of migration. We found that the great reed warblers (all males, no females were included) migrated from the Swedish breeding site in early August. After spending up to three weeks at scattered stopover sites in central to south-eastern Europe, they resumed migration and crossed the Mediterranean Sea and Sahara Desert without lengthy stopovers. They then spread out over a large overwintering area and each bird utilised two (or even three) main wintering sites that were spatially separated by a distinct mid-winter movement. Spring migration initiation date differed widely between individuals (1-27 April). Several males took a more westerly route over the Sahara in spring than in autumn, and in general there were fewer long-distance travels and more frequent shorter stopovers, including one in northern Africa, in spring. The shorter stopovers made spring migration on average faster than autumn migration. There was a strong correlation between the spring departure dates from wintering sites and the arrival dates at the breeding ground. All males had a high migration speed in spring despite large variation in departure dates, indicating a time-minimization strategy to achieve an early arrival at the breeding site; the latter being decisive for high reproductive success in great reed warblers. Our results have important implications for the understanding of long-distance migrants’ ability to predict conditions at distant breeding sites and adapt to rapid environmental change.     

Consistent annual schedules in a migratory shorebird

Many migratory birds start prebreeding moult and premigratory fuelling some months before the breeding season and face severe time constraints, while travelling up to 15,000 km between non-breeding and breeding grounds. Shorebirds typically leave Southern Hemisphere non-breeding areas over a 3-4 week period, but whether they benefit from interannually consistent timing of departure is unknown. Here, I show that individual bar-tailed godwits (Limosa limosa baueri) from New Zealand are highly consistent in their migratory scheduling. Most birds left within the same week each year (between-year repeatability, r, of 0.83) and adult males, which moult into a bright breeding plumage, were also highly repeatable in the extent of their prebreeding moult (r=0.86). This is consistent with the hypothesis that birds have individually optimized migration schedules. Within adult males, but not females, smaller birds tended to migrate earlier than large birds. Whether this reflects differences in size-related migration speed, optimal breeding time at different sites or size-related natural or sexual selection pressures, remains unknown.     

Migratory connectivity and population-specific migration routes in a long-distance migratory bird

Knowledge about migratory connectivity, the degree to which individuals from the same breeding site migrate to the same wintering site, is essential to understand processes affecting populations of migrants throughout the annual cycle. Here, we study the migration system of a long-distance migratory bird, the Montagu''s harrier Circus pygargus, by tracking individuals from different breeding populations throughout northern Europe. We identified three main migration routes towards wintering areas in sub-Saharan Africa. Wintering areas and migration routes of different breeding populations overlapped, a pattern best described by ‘weak (diffuse) connectivity’. Migratory performance, i.e. timing, duration, distance and speed of migration, was surprisingly similar for the three routes despite differences in habitat characteristics. This study provides, to our knowledge, a first comprehensive overview of the migration system of a Palaearctic-African long-distance migrant. We emphasize the importance of spatial scale (e.g. distances between breeding populations) in defining patterns of connectivity and suggest that knowledge about fundamental aspects determining distribution patterns, such as the among-individual variation in mean migration directions, is required to ultimately understand migratory connectivity. Furthermore, we stress that for conservation purposes it is pivotal to consider wintering areas as well as migration routes and in particular stopover sites.     

Latitudinal distribution, migration, and testosterone levels in birds

Tropical bird species usually have lower testosterone (T) levels during breeding than temperate species. However, the potential mechanisms behind the positive interspecific correlation between T and latitude remain unexplored. In a comparative study of more than 100 bird species, we examined whether social constraints during male-male competition arising from migration and breeding synchrony are responsible for the latitude effects. Species that breed at higher latitudes are more likely to migrate and experience more intense intrasexual competition upon spring arrival than nonmigrant species from lower latitudes. Additionally, species from higher latitudes cope with shorter breeding seasons and thus with more synchronous breeding, which selects for high T titers via increased male-male conflicts. Accordingly, peak T levels were associated with migration and the duration of the egg laying period that reflects breeding synchrony. Because migration and breeding synchrony were related to latitudinal distribution, they appear to be important components of the latitude effects on T. A multivariate model controlling for covariation of predictor variables revealed that latitude remained the strongest predictor of peak T. Therefore, selection due to migration and breeding synchrony may partially cause the latitude effect, but other geographically varying factors may also play a role in mediating peak T levels at different latitudes.     

Seasonal matching of habitat quality and fitness in a migratory bird

When species occupy habitats that vary in quality, choice of habitat can be critical in determining individual fitness. In most migratory species, juveniles migrate independently of their parents and must therefore choose both breeding and winter habitats. Using a unique dataset of marked black-tailed godwits (Limosa limosa islandica) tracked throughout their migratory range, combined with analyses of stable carbon isotope ratios, we show that those individuals that occupy higher quality breeding sites also use higher quality winter sites. This seasonal matching can severely inflate inequalities in individual fitness. This population has expanded over the last century into poorer quality breeding and winter habitats and, across the whole population; individual birds tend to occupy either novel or traditional sites in both seasons. Winter and breeding season habitat selection are thus strongly linked throughout this population; these links have profound implications for a wide range of population and evolutionary processes. As adult godwits are highly philopatric, the initial choice of winter habitat by juveniles will be critical in determining future survival, timing of migration and breeding success.     

Breeding latitude predicts timing but not rate of spring migration in a widespread migratory bird in South America

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Migration of the Little Ringed Plover Charadrius dubius breeding in South Sweden tracked by geolocators

Capsule Little Ringed Plovers breeding in South Sweden migrate towards the southeast in the autumn, via the Middle East, to winter in Saharan and sub-Saharan locations or in India, while the spring migration is more directly towards the north.

Aims To study the migration routes and wintering area of Little Ringed Plovers (Charadrius dubius) breeding in South Sweden, and to investigate the migration strategy and speed for this little studied shorebird.

Methods We use light-level geolocators to track the year-round movements of Little Ringed Plovers breeding in South Sweden.

Results Autumn migration proceeded towards the southeast, in three birds via lengthy stopovers in the Middle East, followed by movements towards the west and southwest to final winter destinations in Africa, while one male made a long stopover in northwestern Iran before migrating to India. The birds wintering in Africa probably stayed at freshwater locations in the Sahara or just south or north of the Sahara. Spring migration was more directly back to the breeding area. Overall migration speeds were similar during autumn and spring migration at about 189 and 209?km/day, respectively. The migration was carried out mainly as many short flights between stopovers. In particular, autumn migration was longer than the direct distance between breeding and wintering sites.

Conclusions This study shows that the geolocator method can successfully be used with relatively small (40?g) shorebirds. We found that a local population of Little Ringed Plover may have widely differing wintering sites (low connectivity), from sub-Saharan Africa to the Indian subcontinent. The migration strategy of the Little Ringed Plover, with multiple short flights, deviates from that of many other long-distance migrating shorebirds that, instead, make one or a few long flights.     

Not as the crow flies: a historical explanation for circuitous migration in Swainson's thrush (Catharus ustulatus)

Many migratory songbirds follow circuitous migratory routes instead of taking the shortest path between overwintering and breeding areas. Here, we study the migration patterns in Swainson's thrush (Catharus ustulatus), a neartic-neotropical migrant songbird, using molecular genetic approaches. This species is presently separated into genetically distinct coastal and continental populations that diverged during the Late Pleistocene (as indicated by molecular dating), yet appear to have retained ancestral patterns of migration. Low nucleotide diversity, a star-like haplotype phylogeny and unimodal mismatch distributions all support the hypothesis that both the coastal and the continental populations have undergone recent demographic expansions. Nearctic-neotropical banding and genetic data show nearly complete segregation of migratory routes and of overwintering locations: coastal populations migrate along the Pacific Coast to overwintering sites in Central America and Mexico, whereas continental populations migrate along an eastern route to overwintering sites in Panama and South America. Nearctic-neotropical banding data also show that continental birds north, northwest and east of this migratory divide fly thousands of miles east before turning south. We conclude that circuitous migration in the Swainson's thrush is an artefact of a Late Pleistocene range expansion.     

Migration routes and stopover sites of Black-necked Cranes determined by satellite tracking

ABSTRACT.   Because their breeding and wintering areas are in remote locations, little is known about the biology of Black-necked Cranes ( Grus nigricollis ), including their migratory behavior. Using satellite telemetry, we monitored the migration of Black-necked Cranes ( N = 6) in China to determine migration routes and the location of stopover sites. From 2005 to 2007, four cranes were tracked during two spring migrations and one fall migration, one was tracked during one spring and one fall migration, and one was tracked during one spring migration. On average, the cranes made seven flights over a 5-d period to migrate 651 km to breeding areas in the spring. In the fall, birds averaged six flights in 5 d to migrate 694 km. The routes traveled by cranes during spring and autumn migration were similar. Both the migration distances and duration of migration are the shortest reported for any crane species to date. Most stopover sites were in areas along rivers and close to wetlands in the Daliang Mountains and the Ruoergai Plateau. Conservation measures are needed to reduce habitat loss (wetland and pasture) in the Daliang Mountains and establish a reserve for stopover sites in the Ruoergai marshes, such as Longriba and Bai River in Hongyuan County.     

Evolution of chain migration in an aerial insectivorous bird,the common swift Apus apus

Spectacular long-distance migration has evolved repeatedly in animals enabling exploration of resources separated in time and space. In birds, these patterns are largely driven by seasonality, cost of migration, and asymmetries in competition leading most often to leapfrog migration, where northern breeding populations winter furthest to the south. Here, we show that the highly aerial common swift Apus apus, spending the nonbreeding period on the wing, instead exhibits a rarely found chain migration pattern, where the most southern breeding populations in Europe migrate to wintering areas furthest to the south in Africa, whereas the northern populations winter to the north. The swifts concentrated in three major areas in sub-Saharan Africa during the nonbreeding period, with substantial overlap of nearby breeding populations. We found that the southern breeding swifts were larger, raised more young, and arrived to the wintering areas with higher seasonal variation in greenness (Normalized Difference Vegetation Index) earlier than the northern breeding swifts. This unusual chain migration pattern in common swifts is largely driven by differential annual timing and we suggest it evolves by prior occupancy and dominance by size in the breeding quarters and by prior occupancy combined with diffuse competition in the winter.     

Moult Strategies Affect Age Differences in Autumn Migration Timing in East Mediterranean Migratory Passerines

Adult passerines renew their flight feathers at least once every year. This complete moult occurs either in the breeding areas, just after breeding (summer moult), or, in some long-distance migratory species, at the non-breeding areas, after arrival to the southern wintering area at the end of autumn migration (winter moult). The aim of this study was to relate moult strategies with the DMD, the difference in median migration date, through Israel, between juveniles and adults. Our data on autumn migration timing in juveniles and adults was based on ringing data of 49,125 individuals belonging to 23 passerine species that breed in Europe and Western Asia and migrate through Israel. We found that DMD was associated with moult timing. In all species that perform a winter moult, adults preceded juveniles during autumn. Among migrants who perform a summer moult, we found evidence of both migration timing patterns: juveniles preceding adults or adults preceding juveniles. In addition, in summer moulters, we found a significant, positive correlation between mean breeding latitude and DMD. Although previous studies described that moult duration and extent can be affected by migration, we suggest that moult strategies affect both migration timing and migration strategy. These two moult strategies (summer or winter moult) also represent two unique migration strategies. Our findings highlight the evolutionary interplay between moult and migration strategies.     

Population-scale drivers of individual arrival times in migratory birds

1. In migratory species, early arrival on the breeding grounds can often enhance breeding success. Timing of spring migration is therefore a key process that is likely to be influenced both by factors specific to individuals, such as the quality of winter and breeding locations and the distance between them, and by annual variation in weather conditions before and during migration. 2. The Icelandic black-tailed godwit Limosa limosa islandica population is currently increasing and, throughout Iceland, is expanding into poorer quality breeding areas. Using a unique data set of arrival times in Iceland in different years for individuals of known breeding and wintering locations, we show that individuals breeding in lower quality, recently occupied and colder areas arrive later than those from traditionally occupied areas. The population is also expanding into new wintering areas, and males from traditionally occupied winter sites also arrive earlier than those occupying novel sites. 3. Annual variation in timing of migration of individuals is influenced by large-scale weather systems (the North Atlantic Oscillation), but between-individual variation is a stronger predictor of arrival time than the NAO. Distance between winter and breeding sites does not influence arrival times. 4. Annual variation in timing of migration is therefore influenced by climatic factors, but the pattern of individual arrival is primarily related to breeding and winter habitat quality. These habitat effects on arrival patterns are likely to operate through variation in individual condition and local-scale density-dependent processes. Timing of migration thus appears to be a key component of the intricate relationship between wintering and breeding grounds in this migratory system.     

Borrelia garinii and Francisella tularensis subsp. holarctica detected in migratory shorebirds in Portugal

Migratory shorebirds use, among many, the East Atlantic Flyway that links breeding areas as north as Tundra habitats to aquatic wintering grounds in West Africa. As a consequence, they are potentially important in the spread of global zoonotic diseases transmitted by ticks, such as Lyme borreliosis and tularemia—two diseases previously detected in Portugal. In this study, we looked at the infection status of seven populations of shorebirds during their migration, breeding, or wintering in the Portuguese wetlands to access if they carry these pathogens and to discuss their potential risk in the Portuguese wetlands. A total of 212 migratory shorebirds captured in the Tagus and Sado estuaries; key staging and wintering sites in this flyway and important breeding areas for some species were analyzed for the presence of Borrelia burgdorferi sensu lato and Francisella tularensis. In the present study, B. garinii was identified in seven (3%) specimens (five black-tailed godwits Limosa limosa, one common redshank Tringa totanus, and one little stint Calidris minuta), whereas F. tularensis subsp. holarctica was identified in one (0.4%) little stint. To our knowledge, this is the first evidence that shorebirds that migrate through or winter in Portugal transport these pathogens, potentially contributing for their introduction along the flyway, including the Mediterranean region.     

Ideal free distributions, evolutionary games, and population dynamics in multiple-species environments

In this article, we develop population game theory, a theory that combines the dynamics of animal behavior with population dynamics. In particular, we study interaction and distribution of two species in a two-patch environment assuming that individuals behave adaptively (i.e., they maximize Darwinian fitness). Either the two species are competing for resources or they are in a predator-prey relationship. Using some recent advances in evolutionary game theory, we extend the classical ideal free distribution (IFD) concept for single species to two interacting species. We study population dynamical consequences of two-species IFD by comparing two systems: one where individuals cannot migrate between habitats and one where migration is possible. For single species, predator-prey interactions, and competing species, we show that these two types of behavior lead to the same population equilibria and corresponding species spatial distributions, provided interspecific competition is patch independent. However, if differences between patches are such that competition is patch dependent, then our predictions strongly depend on whether animals can migrate or not. In particular, we show that when species are settled at their equilibrium population densities in both habitats in the environment where migration between habitats is blocked, then the corresponding species spatial distribution need not be an IFD. Thus, when species are given the opportunity to migrate, they will redistribute to reach an IFD (e.g., under which the two species can completely segregate), and this redistribution will also influence species population equilibrial densities. Alternatively, we also show that when two species are distributed according to the IFD, the corresponding population equilibrium can be unstable.     

Isotopic (δ2Hf) evidence of “loop migration” and use of the Gulf of Maine Flyway by both western and eastern breeding populations of Blackpoll Warblers

Declining numbers of Blackpoll Warblers (Setophaga striata) have been documented at long‐term migration monitoring sites as well as in breeding areas. However, the “loop migration” of Blackpoll Warblers makes it difficult to ascribe population change at migration monitoring sites to specific breeding populations. Individuals from all populations across the breeding range of Blackpoll Warblers concentrate in fall along the Atlantic coastline of eastern North America prior to initiating a transoceanic flight to wintering areas. In spring, Blackpoll Warblers return along a different route, moving north into the southeastern United States where birds from eastern and western breeding populations then diverge during migration to reach their respective breeding areas. To monitor breeding populations outside of breeding areas and identify factors potentially affecting those populations, we must be able to identify where birds captured during migration breed and map seasonal variation in population‐specific flyways. To “map” population‐specific migration movements of Blackpoll Warblers, we used feather deuterium (δ²H_f) values and a spatially explicit model to assign molt origins of 289 Blackpoll Warblers moving through sites in the Gulf of Maine (GOM) region and at three locations further west and south (northern Great Lakes area, Pennsylvania, and Florida). The assignment method was validated with feather samples from 35 birds captured during the breeding season at Churchill, Manitoba, Canada. As predicted, the spatial pattern of movement within and between seasons reflected “loop migration.” Blackpoll Warblers captured during fall migration in the GOM region included birds from across their breeding range, whereas birds captured during the spring were exclusively from northeastern populations. During fall migration, Blackpoll Warblers captured at two sites west of the GOM were from breeding areas further northwest than those from western Canada that were captured in the GOM. Blackpoll Warblers captured in eastern Florida during spring migration were assigned exclusively to breeding areas in the northeast, suggesting that eastern and western populations diverge soon after entering the United States. Finally, most Blackpoll Warblers sampled at Manomet Bird Observatory originated from breeding populations in Alaska and western Canada that have shown a similar (70–90%) decline over the same period. Our results, therefore, not only document the “loop migration” pattern of Blackpoll Warblers, but, by mapping patterns of connectivity between breeding and non‐breeding areas, may help target conservation efforts for breeding populations of Blackpoll Warblers where most needed.