Spring phenology of ecological productivity contributes to the use of looped migration strategies by birds

Migration is a common strategy used by birds that breed in seasonal environments. The patterns and determinants of migration routes, however, remain poorly understood. Recent empirical analyses have demonstrated that the locations of two North America migration flyways (eastern and western) shift seasonally, reflecting the influence of looped migration strategies. For the eastern but not western flyway, seasonal variation in atmospheric circulation has been identified as an explanation. Here, we test an alternative explanation based on the phenology of ecological productivity, which may be of greater relevance in western North America, where phenology is more broadly dictated by elevation. Migrants in the western flyway selected lower-elevation spring routes that were wetter, greener and more productive, and higher-elevation autumn routes that were less green and less productive, but probably more direct. Migrants in the eastern flyway showed little season variation but maintained associations with maximum regional greenness. Our findings suggest the annual phenology of ecological productivity is associated with en route timing in both flyways, and the spring phenology of ecological productivity contributes to the use of looped strategies in the western flyway. This fine-tuned spatial synchronization may be disrupted when changing climate induces a mismatch between food availability and needs.     

Around the Mediterranean: an extreme example of loop migration in a long‐distance migratory passerine

An important issue in migration research is how small‐bodied passerines pass over vast geographical barriers; in European–African avian migration, these are represented by the Mediterranean Sea and the Sahara Desert. Eastern (passing eastern Mediterranean), central (passing Apennine Peninsula) and western (via western Mediterranean) major migration flyways are distinguished for European migratory birds. The autumn and spring migration routes may differ (loop migration) and there could be a certain level of individual flexibility in how individuals navigate themselves during a single migration cycle. We used light‐level loggers to map migration routes of barn swallows Hirundo rustica breeding in the centre of a wide putative contact zone between the northeastern and southernwestern European populations that differ in migration flyways utilised and wintering grounds. Our data documented high variation in migration patterns and wintering sites of tracked birds (n = 19 individuals) from a single breeding colony, with evidence for loop migration in all but one of the tracked swallows. In general, two migratory strategies were distinguished. In the first, birds wintering in a belt stretching from southcentral to southern Africa that used an eastern route for both the spring and autumn migration, then shifted their spring migration eastwards (anti‐clockwise loops, n = 12). In the second, birds used an eastern or central route to their wintering grounds in central Africa, shifting the spring migration route westward (clockwise loops, n = 7). In addition, we observed an extremely wide clockwise loop migration encompassing the entire Mediterranean, with one individual utilising both the eastern (autumn) and western (spring) migratory flyway during a single annual migration cycle. Further investigation is needed to ascertain whether clockwise migratory loops encircling the entire Mediterranean also occur other small long‐distance passerine species.     

Migration of red‐backed shrikes from the Iberian Peninsula: optimal or sub‐optimal detour?

The current Northern Hemisphere migration systems are believed to have arisen since the last glaciation. In many cases, birds do not migrate strait from breeding to non‐breeding areas but fly via a detour. All western European populations of red‐backed shrikes Lanius collurio are assumed to reach their southern African wintering grounds detouring via southeast Europe. Based on theoretical considerations under an optimality framework this detour is apparently optimal. Here, we use individual geolocator data on red‐backed shrikes breeding in Spain to show that these birds do indeed detour via southeast Europe en route to southern Africa where they join other European populations of red‐backed shrikes and return via a similar route in spring. Disregarding potential wind assistance, the routes taken for the tracked birds in autumn were not optimal compared to crossing the barrier directly. For spring migration the situation was quite different with the detour apparently being optimal. However, when considering potential wind assistance estimated total air distances during autumn migration were overall similar and the barrier crossing shorter along the observed routes. We conclude that considering the potential benefit of wind assistance makes the route via southeast Europe likely to be less risky in autumn. However, it cannot be ruled out that other factors, such as following a historical colonisation route could still be important.     

Regional wind patterns likely shape a seasonal migration detour

Migrating animals should optimise time and energy use when migrating, travelling directly to their destination. Detours from the most direct route may arise however because of barriers and weather conditions. Identifying how such situations arise from variable weather conditions is crucial to understand population response in the light of increased anthropogenic climate change. Here we used light-level geolocators to follow Cyprus wheatears for their full annual cycle in two separate years migrating between Cyprus, over the Mediterranean and the Sahara to winter in north–east sub-Saharan Africa. We predicted that any route detours would be related to wind conditions experienced during migration. We found that spring migration for all birds included an eastern detour, whilst autumn migrations were direct across the Sahara. The direct autumn migration was likely a consequence of consistent tail-winds, whilst the eastern detour in spring is likely to be more efficient given the wind conditions which are against a direct route. Such variable migration routes shaped by coincidence with prevailing winds are probably common suggesting that some birds may be able to adapt to future changes in wind conditions.     

Rapid long‐distance migration in Norwegian Lesser Black‐backed Gulls Larus fuscus fuscus along their eastern flyway

We studied the long‐distance migration of Lesser Black‐backed Gulls Larus fuscus fuscus breeding in northern Norway along their eastern flyway using geolocators in 2009 and 2010. The majority of birds wintered in lakes in East Africa and the southeast Mediterranean was the most important stopover area. Larus f. fuscus along the eastern flyway travelled at a net travel speed of 399 and 177 km/day during the autumn and spring migration, respectively, higher than published travel speeds for Dutch Larus fuscus migrating along the western flyway. The results suggest that the long‐distance migratory Norwegian L. f. fuscus seek to minimize time spent in transit, whereas lower travel speed during northerly spring migration may reflect differences in wind patterns or food conditions between spring and autumn.     

Loop migration in adult marsh harriers Circus aeruginosus,as revealed by satellite telemetry

Loop migration among birds is characterized by the spring route lying consistently west or east of the autumn route. The existence of loops has been explained by general wind conditions or seasonal differences in habitat distribution. Loop migration has predominantly been studied at the population level, for example by analysing ring recoveries. Here we study loop migration of individual marsh harriers Circus aeruginosus tracked by satellite telemetry. We show that despite a generally narrow migration corridor the harriers travelled in a distinct clockwise loop through Africa and southern Europe, following more westerly routes in spring than in autumn. We used the Normalized Difference Vegetation Index (NDVI) to identify potential feeding habitat in Africa. Suitable habitat seemed always more abundant along the western route, both in spring and autumn, and no important stopover site was found along the eastern route. Observed routes did thus not coincide with seasonal variation in habitat availability. However, favourable habitat might be more important during spring migration, when the crossing of the Sahara seems more challenging, and thus habitat availability might play an indirect role in the harriers’ route choice. Grid‐based wind data were used to reconstruct general wind patterns, and in qualitative agreement with the observed loop marsh harriers predominantly encountered westerly winds in Europe and easterly winds in Africa, both in autumn and in spring. By correlating tail‐ and crosswinds with forward and perpendicular movement rates, respectively, we show that marsh harriers are partially drifted by wind. Thus, we tentatively conclude that wind rather than habitat seems to have an overriding effect on the shape of the migration routes of marsh harriers. General wind conditions seem to play an important role also in the evolution of narrow migratory loops as demonstrated for individual marsh harriers.     

Comparing areas of suitable habitats along travelled and possible shortest routes in migration of White-naped Cranes Grus vipio in East Asia

Many migratory species take detours when migrating from their breeding to wintering grounds rather than following the shortest route available. To test whether the distribution of potentially suitable habitats might be a factor causing the use of less direct detours during migration, we analysed the migratory routes of five White-naped Cranes Grus vipio satellite-tracked from central–east Russia, and compared the total area of wetlands and grasslands along the migratory routes travelled by the Cranes with that along the shortest possible routes to the Cranes' wintering grounds. All five Cranes made an easterly detour, and the distance ratio of the routes used by Cranes to the shortest possible route was 1.13 ± 0.03 sd. Based on National Oceanic and Atmospheric Administration satellite images, we demonstrate that the area of wetlands and grasslands along the migration routes travelled by Cranes was greater than along the shortest possible routes.     

Seasonal variation in migration strategies used to cross ecological barriers in a nearctic migrant wintering in Africa

Ecological barriers such as oceans, mountain ranges or glaciers can have a substantial influence on the evolution of animal migration. Along the migration flyway connecting breeding sites in the North American Arctic and wintering grounds in Europe or Africa, nearctic species are confronted with significant barriers such as the Atlantic Ocean and the Greenland icecap. Using geolocation devices, we identified wintering areas used by ringed plovers nesting in the Canadian High‐Arctic and investigated migration strategies used by these nearctic migrants along the transatlantic route. The main wintering area of the ringed plovers (n = 20) was located in western Africa. We found contrasting seasonal migration patterns, with ringed plovers minimizing continuous flight distances over the ocean in spring by making a detour to stop in Iceland. In autumn, however, most individuals crossed the ocean in one direct flight from southern Greenland to western Europe, as far as southern Spain. This likely resulted from prevailing anti‐clockwise winds associated with the Icelandic low‐pressure system. Moreover, the plovers we tracked largely circumvented the Greenland icecap in autumn, but in spring, some plovers apparently crossed the icecap above the 65°N. Our study highlighted the importance of Iceland as a stepping‐stone during the spring migration and showed that small nearctic migrants can perform non‐stop transatlantic flights from Greenland to southern Europe.     

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.     

The role of wind in passerine autumn migration between Europe and Africa

Large ecological barriers such as oceans and deserts have considerablyshaped the migratory strategies of birds. The ecological barriersposed by the Alps, the Mediterranean Sea, and the Sahara seemto prevent most long-distance migrants from flying on a directsouthward course from Europe to Africa. Migratory routes towardsouthwest and southeast prevail. These two flyways differ withrespect to topography, refueling possibilities, and wind conditions.Aiming at a better understanding of the evolution of both flywaysin spite of differing conditions, we studied potential survivalof passerine birds on their first autumn migration from northernEurope to tropical Africa by means of a computer simulation.Considering real wind conditions at 850 mb (approximately 1500m above sea level), the survival rates of birds with southeasterly(SE) migratory directions were much higher than those of birdswith southwesterly (SW) directions. With the possibility tochoose the altitude (from four levels) with the most favorablewind, both SE and SW migrants had similar high survival, butonly with refueling opportunities in northwest (NW) Africa forSW migrants. Our results suggest that the southwestern flywaydepends on the selection of days, but especially altitudes,with favorable wind conditions and on refueling opportunitiesin NW Africa. The SE flyway is privileged by the frequent favorablewind conditions for crossing the eastern Mediterranean Sea andthe Egyptian desert, where refueling sites are almost absent.Both autumn migration routes would be unlikely without windassistance.     

The pull of the Central Flyway? Veeries breeding in western Canada migrate using an ancestral eastern route

Understanding non‐breeding season movements and identifying wintering areas of different populations of migratory birds is important for establishing patterns of migratory connectivity over the annual cycle. We analyzed archival solar geolocation (N = 5) and global positioning data (N = 1) to investigate migration routes, stopover sites, and wintering areas of a western‐most breeding population of Veeries (Catharus fuscescens) in the Pemberton Valley, British Columbia, Canada. Geolocation data were analyzed using a Bayesian state‐space model to improve likely position estimates. We compared our results with those from a Veery population located ~250 km east across a mountain chain in the Okanagan Valley, British Columbia, and with an eastern population in Delaware, U.S.A. Migrating Veeries from the Pemberton Valley used an eastern trajectory through the Rocky Mountains to the Great Plains to join a central flyway during fall and spring migration, a route similar to that used by Veeries breeding in the Okanagan Valley. However, wintering destinations of Pemberton Valley birds were more varied, with inter‐individual wintering distances ~1000 km greater than birds from the Okanagan Valley population and ~500 km from the previously known winter range of Veeries. The observed eastern migration path likely follows an ancestral route that evolved following the most recent glacial retreat. Consistent with patterns observed from the Okanagan and Delaware populations, Veeries from the Pemberton Valley undertook an intra‐tropical migration on the wintering grounds, but this winter movement differed from those of previously studied populations. Such winter movements may thus be idiosyncratic or show coarse population associations. Intra‐wintering‐ground movements likely occur either in response to seasonal changes in habitat suitability or as a means of optimizing pre‐migratory fueling prior to long‐distance spring movements to North America.     

Tracking the Stejneger's stonechat Saxicola stejnegeri along the East Asian–Australian Flyway from Japan via China to southeast Asia

The East Asian–Australian Flyway spans from north Asia to Australia and is the world's richest birds' flyway because it involves > 40% of global migratory bird species. However, information is lacking on individual migratory routes and non‐breeding grounds for small land birds using this flyway. Here, we present the first migration tracks of the songbird Stejneger's stonechat Saxicola stejnegeri from this part of the world using light‐level geolocators. This species depends on grasslands during the entire annual cycle and was captured and equipped with tracking devices in Hokkaido, northern Japan. All individuals traveled through southern Primorye or eastern Heilongjiang (Russia/China) before flying southward via central China toward their major non‐breeding grounds in southeast Asia (China, Laos, Cambodia, Thailand, and Vietnam). Individual stonechats spent 42–70 d en route during their autumn migration. Both the major non‐breeding grounds and the stopover sites are likely to pose challenges to the persistence of this species, because these habitats are currently degraded and will likely be lost in the near future due to intensified agriculture and the establishment of permanent croplands. Moreover, the areas used by Stejneger's stonechat during migration largely overlapped with illegal trapping areas in northeastern China.     

High juvenile mortality during migration in a declining population of a long‐distance migratory raptor

Many populations of long‐distance migrants are declining and there is increasing evidence that declines may be caused by factors operating outside the breeding season. Among the four vulture species breeding in the western Palaearctic, the species showing the steepest population decline, the Egyptian Vulture Neophron percnopterus, is a long‐distance migrant wintering in Africa. However, the flyways and wintering areas of the species are only known for some populations, and without knowledge of where mortality occurs, effective conservation management is not possible. We tracked 19 juvenile Egyptian Vultures from the declining breeding population on the Balkan Peninsula between 2010 and 2014 to estimate survival and identify important migratory routes and wintering areas for this species. Mortality during the first autumn migration was high (monthly survival probability 0.75) but mortality during migration was exclusively associated with suboptimal navigation. All birds from western breeding areas and three birds from central and eastern breeding areas attempted to fly south over the Mediterranean Sea, but only one in 10 birds survived this route, probably due to stronger tailwind. All eight birds using the migratory route via Turkey and the Middle East successfully completed their first autumn migration. Of 14 individual and environmental variables examined to explain why juvenile birds did or did not successfully complete their first migration, the natal origin of the bird was the most influential. We speculate that in a declining population with fewer experienced adults, an increasing proportion of juvenile birds are forced to migrate without conspecific guidance, leading to high mortality as a consequence of following sub‐optimal migratory routes. Juvenile Egyptian Vultures wintered across a vast range of the Sahel and eastern Africa, and had large movement ranges with core use areas at intermediate elevations in savannah, cropland or desert. Two birds were shot in Africa, where several significant threats exist for vultures at continental scales. Given the broad distribution of the birds and threats, effective conservation in Africa will be challenging and will require long‐term investment. We recommend that in the short term, more efficient conservation could target narrow migration corridors in southern Turkey and the Middle East, and known congregation sites in African wintering areas.     

Migration patterns of the Osprey Pandion haliaetus on the Eastern European–East African flyway

We analysed migration strategies of the Osprey Pandion haliaetus on the poorly studied Eastern European–East African flyway. Four adult birds were equipped with GPS-based satellite-transmitters or data-loggers in their breeding sites in Estonia (north-eastern Europe) and tracked to their wintering grounds in Africa and back, during up to six migration cycles. Departure times, migration routes, as well as wintering and stopover sites varied remarkably between individuals but not much between years. Stopovers (2–30 days) were made mostly in Europe and less in the Middle East (Turkey) and north-eastern Africa (Egypt). The Ospreys did not avoid flying long distances over the sea, and the sea was crossed four times during the night. The current study adds to current knowledge on Osprey migration and should help to concentrate actions on protecting important flyways and stopover locations.     

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.     

The annual cycle of a trans-equatorial Eurasian-African passerine migrant: different spatio-temporal strategies for autumn and spring migration

The small size of the billions of migrating songbirds commuting between temperate breeding sites and the tropics has long prevented the study of the largest part of their annual cycle outside the breeding grounds. Using light-level loggers (geolocators), we recorded the entire annual migratory cycle of the red-backed shrike Lanius collurio, a trans-equatorial Eurasian-African passerine migrant. We tested differences between autumn and spring migration for nine individuals. Duration of migration between breeding and winter sites was significantly longer in autumn (average 96 days) when compared with spring (63 days). This difference was explained by much longer staging periods during autumn (71 days) than spring (9 days). Between staging periods, the birds travelled faster during autumn (356 km d(-1)) than during spring (233 km d(-1)). All birds made a protracted stop (53 days) in Sahelian sub-Sahara on southbound migration. The birds performed a distinct loop migration (22 000 km) where spring distance, including a detour across the Arabian Peninsula, exceeded the autumn distance by 22 per cent. Geographical scatter between routes was particularly narrow in spring, with navigational convergence towards the crossing point from Africa to the Arabian Peninsula. Temporal variation between individuals was relatively constant, while different individuals tended to be consistently early or late at different departure/arrival occasions during the annual cycle. These results demonstrate the existence of fundamentally different spatio-temporal migration strategies used by the birds during autumn and spring migration, and that songbirds may rely on distinct staging areas for completion of their annual cycle, suggesting more sophisticated endogenous control mechanisms than merely clock-and-compass guidance among terrestrial solitary migrants. After a century with metal-ringing, year-round tracking of long-distance migratory songbirds promises further insights into bird migration.     

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.     

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.     

Population-specific adjustment of the annual cycle in a super-swift trans-Saharan migrant

For migratory birds optimal timing of the onset of reproduction is vital, especially when suitable conditions for reproduction occur only for a short while during the year. With increasing latitude the suitable period becomes shorter and we expect the organization of annual cycle to be more synchronized to the local conditions across individuals of same population. This should result in low variation of arrival and departure date in breeding sites at higher latitudes. We quantify the temporal and geographical variation in pre- and post-breeding migration between individuals from four different populations of alpine swifts Tachymarptis melba along a latitudinal gradient. We tracked 215 individuals in three years with geolocators. The two western and two eastern populations showed separate migratory flyways and places of residence in Africa. Length of stay at the breeding sites was negatively correlated with latitude and differed by more than a month between populations. Duration of migration was similarly short in all populations (median 6.2 days in autumn and 8.7 days in spring). However, variation in timing of migration was unrelated to latitude and individuals everywhere arrived in the same asynchrony at the breeding site.