Where eagles soar: Fine‐resolution tracking reveals the spatiotemporal use of differential soaring modes in a large raptor

Unlike smaller raptors, which can readily use flapping flight, large raptors are mainly restricted to soaring flight due to energetic constraints. Soaring comprises of two main strategies: thermal and orographic soaring. These soaring strategies are driven by discrete uplift sources determined by the underlying topography and meteorological conditions in an area. High‐resolution GPS tracking of raptor flight allows the identification of these flight strategies and interpretation of the spatiotemporal occurrence of thermal and orographic soaring. In this study, we develop methods to identify soaring flight behaviors from high‐resolution GPS tracking data of Verreaux’s eagle Aquila verreauxii and analyze these data to understand the conditions that promote the use of thermal and orographic soaring. We use these findings to predict the use of soaring flight both spatially (across the landscape) and temporally (throughout the year) in two topographically contrasting regions in South Africa. We found that topography is important in determining the occurrence of soaring flight and that thermal soaring occurs in relatively flat areas which are likely to have good thermal uplift availability. The predicted use of orographic soaring was predominately determined by terrain slope. Contrary to our expectations, the topography and meteorology of eagle territories in the Sandveld promoted the use of soaring flight to a greater extent than in territories in the more mountainous Cederberg region. Spatiotemporal mapping of predicted flight behaviors can broaden our understanding of how large raptors like the Verreaux’s eagle use their habitat and how that links to energetics (as the preferential use of areas that maximize net energy gain is expected), reproductive success, and ultimately population dynamics. Understanding the fine‐scale landscape use and environmental drivers of raptor flight can also help to predict and mitigate potential detrimental effects of anthropogenic developments, such as mortality via collision with wind turbines.     

Testing an emerging paradigm in migration ecology shows surprising differences in efficiency between flight modes

To maximize fitness, flying animals should maximize flight speed while minimizing energetic expenditure. Soaring speeds of large-bodied birds are determined by flight routes and tradeoffs between minimizing time and energetic costs. Large raptors migrating in eastern North America predominantly glide between thermals that provide lift or soar along slopes or ridgelines using orographic lift (slope soaring). It is usually assumed that slope soaring is faster than thermal gliding because forward progress is constant compared to interrupted progress when birds pause to regain altitude in thermals. We tested this slope-soaring hypothesis using high-frequency GPS-GSM telemetry devices to track golden eagles during northbound migration. In contrast to expectations, flight speed was slower when slope soaring and eagles also were diverted from their migratory path, incurring possible energetic costs and reducing speed of progress towards a migratory endpoint. When gliding between thermals, eagles stayed on track and fast gliding speeds compensated for lack of progress during thermal soaring. When thermals were not available, eagles minimized migration time, not energy, by choosing energetically expensive slope soaring instead of waiting for thermals to develop. Sites suited to slope soaring include ridges preferred for wind-energy generation, thus avian risk of collision with wind turbines is associated with evolutionary trade-offs required to maximize fitness of time-minimizing migratory raptors.     

Using a Doppler light detection and ranging (lidar) system to characterize an atmospheric thermal providing lift for soaring raptors

ABSTRACT.   Raptors and other large birds in soaring flight take advantage of upward drafts of air called thermals to maintain altitude with minimal flapping. I used a Doppler light detection and ranging (lidar) system to characterize a thermal in which raptors were soaring. Doppler lidar allows imaging of wind fields to reveal the structure of updrafts and downdrafts in a thermal. The thermal I monitored was in the form of a horizontal convective roll created at a transition from clear sky to partly cloudy sky, and gave both lift and lateral motion to the soaring birds. The thermal was 700 m high with a vertical wind speed that peaked at 3 m/s, so raptors could have soared to and maintained that altitude as the horizontal wind moved the thermal. My results suggest that imaging wind fields with Doppler lidar can be a useful tool for studying thermals and how they are used by soaring birds. An effective combination for further study of bird flight interaction with wind phenomena would be to add lidar measurements to an established means of tracking bird flight by radio or GPS transmitters, aircraft tracking, or radar.     

Regional and seasonal flight speeds of soaring migrants and the role of weather conditions at hourly and daily scales

Given that soaring birds travel faster with supportive winds or in good thermal soaring conditions, we expect weather conditions en route of migration to explain commonly observed regional and seasonal patterns in the performance of soaring migrants. We used GPS‐loggers to track 13 honey buzzards and four Montagu's harriers for two to six migrations each. We determined how tailwinds, crosswinds, boundary layer height (a proxy for thermal convection) and precipitation affected hourly speeds, daily distances and daily mean speeds with linear regression models. Honey buzzards mostly travel by soaring while Montagu's harriers supplement soaring with flapping. Therefore, we expect that performance of harriers will be less affected by weather than for buzzards. Weather conditions explained between 30 and 50% of variation in migration performance of both species. Tailwind had the largest effect on hourly speeds, daily mean speeds and daily travel distances. Honey buzzards travelled significantly faster and farther, and Montagu's harriers non‐significantly faster, under better convective conditions. Honey buzzards travelled at slower speeds and shorter distances in crosswinds, whereas harriers maintained high speeds in crosswinds. Weather conditions varied between regions and seasons, and this variation accounted for nearly all regional and seasonal variation in flight performance. Hourly performance was higher than predicted at times when we suspect birds had switched to intermittent or continuous flapping flight, for example during sea‐crossings. The daily travel distance of Montagu's harriers was determined to a significant extent by their daily travel time, which differed between regions, possibly also due to weather conditions. We conclude with the implications of our work for studies on migration phenology and we suggest an important role for high‐resolution telemetry in understanding migratory behavior across entire migratory journeys.     

Flight strategies of migrating raptors; a comparative study of interspecific variation in flight characteristics

The comparison of flight styles and flight parameters of migrating raptors in Israel revealed the following. (1) Climbing rate in thermal circling did not differ between species, indicating that chiefly the strength of thermal updrafts determined the climbing rate and that morphological features were less relevant. (2) In interthermal gliding, air speed was positively and gliding angle negatively related to the species' average body mass. Heavier species glided faster and had smaller gliding angles. (3) In soaring and gliding flight, cross-country speed relative to the air was positively related to the species' body mass; it was obviously the result of the gliding ability increasing with body mass. (4) Eagles and buzzards used soaring and gliding flight for more than 95% of the observation time. Additional soaring in a straight line whilst gliding was extensively used by the Steppe Eagle Aquila nipalensis, Lesser Spotted Eagle Aquila pomarina and Booted Eagle Hieraætus pennatus and even more frequently by the resident species, the Griffon Vulture Gyps fulvus and Shorttoed Eagle Circaetus gallicus. Smaller species, such as the Levant Sparrowhawk Accipiter brevipes, harriers (Circus sp.) and small falcons (Falco sp.). showed the highest proportion of flapping and gliding flight (9–33%). (5) In a comparison of the flight parameters and proportions of flight styles, a cluster analysis distinguished two main groups: The first consisted of Montagu's Harrier Circus pygargus, Pallid Harrier Circus macrourus, Levant Sparrowhawk and small falcons; their flight behaviour was characterized by both the high proportion of flapping and the low gliding performance. The second group comprised the typical soaring migrants: Steppe Eagle, Lesser Spotted Eagle, Booted Eagle, Steppe Buzzard Buteo buteo vulpinus, Honey Buzzard Pernis apivorus and Egyptian Vulture Neophron percnopterus, and they had very similar flight behaviour and were closely clustered. The Black Kite Milvus migrans and Marsh Harrier Circus aeruginosus were intermediate between typical soarers and flappers. The two resident species, Griffon Vulture and Short-toed Eagle, were grouped separately from the soaring migrants.     

Soaring across continents: decision‐making of a soaring migrant under changing atmospheric conditions along an entire flyway

Thermal soaring birds reduce flight‐energy costs by alternatingly gaining altitude in thermals and gliding across the earth's surface. To find out how soaring migrants adjust their flight behaviour to dynamic atmospheric conditions across entire migration routes, we combined optimal soaring migration theory with high‐resolution GPS tracking data of migrating honey buzzards Pernis apivorus and wind data from a global numerical atmospheric model. We compared measurements of gliding air speeds to predictions based on two distinct behavioural benchmarks for thermal soaring flight. The first being a time‐optimal strategy whereby birds alter their gliding air speeds as a function of climb rates to maximize cross‐country air speed over a full climb– glide cycle (V_opt). The second a risk‐averse energy‐efficient strategy at which birds alter their gliding air speed in response to tailwinds/headwinds to maximize the distance travelled in the intended direction during each glide phase (V_bgw). Honey buzzards were gliding on average 2.05 ms^{– 1} slower than V_opt and 3.42 ms^{– 1} faster than V_bgw while they increased air speeds with climb rates and reduced air speeds in tailwinds. They adopted flexible flight strategies gliding mostly near V_bgw under poor soaring conditions and closer to V_opt in good soaring conditions. Honey buzzards most adopted a time‐optimal strategy when crossing the Sahara, and at the onset of spring migration, where and when they met with the best soaring conditions. The buzzards nevertheless glided slower than V_opt during most of their journeys, probably taking time to navigate, orientate and locate suitable thermals, especially in areas with poor thermal convection. Linking novel tracking techniques with optimal migration models clarifies the way birds balance different tradeoffs during migration.     

How Cheap Is Soaring Flight in Raptors? A Preliminary Investigation in Freely-Flying Vultures

Measuring the costs of soaring, gliding and flapping flight in raptors is challenging, but essential for understanding their ecology. Among raptors, vultures are scavengers that have evolved highly efficient soaring-gliding flight techniques to minimize energy costs to find unpredictable food resources. Using electrocardiogram, GPS and accelerometer bio-loggers, we report the heart rate (HR) of captive griffon vultures (Gyps fulvus and G. himalayensis) trained for freely-flying. HR increased three-fold at take-off (characterized by prolonged flapping flight) and landing (>300 beats-per-minute, (bpm)) compared to baseline levels (80–100 bpm). However, within 10 minutes after the initial flapping phase, HR in soaring/gliding flight dropped to values similar to baseline levels, i.e. slightly lower than theoretically expected. However, the extremely rapid decrease in HR was unexpected, when compared with other marine gliders, such as albatrosses. Weather conditions influenced flight performance and HR was noticeably higher during cloudy compared to sunny conditions when prolonged soaring flight is made easier by thermal ascending air currents. Soaring as a cheap locomotory mode is a crucial adaptation for vultures who spend so long on the wing for wide-ranging movements to find food.     

Wing size but not wing shape is related to migratory behavior in a soaring bird

Both wing size and wing shape affect the flight abilities of birds. Intra and inter‐specific studies have revealed a pattern where high aspect ratio and low wing loading favour migratory behaviour. This, however, have not been studied in soaring migrants. We assessed the relationship between the wing size and shape and the characteristics of the migratory habits of the turkey vulture Cathartes aura, an obligate soaring migrant. We compared wing size and shape with migration strategy among three fully migratory, one partially migratory and one non‐migratory (resident) population distributed across the American continent. We calculated the aspect ratio and wing loading using wing tracings to characterize the wing morphology. We used satellite‐tracking data from the migratory populations to calculate distance, duration, speed and altitude during migration. Wing loading, but not aspect ratio, differed among the populations, segregating the resident population from the completely migratory ones. Unlike what has been reported in species using flapping flight during migration, the migratory flight parameters of turkey vultures were not related to the aspect ratio. By contrast, wing loading was related to most flight parameters. Birds with lower wing loading flew farther, faster, and higher during their longer journeys. Our results suggest that wing morphology in this soaring species enables lower‐cost flight, through low wing‐loading, and that differences in the relative sizes of wings may increase extra savings during migration. The possibility that wing shape is influenced by foraging as well as migratory flight is discussed. We conclude that flight efficiency may be improved through different morphological adaptations in birds with different flight mechanisms.     

Estimating flight height and flight speed of breeding Piping Plovers

Collisions with wind turbines are an increasing conservation concern for migratory birds that already face many threats. Existing collision‐risk models take into account parameters of wind turbines and bird flight behavior to estimate collision probability and mortality rates. Two behavioral characteristics these models require are the proportion of birds flying at the height of the rotor swept‐zone and the flight speed of birds passing through the rotor swept‐zone. In recent studies, investigators have measured flight height and flight speed of migrating birds using fixed‐beam radar and thermal imaging. These techniques work well for fixed areas where migrants commonly pass over, but they cannot readily provide species‐specific information. We measured flight heights of a nesting shorebird, the federally threatened Piping Plover (Charadrius melodus), using optical range finding and measured flight speed using videography. Several single‐turbine wind projects have been proposed for the Atlantic coast of the United States where they may pose a potential threat to these plovers. We studied Piping Plovers in New Jersey and Massachusetts during the breeding seasons of 2012 and 2013. Measured flight heights ranged from 0.7 to 10.5 m with a mean of 2.6 m (N = 19). Concurrent visually estimated flight heights were all within 2 m of measured heights and most within 1 m. In separate surveys, average visually estimated flight height was 2.6 m (N = 1674) and ranged from 0.25 m to 40 m. Average calculated flight speed was 9.30 m/s (N = 17). Optical range finding was challenging, but provided a useful way to calibrate visual estimates where frames of reference were lacking in the environment. Our techniques provide comparatively inexpensive, replicable procedures for estimating turbine collision‐risk parameters where the focus is on discrete nesting areas of specific species where birds follow predictable flight paths.     

Flight mode affects allometry of migration range in birds

Billions of birds migrate to exploit seasonally available resources. The ranges of migration vary greatly among species, but the underlying mechanisms are poorly understood. I hypothesise that flight mode (flapping or soaring) and body mass affect migration range through their influence on flight energetics. Here, I compiled the tracks of migratory birds (196 species, weighing 12–10 350 g) recorded by electronic tags in the last few decades. In flapping birds, migration ranges decreased with body mass, as predicted from rapidly increasing flight cost with increasing body mass. The species with higher aspect ratio and lower wing loading had larger migration ranges. In soaring birds, migration ranges were mass‐independent and larger than those of flapping birds, reflecting their low flight costs irrespective of body mass. This study demonstrates that many animal‐tracking studies are now available to explore the general patterns and the underlying mechanisms of animal migration.     

Meteorological and environmental variables affect flight behaviour and decision‐making of an obligate soaring bird,the California Condor Gymnogyps californianus

The movements of animals are limited by evolutionary constraints and ecological processes and are strongly influenced by the medium through which they travel. For flying animals, variation in atmospheric conditions is critically influential in movement. Obligate soaring birds depend on external sources of updraft more than do other flying species, as without that updraft they are unable to sustain flight for extended periods. These species are therefore good models for understanding how the environment can influence decisions about movement. We used meteorological and topographic variables to understand the environmental influences on the decision to engage in flight by obligate soaring and critically endangered California Condors Gymnogyps californianus. Condors were more likely to fly, soared at higher altitudes and flew over smoother terrain when weather conditions promoted either thermal or orographic updrafts, for example when turbulence and solar radiation were higher and when winds from the east and north were stronger. However, increased atmospheric stability, which is inconsistent with thermal development but may be associated with orographic updrafts, was correlated with a somewhat higher probability of being in flight at lower altitudes and over rougher terrain. The close and previously undescribed linkages between Condor flight and conditions that support development of thermal and orographic updrafts provide important insight into the behaviour of obligate soaring birds and into the environmental parameters that may define the currently expanding distribution of Condors within and outside the state of California.     

OBSERVATIONS OF THE MIGRATION OF RAPTORS AND OTHER LARGE SOARING BIRDS IN BULGARIA, 1975–1978

Observations are given, from the years 1975 to 1978, on the migrations of raptors and large soaring birds along the Bulgarian section of the Black Sea coast, with casual observations on passage in western and central Bulgaria. The winter status of pelicans, and of some raptors, is discussed.
The author's findings suggest that the main spring migration route follows the coast north from Burgas, rather than turning west there, as had been suggested.     

Directed flight and optimal airspeeds: homeward‐bound gulls react flexibly to wind yet fly slower than predicted

Birds in flight are proposed to adjust their body orientation (heading) and airspeed to wind conditions adaptively according to time and energy constraints. Airspeeds in goal‐directed flight are predicted to approach or exceed maximum‐range airspeeds, which minimize transport costs (energy expenditure per unit distance) and should increase in headwinds and crosswinds. Diagnosis of airspeed adjustment is however obscured by uncertainty regarding birds' goal‐directions, transport costs, interrelations with orientation strategy and the attainability of predicted behaviour. To address these issues, we tested whether gulls minimized transport costs through adjustment of airspeed and heading to wind conditions during extended inbound flight over water (180–360 km) to their breeding colony, and introduce a methodology to assess transport (energy) efficiency given wind conditions. Airspeeds, heading, flight mode and energy expenditure were estimated using GPS tracking, accelerometer and wind data. Predicted flight was determined by simulating each trip according to maximum‐range airspeeds and various orientation strategies. Gulls employed primarily flapping flight (93%), and negotiated crosswinds flexibly to exploit both high altitude tailwinds and coastal soaring opportunities. We demonstrate that predicted airspeeds in heavy crosswinds depend strongly on orientation strategy and presumed preferred direction. Measured airspeeds increased with headwind and crosswind similarly to maximum‐range airspeeds based on full compensation for wind drift, yet remained ～ 30% lower than predicted by all strategies, resulting in slower and 30–35% costlier flight. Interestingly, more energy could be saved through adjustment of airspeed (median 40%) than via orientation strategy (median 4%). Therefore, despite remarkably flexible reaction to wind at sea, these gulls evidently minimized neither time nor energy expenditure. However, airspeeds were possibly over‐predicted by current aerodynamic models. This study emphasizes the importance of accounting for orientation strategy when assessing airspeed adjustments to wind and indicates that either the cost or adaptive ‘currency’ of extended flight among gulls may require revision.     

Cross sectional geometry of the forelimb skeleton and flight mode in pelecaniform birds

Avian wing elements have been shown to experience both dorsoventral bending and torsional loads during flapping flight. However, not all birds use continuous flapping as a primary flight strategy. The pelecaniforms exhibit extraordinary diversity in flight mode, utilizing flapping, flap‐gliding, and soaring. Here we (1) characterize the cross‐sectional geometry of the three main wing bone (humerus, ulna, carpometacarpus), (2) use elements of beam theory to estimate resistance to loading, and (3) examine patterns of variation in hypothesized loading resistance relative to flight and diving mode in 16 species of pelecaniform birds. Patterns emerge that are common to all species, as well as some characteristics that are flight‐ and diving‐mode specific. In all birds examined, the distal most wing segment (carpometacarpus) is the most elliptical (relatively high I_max/I_min) at mid‐shaft, suggesting a shape optimized to resist bending loads in a dorsoventral direction. As primary flight feathers attach at an oblique angle relative to the long axis of the carpometacarpus, they are likely responsible for inducing bending of this element during flight. Moreover, among flight modes examined the flapping group (cormorants) exhibits more elliptical humeri and carpometacarpi than other flight modes, perhaps pertaining to the higher frequency of bending loads in these elements. The soaring birds (pelicans and gannets) exhibit wing elements with near‐circular cross‐sections and higher polar moments of area than in the flap and flap‐gliding birds, suggesting shapes optimized to offer increased resistance to torsional loads. This analysis of cross‐sectional geometry has enhanced our interpretation of how the wing elements are being loaded and ultimately how they are being used during normal activities. J. Morphol., 2011. © 2011 Wiley‐Liss,Inc.     

Narrow sea crossings present major obstacles to migrating Griffon Vultures Gyps fulvus

The flight behaviour of Griffon Vultures Gyps fulvus was studied at a major migration bottleneck, the Strait of Gibraltar in southernmost Spain, during the autumns of 2004 to 2007. The 14‐km‐wide sea channel significantly impeded the southern migration of the species into Africa, with many birds attempting repeated passage for weeks before crossing, and others not crossing at all and overwintering in Southern Spain. Water‐crossing attempts were restricted to times between 11:00 and 14:00 h on days with light or variable winds, or on days with strong winds from the north or west. No crossing attempts were made on days with strong winds from the south or east. Vultures attempted to cross the Strait in large flocks and never attempted to do so alone. Although 29% of the birds soared during crossing attempts, at least until they flew beyond visible range of approximately 4 km, most engaged in considerable flapping flight when attempting to cross. Overall, birds flying over water flapped more than 10 times as frequently as those flying over land prior to crossing attempts. Vultures did not flap continuously, but intermittently in brief bouts of flapping interspersed with periods of gliding or soaring flight. The number of flaps per bout over water was significantly greater than the number of flaps per bout over land. Vultures flying over water that flapped at rates of 20 flaps or more per minute typically aborted attempted crossings and returned to Spain in intermittent flapping and gliding flight. There are numerous reports of Vultures falling into the Strait and drowning while attempting to cross, as well as reports of returning Vultures collapsing on the beach having reached Spain in spring ( Barrios Partida 2006 ). Our observations indicate that passage of Griffon Vultures at the Strait of Gibraltar is limited by the species’ over‐water flapping‐flight abilities, including its inability to flap continuously for even short periods of time. We suggest that even relatively short sea crossings represent significant obstacles to migrating Vultures and discuss the implications of this limitation on the distribution and abundance of the species.     

Extremely detoured migration in an inexperienced bird: interplay of transport costs and social interactions

We tagged two juvenile short‐toed eagles in southern Italian peninsula with GPS satellite transmitters. According to previous visual observations, two different migratory routes for Italian short‐toed eagles to reach Africa in autumn have been proposed: via Sicily and via Gibraltar. These routes include different over‐water distances to cross the Mediterranean Sea, and thus different proportions of flight modes (soaring–gliding vs flapping–gliding) with resulting different transport costs. Considering different scenarios of energy cost of transport, with flapping–gliding flight over water being more costly than flying over land using soaring–gliding flight, we predicted a maximum optimal detour of 1218 km. Both individuals reached Africa using the longest, detoured, route, avoiding the longest water crossing. To achieve this they began migrating northwards, keeping for ca 700 km a direction opposite to that followed by any other migrating bird from the Northern hemisphere in autumn. The comparison of optimal detour predictions with observed migratory tracks suggests that this migratory strategy prioritizes not only energy minimization, but also safety, given the mortality risk associated with the sea crossing. Finally, it is unlike that these inexperienced individuals followed such a complex route relying only on endogenous information and we therefore suggest, also on the basis of field observations, that social interactions (adult guidance) allow these individuals to learn the detoured route.     

Gliding flight in the American Kestrel (Falco sparverius): An electromyographic study

Electromyographic (EMG) activity was studied in American Kestrels (Falco sparverius) gliding in a windtunnel tilted to 8 degrees below the horizontal. Muscle activity was observed in Mm. biceps brachii, triceps humeralis, supracoracoideus, and pectoralis, and was absent in M. deltoideus major and M. thoracobrachialis (region of M. pectoralis). These active muscles are believed to function in holding the wing protracted and extended during gliding flight. Quantification of the EMG signals showed a lower level of activity during gliding than during flapping flight, supporting the idea that gliding is a metabolically less expensive form of locomotion than flapping flight. Comparison with the pectoralis musculature of specialized gliding and soaring birds suggests that the deep layer of the pectoralis is indeed used during gliding flight and that the slow tonic fibers found in soaring birds such as vultures represents a specialization for endurant gliding. It is hypothesized that these slow fibers should be present in the wing muscles that these birds use for wing protraction and extension, in addition to the deep layer of the pectoralis. © 1993 Wiley-Liss, Inc.     

Influence of weather conditions on the flight of migrating black storks

This study tested the potential influence of meteorological parameters (temperature, humidity, wind direction, thermal convection) on different migration characteristics (namely flight speed, altitude and direction and daily distance) in 16 black storks (Ciconia nigra). The birds were tracked by satellite during their entire autumnal and spring migration, from 1998 to 2006. Our data reveal that during their 27-day-long migration between Europe and Africa (mean distance of 4100 km), the periods of maximum flight activity corresponded to periods of maximum thermal energy, underlining the importance of atmospheric thermal convection in the migratory flight of the black stork. In some cases, tailwind was recorded at the same altitude and position as the birds, and was associated with a significant rise in flight speed, but wind often produced a side azimuth along the birds'' migratory route. Whatever the season, the distance travelled daily was on average shorter in Europe than in Africa, with values of 200 and 270 km d⁻¹, respectively. The fastest instantaneous flight speeds of up to 112 km h⁻¹ were also observed above Africa. This observation confirms the hypothesis of thermal-dependant flight behaviour, and also reveals differences in flight costs between Europe and Africa. Furthermore, differences in food availability, a crucial factor for black storks during their flight between Europe and Africa, may also contribute to the above-mentioned shift in daily flight speeds.     

Inland flights of young red‐eyed vireos Vireo olivaceus in relation to survival and habitat in a coastal stopover landscape

Inland dispersal of migrating land birds away from the coast, often opposite to the direction of migration, occurs frequently. Many of these movements may involve migrants seeking improved stopover conditions farther inland, but direct study of inland flights and of the ecological factors influencing their occurrence is limited. We used an automated telemetry array and ground‐tracking to assess flight behaviours, survival, and habitat use of young red‐eyed vireos Vireo olivaceus during fall migration at a coastal island and an inland stopover site in southwest Nova Scotia. We recorded inland flights for 41% (11/27) of individuals that departed the island. At least 25% of 16 individuals tagged at the inland site also relocated within the landscape prior to continuing migration, but due to the higher proportion of ambiguous flights at the inland site (44%) compared to the island (15%), we could not be sure if actual proportions of relocations differed between sites. Mortality on the island (at least 10 of 39 individuals) was significantly higher than at the inland site (0 of 16 individuals). At mainland sites near the coast where we found 6 of 11 individuals after they relocated away from the island, mortality remained high (2/6). Lack of deciduous canopy cover may have contributed to the high mortality on the island, but coastal mainland sites had a relatively high amount of deciduous canopy cover, similar to at the inland site where there was no mortality. Although coastal stopover sites may be important for migrating songbirds, especially before or after making a large water crossing, our results show that mortality can be much higher, and habitat poorer, at the coast compared to farther inland. Therefore, relocating inland may be an adaptive strategy for individuals that initially settle at the coast and that need to rest and refuel before they continue migration.     

Optimal flight behavior of soaring migrants: a case study of migrating steppe buzzards, Buteo buteo vulpinus

This article presents tests of the theoretical predictions onoptimal soaring and gliding flight of large, diurnal migrantsusing Pennycuick's program 2 for "bird flight performance."Predictions were compared with 141 observed flight paths ofmigrating steppe buzzards, Buteo buteo vulpinus. Calculationsof cross-country speed relative to the air included bird's airspeedsand sinking rates in interthermal gliding and climbing ratesin thermal circling. Steppe buzzards adjusted interthermal glidingairspeed . according to their actual climbing rate in thermalcircling. By optimizing their gliding airspeed, the birds maximizedtheir crosscountry performance relative to the air. Despitethis general agreement with the model, there was much scatterin the data, for the model neglects horizontal winds and updraftsduring the gliding phase. Lower sinking rates due to updraftsduring the gliding phases allowed many birds to achieve highercross-country speeds than predicted. In addition, birds reactedto different wind directions and speeds: in side and opposingwinds, the steppe buzzards compensated for wind displacementduring soaring and increased their gliding airspeed with decreasingtailwind component Nevenheless, cross-country speed relativeto the ground, which is the important measure for a migratorybird, was still higher under following winds. This study showsthat Pennycuick's program 2 provides reliable predictions onoptimal soaring and gliding behavior using realistic assumptionsand constants in the model, but a great deal of variation aroundthe mean is generated by factors not included in the model