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.     

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.     

Thermal soaring flight of birds and unmanned aerial vehicles

Thermal soaring saves much energy, but flying large distances in this form represents a great challenge for birds, people and unmanned aerial vehicles (UAVs). The solution is to make use of the so-called thermals, which are localized, warmer regions in the atmosphere moving upward with a speed exceeding the descent rate of birds and planes. Saving energy by exploiting the environment more efficiently is an important possibility for autonomous UAVs as well. Successful control strategies have been developed recently for UAVs in simulations and in real applications. This paper first presents an overview of our knowledge of the soaring flight and strategy of birds, followed by a discussion of control strategies that have been developed for soaring UAVs both in simulations and applications on real platforms. To improve the accuracy of the simulation of thermal exploitation strategies we propose a method to take into account the effect of turbulence. Finally, we propose a new GPS-independent control strategy for exploiting thermal updrafts.     

Estimating updraft velocity components over large spatial scales: contrasting migration strategies of golden eagles and turkey vultures

Soaring birds migrate in massive numbers worldwide. These migrations are complex and dynamic phenomena, strongly influenced by meteorological conditions that produce thermal and orographic uplift as the birds traverse the landscape. Herein we report on how methods were developed to estimate the strength of thermal and orographic uplift using publicly available digital weather and topography datasets at continental scale. We apply these methods to contrast flight strategies of two morphologically similar but behaviourally different species: golden eagle, Aquila chrysaetos, and turkey vulture, Cathartes aura, during autumn migration across eastern North America tracked using GPS tags. We show that turkey vultures nearly exclusively used thermal lift, whereas golden eagles primarily use orographic lift during migration. It has not been shown previously that migration tracks are affected by species-specific specialisation to a particular uplift mode. The methods introduced herein to estimate uplift components and test for differences in weather use can be applied to study movement of any soaring species.     

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.     

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.     

The influence of weather on the flight altitude of nocturnal migrants in mid‐latitudes

By altering its flight altitude, a bird can change the atmospheric conditions it experiences during migration. Although many factors may influence a bird's choice of altitude, wind is generally accepted as being the most influential. However, the influence of wind is not clearly understood, particularly outside the trade‐wind zone, and other factors may play a role. We used operational weather radar to measure the flight altitudes of nocturnally migrating birds during spring and autumn in the Netherlands. We first assessed whether the nocturnal altitudinal distribution of proportional bird density could be explained by the vertical distribution of wind support using three different methods. We then used generalized additive models to assess which atmospheric variables, in addition to altitude, best explained variability in proportional bird density per altitudinal layer each night. Migrants generally remained at low altitudes, and flight altitude explained 52 and 73% of the observed variability in proportional bird density in spring and autumn, respectively. Overall, there were weak correlations between altitudinal distributions of wind support and proportional bird density. Improving tailwind support with height increased the probability of birds climbing to higher altitude, but when birds did fly higher than normal, they generally concentrated around the lowest altitude with acceptable wind conditions. The generalized additive model analysis also indicated an influence of temperature on flight altitudes, suggesting that birds avoided colder layers. These findings suggested that birds increased flight altitudes to seek out more supportive winds when wind conditions near the surface were prohibitive. Thus, birds did not select flight altitudes only to optimize wind support. Rather, they preferred to fly at low altitudes unless wind conditions there were unsupportive of migration. Overall, flight altitudes of birds in relation to environmental conditions appear to reflect a balance between different adaptive pressures.     

The gliding speed of migrating birds: slow and safe or fast and risky?

Aerodynamic theory postulates that gliding airspeed, a major flight performance component for soaring avian migrants, scales with bird size and wing morphology. We tested this prediction, and the role of gliding altitude and soaring conditions, using atmospheric simulations and radar tracks of 1346 birds from 12 species. Gliding airspeed did not scale with bird size and wing morphology, and unexpectedly converged to a narrow range. To explain this discrepancy, we propose that soaring‐gliding birds adjust their gliding airspeed according to the risk of grounding or switching to costly flapping flight. Introducing the Risk Aversion Flight Index (RAFI, the ratio of actual to theoretical risk‐averse gliding airspeed), we found that inter‐ and intraspecific variation in RAFI positively correlated with wing loading, and negatively correlated with convective thermal conditions and gliding altitude, respectively. We propose that risk‐sensitive behaviour modulates the evolution (morphology) and ecology (response to environmental conditions) of bird soaring flight.     

Flight by night or day? Optimal daily timing of bird migration

Many migratory bird species fly mainly during the night (nocturnal migrants), others during daytime (diurnal migrants) and still others during both night and day. Need to forage during the day, atmospheric structure, predator avoidance and orientation conditions have been proposed as explanations for the widespread occurrence of nocturnal migration. However, the general principles that determine the basic nocturnal-diurnal variation in flight habits are poorly known. In the present study optimal timing of migratory flights, giving the minimum total duration of the migratory journey, is evaluated in a schematic way in relation to ecological conditions for energy gain in foraging and for energy costs in flight. There exists a strong and fundamental advantage of flying by night because foraging time is maximized and energy deposition can take place on days immediately after and prior to the nocturnal flights. The increase in migration speed by nocturnal compared with diurnal migration will be largest for birds with low flight costs and high energy deposition rates. Diurnal migration will be optimal if it is associated with efficient energy gain immediately after a migratory flight because suitable stopover/foraging places have been located during the flight or if energy losses during flight are substantially reduced by thermal soaring and/or by fly-and-forage migration. A strategy of combined diurnal and nocturnal migration may be optimal when birds migrate across regions with relatively poor conditions for energy deposition (not only severe but also soft barriers). Predictions about variable timing of migratory flights depending on changing foraging and environmental conditions along the migration route may be tested for individual birds by analysing satellite tracking results with respect to daily travel routines in different regions. Documenting and understanding the adaptive variability in daily travel schedules among migrating animals constitute a fascinating challenge for future research.     

Environmental effects on flying migrants revealed by radar

Migratory animals are affected by various factors during their journeys, and the study of animal movement by radars has been instrumental in revealing key influences of the environment on flying migrants. Radars enable the simultaneous tracking of many individuals of almost all sizes within the radar range during day and night, and under low visibility conditions. We review how atmospheric conditions, geographic features and human development affect the behavior of migrating insects and birds as recorded by radars. We focus on flight initiation and termination, as well as in‐flight behavior that includes changes in animal flight direction, speed and altitude. We have identified several similarities and differences in the behavioral responses of aerial migrants including an overlooked similarity in the use of thermal updrafts by very small (e.g. aphids) and very large (e.g. vultures) migrants. We propose that many aerial migrants modulate their migratory flights in relation to the interaction between atmospheric conditions and geographic features. For example, aerial migrants that encounter crosswind may terminate their flight or continue their migration and may also drift or compensate for lateral displacement depending on their position (over land, near the coast or over sea). We propose several promising directions for future research, including the development and application of algorithms for tracking insects, bats and large aggregations of animals using weather radars. Additionally, an important contribution will be the spatial expansion of aeroecological radar studies to Africa, most of Asia and South America where no such studies have been undertaken. Quantifying the role of migrants in ecosystems and specifically estimating the number of departing birds from stopover sites using low‐elevation radar scans is important for quantifying migrant–habitat relationships. This information, together with estimates of population demographics and migrant abundance, can help resolve the long‐term dynamics of migrant populations facing large‐scale environmental changes.     

Flight responses by a migratory soaring raptor to changing meteorological conditions

Soaring birds that undertake long-distance migration should develop strategies to minimize the energetic costs of endurance flight. This is relevant because condition upon completion of migration has direct consequences for fecundity, fitness and thus, demography. Therefore, strong evolutionary pressures are expected for energy minimization tactics linked to weather and topography. Importantly, the minute-by-minute mechanisms birds use to subsidize migration in variable weather are largely unknown, in large part because of the technological limitations in studying detailed long-distance bird flight. Here, we show golden eagle (Aquila chrysaetos) migratory response to changing meteorological conditions as monitored by high-resolution telemetry. In contrast to expectations, responses to meteorological variability were stereotyped across the 10 individuals studied. Eagles reacted to increased wind speed by using more orographic lift and less thermal lift. Concomitantly, as use of thermals decreased, variation in flight speed and altitude also decreased. These results demonstrate how soaring migrant birds can minimize energetic expenditures, they show the context for avian decisions and choices of specific instantaneous flight mechanisms and they have important implications for design of bird-friendly wind energy.     

Effect of wind,thermal convection,and variation in flight strategies on the daily rhythm and flight paths of migrating raptors at Georgia's Black Sea coast

Every autumn, large numbers of raptors migrate through geographical convergence zones to avoid crossing large bodies of water. At coastal convergence zones, raptors may aggregate along coastlines because of convective or wind conditions. However, the effect of wind and thermal convection on migrating raptors may vary depending on local landscapes and weather, and on the flight strategies of different raptors. From 20 August to 14 October 2008 and 2009, we studied the effect of cloud development and crosswinds on the flight paths of raptors migrating through the eastern Black Sea convergence zone, where coastal lowlands at the foothills of the Pontic Mountains form a geographical bottleneck 5‐km‐wide near Batumi, the capital of the Independent Republic of Ajaria in southwestern Georgia. To identify key correlates of local aggregation, we examined diurnal variation in migration intensity and coastal aggregation of 11 species of raptors categorized based on size and flight strategies. As reported at other convergence zones, migration intensity of large obligate‐soaring species peaked during the core period of thermal activity at mid‐day. When clouds developed over interior mountains and limited thermal convection, these large obligate‐soaring species aggregated near the coast. However, medium‐sized soaring migrants that occasionally use flapping flight did not aggregate at the coast when clouds over the mountains weakened thermal convection. Numbers of alternate soaring‐flapping harriers (Circus spp.) peaked during early morning, with these raptors depending more on flapping flight during a time of day with poor thermal convection. Small sparrowhawks (Accipiter spp.) aggregated at the coast during periods when winds blew offshore, suggesting aggregation caused by wind drift. Thus, weather conditions, including cloud cover and wind speed and direction, can influence the daily rhythm and flight paths of migrating raptors and, therefore, should be accounted for before inferring population trends from migration counts.     

Scaling of Soaring Seabirds and Implications for Flight Abilities of Giant Pterosaurs

The flight ability of animals is restricted by the scaling effects imposed by physical and physiological factors. In comparisons of the power available from muscle and the mechanical power required to fly, it is predicted that the margin between the powers should decrease with body size and that flying animals have a maximum body size. However, predicting the absolute value of this upper limit has proven difficult because wing morphology and flight styles varies among species. Albatrosses and petrels have long, narrow, aerodynamically efficient wings and are considered soaring birds. Here, using animal-borne accelerometers, we show that soaring seabirds have two modes of flapping frequencies under natural conditions: vigorous flapping during takeoff and sporadic flapping during cruising flight. In these species, high and low flapping frequencies were found to scale with body mass (mass ^−0.30 and mass ^−0.18) in a manner similar to the predictions from biomechanical flight models (mass ^−1/3 and mass ^−1/6). These scaling relationships predicted that the maximum limits on the body size of soaring animals are a body mass of 41 kg and a wingspan of 5.1 m. Albatross-like animals larger than the limit will not be able to flap fast enough to stay aloft under unfavourable wind conditions. Our result therefore casts doubt on the flying ability of large, extinct pterosaurs. The largest extant soarer, the wandering albatross, weighs about 12 kg, which might be a pragmatic limit to maintain a safety margin for sustainable flight and to survive in a variable environment.     

SOARING BEHAVIOUR AND PERFORMANCE OF SOME EAST AFRICAN BIRDS, OBSERVED FROM A MOTOR-GLIDER

Various species of soaring birds were studied by following them in a motor-glider, mainly over the Serengeti National Park, Tanzania. The characteristics of thermal convection in the study area are described in general terms. The two vulture species of the genus Gyps live by scavenging among the herds of migratory ungulates, especially Wildebeest. They are not territorial, and gather in large numbers on kills. When raising young they may be obliged by game movements to forage at long distances from their nests. Their cross-country performance is adequate for a foraging radius of over 100 km in dry-season conditions. Their ability to compete with Spotted Hyaenas is thought to depend partly on this factor and partly on an advantage in arriving early at kills. These two species appear to find food more by watching other vultures than by searching for it directly. The Lappet-faced and White-headed Vultures are thought to be sedentary, and to depend on thorough searching of a fixed foraging territory, rather than on following migratory game. They have lower wing loadings than the Gyps vultures, and were not seen cross-country flying. They never gather in large numbers. The Hooded Vulture is a solitary nester, but it does fly across country, and does gather at kills. Vultures soar individually, and seem to be good at exploiting such phenomena as thermal streets. They do not travel in flocks. Tawny and Martial Eagles react positively to the glider, and are suspected of regarding it as potential prey. White Storks migrate between Europe and Africa, and also travel about within East Africa, by thermal soaring. They soar in flocks, and unlike vultures rely on co-ordinated social behaviour to locate thermals. In choosing their route, they often fail to react to obvious weather signs. They enter cumulus clouds from the bottom when thermalling, but probably do not climb far above cloudbase. Marabou Storks soar individually, but also sometimes travel in flocks. When doing so, they show less lateral spreading than White Storks, which reduces the effectiveness of the flock as a thermal-finding unit; on the other hand, they do seem to react to visible weather signs, like vultures or glider pilots. White Pelicans, which travel by thermal soaring between different lakes in the Rift Valley, show the most highly co-ordinated social soaring behaviour. Unlike White Storks, they fly in formation even when circling. Storks and pelicans showed more signs of alarm when approached by the glider than did the vultures or birds of prey. This could be due to their being preyed upon in flight, for instance by Martial Eagles. The basis of conventional thermal cross-country flying is outlined, and it is explained why the high wing loadings of the Gyps vultures are appropriate to their peripatetic habits. A method of thermal soaring without circling is discussed, and shown to be more readily feasible for small than for large birds. Some differences in soaring techniques between birds and glider pilots are interpreted in the light of this calculation. A case in which Black Kites apparently used this technique to soar in random turbulence is described. The cross-country speed attainable by thermal soaring should be similar to the cruising speed under power in both large and small birds. Rough calculations of the energy costs suggest that a large bird (White Stork) should reduce its fuel consumption by a factor of 23 by soaring rather than flying under power, whereas this factor would be only 2–4 for a small bird (Bonelli's Warbler). Other reasons why thermal soaring is an advantageous means of travel for large but not for small birds are also indicated.     

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.     

Wind estimation based on thermal soaring of birds

The flight performance of birds is strongly affected by the dynamic state of the atmosphere at the birds' locations. Studies of flight and its impact on the movement ecology of birds must consider the wind to help us understand aerodynamics and bird flight strategies. Here, we introduce a systematic approach to evaluate wind speed and direction from the high‐frequency GPS recordings from bird‐borne tags during thermalling flight. Our method assumes that a fixed horizontal mean wind speed during a short (18 seconds, 19 GPS fixes) flight segment with a constant turn angle along a closed loop, characteristic of thermalling flight, will generate a fixed drift for each consequent location. We use a maximum‐likelihood approach to estimate that drift and to determine the wind and airspeeds at the birds' flight locations. We also provide error estimates for these GPS‐derived wind speed estimates. We validate our approach by comparing its wind estimates with the mid‐resolution weather reanalysis data from ECMWF, and by examining independent wind estimates from pairs of birds in a large dataset of GPS‐tagged migrating storks that were flying in close proximity. Our approach provides accurate and unbiased observations of wind speed and additional detailed information on vertical winds and uplift structure. These precise measurements are otherwise rare and hard to obtain and will broaden our understanding of atmospheric conditions, flight aerodynamics, and bird flight strategies. With an increasing number of GPS‐tracked animals, we may soon be able to use birds to inform us about the atmosphere they are flying through and thus improve future ecological and environmental studies.     

The Flight Behavior of Migrating Birds in Changing Wind Fields: Radar and Visual Analyses

This paper examines the influence of atmospheric structure andmotion (principally winds aloft) on the flight behavior andaltitudinal distribution of migrating songbirds. Bird migrationdata that I gathered using surveillance radars operated by theUnited States National Weather Service and the Federal AviationAdministration and a vertically directed fixed-beam marine radarmounted on a mobile laboratory are analyzed in relation to windsaloft. Migrating birds appear to fly at altitudes where windswill minimize the cost of transport and assist movements inseasonally appropriate directions. When migratory flights occurat altitudes that are higher than usual, a significant correlationexists between the altitude of densest migration and the altitudeof most favorable wind. Lower altitudes may be favored overslightly more favorable winds at much higher altitudes. Radardata on the flight behavior of migrating birds in the vicinityof frontal systems is also examined. The flight strategies ofmigrants (fly over the front, change the direction of flight,or land and terminate the flight) differ depending on seasonand the "thickness" of the front. Recent migration studies thatare related to atmospheric structure and motion are summarizedand related to atmospheric processes operating simultaneouslyat vastly different spatial and temporal scales.     

Predicting spatial patterns of eagle migration using a mesoscale atmospheric model: a case study associated with a mountain-ridge wind development

High resolution numerical atmospheric modeling around a mountain ridge in Northeastern British Columbia (BC), Canada was performed in order to examine the influence of meteorology and topography on Golden Eagle migration pathways at the meso-scale (tens of km). During three eagle fall migration periods (2007–2009), local meteorological conditions on the day of peak bird counts were modeled using the Regional Atmospheric Modeling System (RAMS) mesoscale model. Hourly local surface wind speed, wind direction, temperature, pressure and relative humidity were also monitored during these migration periods. Eagle migration flight paths were observed from the ground and converted to three-dimensional tracks using ArcGIS. The observed eagle migration flight paths were compared with the modeled vertical velocity wind fields. Flight tracks across the study area were also simulated using the modeled vertical velocity field in a migration model based on a fluid-flow analogy. It was found that both the large-scale weather conditions and the horizontal wind fields across the study area were broadly similar on each of the modeled migration days. Nonetheless, the location and density of flight tracks across the domain varied between days, with the 2007 event producing more tracks to the southwest of the observation location than the other 2 days. The modeled wind fields suggest that it is not possible for the eagles to traverse the study area without leaving updraft regions, but birds do converge on the locations of updrafts as they move through the area. Statistical associations between observed eagles positions and the vertical velocity field suggest that to the northwest (and to a lesser extent the southwest) of the main study ridge (Johnson col), eagles can always find updrafts but that they must pass through downdraft regions in the NE and SE as they make their way across the study area. Finally, the simulated flight tracks based on the fluid-flow model and the vertical velocity fields are in general agreement with the observed flight track patterns. Our results suggest that use of high resolution meteorological fields to locate the occurrence of updrafts in proposed ridge-line wind installations could aid in predicting, and mitigating for, convergence points in raptor migrations.     

Obligate vertebrate scavengers must be large soaring fliers

Among extant vertebrates, only the 23 species of vulture are obligate scavengers. We use an energetic modelling approach to explore the constraints imposed by an obligate scavenging lifestyle, and to ask whether obligate scavengers must always be avian and generally large-bodied users of soaring flight. Our model found that aerial scavengers always out-competed postulated terrestrial ones, mainly because flight allows area to be searched much more rapidly for carrion. Soaring was favoured over flapping flight because the reduction in flight speed (and so rate of area search) was more than compensated for by the decrease in the costs of transport. Large individual size is selected for if carrion is available in large packages, when obligate scavenger feed only infrequently, and so must be able to survive on body reserves in the periods between discovering food falls. In the absence of avian radiation, an obligate terrestrial scavenger seems energetically feasible, but we argue that such a beast is unlikely to have evolved. In birds, in order to become exclusive scavengers, vultures have needed to specialize for efficient soaring flight as a low energy form of travel, and as a consequence they have lost the agility needed to kill prey. In mammals, however, no comparable trade-off occurs. So for terrestrial carnivores there is probably no strong selection pressure towards being an exclusive scavenger. Indeed it will perhaps always be more advantageous to retain the flexibility of obtaining food by either predation or scavenging.     

Soaring energetics and glide performance in a moving atmosphere

Here, we analyse the energetics, performance and optimization of flight in a moving atmosphere. We begin by deriving a succinct expression describing all of the mechanical energy flows associated with gliding, dynamic soaring and thermal soaring, which we use to explore the optimization of gliding in an arbitrary wind. We use this optimization to revisit the classical theory of the glide polar, which we expand upon in two significant ways. First, we compare the predictions of the glide polar for different species under the various published models. Second, we derive a glide optimization chart that maps every combination of headwind and updraft speed to the unique combination of airspeed and inertial sink rate at which the aerodynamic cost of transport is expected to be minimized. With these theoretical tools in hand, we test their predictions using empirical data collected from a captive steppe eagle (Aquila nipalensis) carrying an inertial measurement unit, global positioning system, barometer and pitot tube. We show that the bird adjusts airspeed in relation to headwind speed as expected if it were seeking to minimize its aerodynamic cost of transport, but find only weak evidence to suggest that it adjusts airspeed similarly in response to updrafts during straight and interthermal glides.This article is part of the themed issue ‘Moving in a moving medium: new perspectives on flight’.