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.     

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’.     

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 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.     

Confronting the winds: orientation and flight behaviour of roosting swifts, Apus apus

Swifts, Apus apus, spend the night aloft and this offers an opportunity to test the degree of adaptability of bird orientation and flight to different ecological situations. We predicted the swifts' behaviour by assuming that they are adapted to minimize energy expenditure during the nocturnal flight and during a compensatory homing flight if they become displaced by wind. We tested the predictions by recording the swifts' altitudes, speeds and directions under different wind conditions with tracking radar; we found an agreement between predictions and observations for orientation behaviour, but not for altitude and speed regulation. The swifts orientated consistently into the head wind, with angular concentration increasing with increasing wind speed. However, contrary to our predictions, they did not select altitudes with slow or moderate winds, nor did they increase their airspeed distinctly when flying into strong head winds. A possible explanation is that their head-wind orientation is sufficient to keep nocturnal displacement from their home area within tolerable limits, leaving flight altitude to be determined by other factors (correlated with temperature), and airspeed to show only a marginal increase in strong winds. The swifts were often moving "backwards", heading straight into the wind but being overpowered by wind speeds exceeding their airspeed. The regular occurrence of such flights is probably uniquely associated with the swifts' remarkable habit of roosting on the wing.     

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.     

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.     

Commuting fruit bats beneficially modulate their flight in relation to wind

When animals move, their tracks may be strongly influenced by the motion of air or water, and this may affect the speed, energetics and prospects of the journey. Flying organisms, such as bats, may thus benefit from modifying their flight in response to the wind vector. Yet, practical difficulties have so far limited the understanding of this response for free-ranging bats. We tracked nine straw-coloured fruit bats (Eidolon helvum) that flew 42.5 ± 17.5 km (mean ± s.d.) to and from their roost near Accra, Ghana. Following detailed atmospheric simulations, we found that bats compensated for wind drift, as predicted under constant winds, and decreased their airspeed in response to tailwind assistance such that their groundspeed remained nearly constant. In addition, bats increased their airspeed with increasing crosswind speed. Overall, bats modulated their airspeed in relation to wind speed at different wind directions in a manner predicted by a two-dimensional optimal movement model. We conclude that sophisticated behavioural mechanisms to minimize the cost of transport under various wind conditions have evolved in bats. The bats’ response to the wind is similar to that reported for migratory birds and insects, suggesting convergent evolution of flight behaviours in volant organisms.     

Nocturnal passerine migration without tailwind assistance

Nocturnal passerine migrants could substantially reduce the amount of energy spent per distance covered if they fly with tailwind assistance and thus achieve ground speeds that exceed their airspeeds (the birds’ speed in relation to the surrounding air). We analysed tracking radar data from two study sites in southern and northern Scandinavia and show that nocturnally migrating passerines, during both spring and autumn migration, regularly travelled without tailwind assistance. Average ground and airspeeds of the birds were strikingly similar for all seasonal and site‐specific samples, demonstrating that winds had little overall influence on the birds’ resulting travel speeds. Distributions of wind effects, measured as (1) the difference between ground and airspeed and (2) the tail/headwind component along the birds’ direction of travel, showed peaks close to a zero wind effect, indicating that the migratory flights often occurred irrespective of wind direction. An assessment of prevailing wind speeds at the birds’ mean altitude indicated a preference for lower wind speeds, with flights often taking place in moderate winds of 3–10 m/s. The limited frequency of wind‐assisted flights among the nocturnal passerine migrants studied is surprising and in clear contrast to the strong selectivity of tailwinds exhibited by some other bird groups. Relatively high costs of waiting for favourable winds, rather low probabilities of occurrence of tailwind conditions and a need to use a large proportion of nights for flying are probably among the factors that explain the lack of a distinct preference for wind‐assisted flights among nocturnal passerine migrants.     

Testing predictions from flight mechanical theory: a case study of Cory’s shearwater and Audouin’s gull

Predictions from flight mechanical theory concerning optimal flight speeds were tested in the field in two Mediterranean seabirds, the Cory’s shearwater Calonectris diomedea and the Audouin’s gull Larus audouinii. Both species were commuting off the coast of Isola di San Pietro, 6 km south-west of the coast of Sardinia. Heading and airspeed were obtained by vector calculation of flight tracks and measured wind. The Cory’s shearwater used a mixture of gliding and active flight. At low wind speeds the proportion of active flight was large but it decreased with increasing wind speed. The mean airspeed was 12.0 m s^–1, which is not significantly different from minimum power speed (V _mp) in active flight or the speed for best glide (V _bg) used in gliding flight. However, the shearwaters showed a significant response to wind increment/decrement, indicating that they were not flying at V _mp, which should be unaffected by head and tailwind. Furthermore, shearwaters can potentially reduce induced drag by the ground effect while flying close to the sea surface at weak winds, which leads to a reduction in characteristic flight speed. We suspect that the predictions for gliding flight are most valid for shearwaters at moderate to high wind speeds, when they should be maximising distance by using V _bg. Audouin’s gulls used active flight exclusively, with a mean airspeed of 11.3 m s^–1 that was significantly different from the predicted V _mp. Interestingly, though, the gulls did not show any significant wind response, indicating that they were flying close to their true V _mp when foraging along the coast. Received: 17 May 2000 / Received in revised form: 21 November 2000 / Accepted: 8 January 2001     

Flight dynamics of Cory’s shearwater foraging in a coastal environment

Flight dynamics theories are influenced by two major topics: how birds adapt their flight to cope with heterogeneous habitats, and whether birds plan to use the wind field or simply experience it. The aim of this study was to understand the flight dynamics of free-flying Cory’s shearwaters in relation to the wind characteristics on the coastal upwelling region of continental Portugal. We deployed recently miniaturised devices—global positioning system loggers to collect precise and detailed information on birds’ positions and motions. Prevalent winds were blowing from the north-east and adults used those winds by adjusting their flight directions mainly towards north-west and south-west, flying with cross and tail winds, respectively, and avoiding head winds. This is confirmation that Cory’s shearwaters use a shear soaring flying strategy while exploiting the environment for food: adults foraged mainly with cross winds and their ground speed was not constant during all foraging trips as it changed dynamically as a result of the ocean surface shear winds. During travelling phases, ground speed was strongly influenced by the position of the bird with regard to the wind direction, as ground speed increased significantly with increasing tail wind component (TWC) values. Adults appear to choose foraging directions to exploit ambient wind, in order to improve shear soaring efficiency (cross winding) and exploit diurnal changes in tail wind strength to maximise commuting efficiency. We report, for the first time, precise ground speed values (GPS-derived data) and computed actual flight speed values (using TWC analysis) for Cory’s shearwater.     

Flight speed of seabirds in relation to wind speed and direction

We studied flight speed among all major seabird taxa. Our objectives were to provide further insight into dynamics of seabird flight and to develop allometric equations relating ground speed to wind speed and direction for use in adjusting seabird density estimates (calculated from surveys at sea) for the effect of bird movement. We used triangulation at sea to estimate ground speeds of 1562 individuals of 98 species. Species sorted into 25 “groups” based on similarity in ground speeds and taxonomy. After they were controlled for differences inground speed, the 25 groups sorted into eight major “types” on the basis of response to wind speed and wind direction. Wind speed and direction explained 1664% of the variation in ground speed among seabird types. For analyses on air speed (ground speed minus apparent wind speed), we divided the 25 groups according to four flight styles: gliding, flap-gliding, glide-flapping and flapping. Tailwind speed had little effect on air speed of gliders (albatrosses and large gadfly petrels), but species that more often used flapping decreased air speed with increase in tailwinds. All species increased air speeds significantly with increased headwinds. Gliders showed the greatest increase relative to increase in headwind speed and flappers the least. With tailwind flight, air speeds were greatest among species with highest wing loading for each flight style except gliders, which showed no relationship. For headwind flight, species with higher wing loading had higher air speeds; however, the relation was weaker in flappers compared with species using some amount of gliding. In contrast, analyses for air speed ratio (i.e. difference between air speed in acrosswinds [with no apparent wind] and speed flown into headwinds, or with tailwinds, divided by speed acrosswind) revealed that among species using some flapping, and with lower wing loading (surface-feeding shearwaters, small gadfly petrels, storm petrels, phalaropes, gulls and terns), adjusted air speeds more than those with higher wing loading (alcids, “diving shearwaters”, “Manx-type shearwaters”, pelicans, boobies and cormorants). As a result, most flappers of low wing loading flew much faster than V_mr (the most energy efficient air speed per distance flown) when flying into headwinds. We suggest that better-than-predicted gliding performance with acrosswinds and tailwinds of large gadfly petrels, compared with albatrosses, resulted from a different type of “soaring” not previously described in seabirds.     

Skylark optimal flight speeds for flying nowhere and somewhere

Flight is energetically very costly. For birds the mechanicalpower in relation to airspeed is characterized by a U-shapedfunction. From this function we can derive optimal flight speedsassociated with minimum power (V_mp), minimum cost of transport(V_mr) and minimum overall time of migration (V_mt). Since flightis energetically so costly, aerial displays and song flightcan potentially serve as signals reliably indicating the individualquality or resource potential of the signaler. In order to maximizethe amount of song flight produced, we expect V_mp during songflight, while during migration we rather expect V_mr or V_mv Wecompared flight speeds of skylarks (Alauda arvensis) duringsong flight and migration flight, respectively. In this speciespredicted V_mp = 5.5 m/s, V_mr = 10.5 m/s, and V_mt = 12.1 m/s.The preferred airspeed during song flight did not differ significantlyfrom the predicted V_mp, while airspeed during migration wassignificantly higher than V_mr and Vmp indicating that flightspeed is a flexible trait that birds adjust to different situations.Why the skylarks speed up so much on migration is still unclear,but it may be that due to the shape of the predicted power curve,variation in cost of transport at high speeds is relativelysmall.     

Effects of host-odour plume altitude and changing wind velocity on upwind flight manoeuvres of a specialist braconid parasitoid

Abstract. Females of the specialist parasitoid, Microplitis croceipes (Cresson) (Hymenoptera: Braconidae), were released in a wind tunnel into host-odour plumes dispersed by winds of three velocities and winds whose speed was changed while the wasps were engaged in upwind flight. In steady winds of 61, 122 and 183 cms-^-1, wasps maintained similar 'preferred' ground speeds by adjusting their airspeed, while turning to a lesser degree as wind velocity increased. In winds of changing velocity (either increasing or decreasing within a 60–100 cm s-¹ range), wasps lowered their rate of upwind progress, leading to more tortuous tracks. During changing wind speeds longitudinal image flow decreased. Wasps flying in host-odour plumes 10 cm and 20 cm above the flight tunnel floor in a 122 cm s-¹ wind had similar ground speeds; thus their rate of ventral visual image flow varied two-fold. M.croceipes may 'aim' upwind by comparing how changes in the course angle vary with the direction of visual image flow. During changing wind velocities the relationship between changes in visual and flight muscle generated torque is ambiguous. Under these conditions most wasps cast, a manoeuvre characterized by wide lateral excursions across the wind without upwind progress. Once wind speed stabilizes, flight straightens out and upwind flight resumes.     

Interactive effects of body mass changes and species‐specific morphology on flight behavior of chick‐rearing Antarctic fulmarine petrels under diurnal wind patterns

For procellariiform seabirds, wind and morphology are crucial determinants of flight costs and flight speeds. During chick‐rearing, parental seabirds commute frequently to provision their chicks, and their body mass typically changes between outbound and return legs. In Antarctica, the characteristic diurnal katabatic winds, which blow stronger in the mornings, form a natural experimental setup to investigate flight behaviors of commuting seabirds in response to wind conditions. We GPS‐tracked three closely related species of sympatrically breeding Antarctic fulmarine petrels, which differ in wing loading and aspect ratio, and investigated their flight behavior in response to wind and changes in body mass. Such information is critical for understanding how species may respond to climate change. All three species reached higher ground speeds (i.e., the speed over ground) under stronger tailwinds, especially on return legs from foraging. Ground speeds decreased under stronger headwinds. Antarctic petrels (Thalassoica antarctica; intermediate body mass, highest wing loading, and aspect ratio) responded stronger to changes in wind speed and direction than cape petrels (Daption capense; lowest body mass, wing loading, and aspect ratio) or southern fulmars (Fulmarus glacialoides; highest body mass, intermediate wing loading, and aspect ratio). Birds did not adjust their flight direction in relation to wind direction nor the maximum distance from their nests when encountering headwinds on outbound commutes. However, birds appeared to adjust the timing of commutes to benefit from strong katabatic winds as tailwinds on outbound legs and avoid strong katabatic winds as headwinds on return legs. Despite these adaptations to the predictable diurnal wind conditions, birds frequently encountered unfavorably strong headwinds, possibly as a result of weather systems disrupting the katabatics. How the predicted decrease in Antarctic near‐coastal wind speeds over the remainder of the century will affect flight costs and breeding success and ultimately population trajectories remains to be seen.     

The effect of optic flow cues on honeybee flight control in wind

To minimize the risk of colliding with the ground or other obstacles, flying animals need to control both their ground speed and ground height. This task is particularly challenging in wind, where head winds require an animal to increase its airspeed to maintain a constant ground speed and tail winds may generate negative airspeeds, rendering flight more difficult to control. In this study, we investigate how head and tail winds affect flight control in the honeybee Apis mellifera, which is known to rely on the pattern of visual motion generated across the eye—known as optic flow—to maintain constant ground speeds and heights. We find that, when provided with both longitudinal and transverse optic flow cues (in or perpendicular to the direction of flight, respectively), honeybees maintain a constant ground speed but fly lower in head winds and higher in tail winds, a response that is also observed when longitudinal optic flow cues are minimized. When the transverse component of optic flow is minimized, or when all optic flow cues are minimized, the effect of wind on ground height is abolished. We propose that the regular sidewards oscillations that the bees make as they fly may be used to extract information about the distance to the ground, independently of the longitudinal optic flow that they use for ground speed control. This computationally simple strategy could have potential uses in the development of lightweight and robust systems for guiding autonomous flying vehicles in natural environments.     

Flight speeds and climb rates of Brent Geese: mass-dependent differences between spring and autumn migration

Aerodynamic theories of bird flight predict that horizontal flight speed will increase with increasing load whereas vertical flight speed will decrease. Horizontal flight speed for birds minimizing overall time on migration is predicted to be higher than flight speed for birds minimizing energy expenditure. In this study we compare flight speeds of Brent Geese Branta b. bernicla recorded by tracking radar and optical range finder during spring and autumn migration in southernmost Sweden, testing the above-mentioned predictions. Geese passing Sweden in spring are substantially heavier than in autumn and there might also be a stronger element of time-selection in spring than in autumn. Recorded airspeeds were significantly higher in spring (mean 19.0 m s⁻¹) than in autumn (mean 17.3 m s⁻¹), the average difference being slightly larger than predicted due to the mass difference alone. The effects on airspeed of wind, vertical speed, flock size and altitude were also analysed, but none of these factors could explain the seasonal difference in airspeed. Hence, the results support the hypothesis of mass-dependent flight speed adjustment. The difference between the two seasons was not large enough to corroborate the hypothesis of a stronger element of time-selection in spring, but this hypothesis cannot be rejected. Vertical flight speeds were lower in spring than in autumn, supporting a negative effect of load on birds' flight power margin.     

Climb and flight speeds of shorebirds embarking on an intercontinental flight; do they achieve the predicted optimal behaviour?

Most Arctic-breeding waders wintering in West Africa cover the first 4000 km of their northward journey in spring by a single flight to western Europe. We examined the extent to which waders economize their flight behaviour during departure by comparing climb rates and forward flight speeds with predictions based on flight mechanic theory and the relevant morphological measurements made of birds collected on the site. With an optical range finder, we followed 98 wader flocks on their departure from Banc d'Arguin in Mauritania. We also measured wind speed and direction at different altitudes by tracking helium-filled balloons and thus were able to deduce airspeeds from groundspeeds of the departing flocks. Of the nine species examined, six showed the predicted negative relationship between climb rate and airspeed, although only one was statistically significant. By normalizing the data, we found a statistically significant negative correlation across all species. Although 17% of the observed climb rates were greater than the predicted theoretical maximum, the average observed climb rate was lower than the predicted optimum and the average observed airspeed was higher. The absolute deviations of climb rates from theory may have been because of the existence of pockets of rising and sinking air at the boundary of desert and ocean. That the absolute deviations in average climb rate and airspeed followed the predicted negative relationship is in accordance with the current theory of flight mechanics.     

Predator versus prey: on aerial hunting and escape strategies in birds

Predator and prey attack-escape performance is likely to bethe outcome of an evolutionary arms race. Predatory birds aretypically larger than their prey, suggesting different flightperformances. We analyze three idealized attack-escape situationsbetween predatory and prey birds: climbing flight escape, horizontalspeeding, and turning and escape by diving. Generally a smallerbird will outclimb a larger predator and hence outclimbing shouldbe a common escape strategy. However, some predators such asthe Eleonora's falcon (Falco elenorae) has a very high rateof climb for its size. Prey species with an equal or highercapacity to climb fast, such as the swift Apus apus, usuallyadopt climbing escape when attacked by Eleonora's falcons.To analyze the outcome of the turning gambit between predatorand prey we use a Howland diagram, where the relative lineartop speeds and minimum turning radii of prey and predator definethe escape and danger zones. Applied to the Eleonora's falconand some potential prey species, this analysis indicates thatthe falcon usually wins against the example prey species; thatis, the prey will be captured. Level maneuvering hunting isthe most common strategy seen in Eleonora's falcons. To avoidcapture via use of this strategy by a predator, the prey shouldbe able to initiate tight turns at high linear speed, whichis facilitated by a low wing loading (weight per unit of wingarea). High diving speed is favored by large size. If close enough to safe cover, a prey might still opt for a verticaldive to escape in spite of lower terminal diving speed thanthat of the predator. On the basis of aerodynamic considerationswe discuss escape flight strategies in birds in relation tomorphological adaptations.     

Flight characteristics of birds:

This is the first part of a study on flight characteristics of birds and presents an annotated list of flight speeds of 139 western Palearctic species. All measurements were taken with the same tracking radar and corrected for wind influence according to radar-tracked wind-measuring balloons. Graphical presentation of the birds' air speeds emphasizes the wide variation of speeds within species and allows easy comparison between taxonomic groups, species, and types of flight. Unlike theoretical predictions, speeds increase only slightly with size. The larger species seem to be increasingly limited to speeds close to their speed of minimum power consumption V_mp',. Released birds, apparently reluctant to depart with migratory speed, fly at considerably lower speeds than migrating conspecifics. While large birds seem to be limited to speeds around V _mp', smaller birds seem to be capable of selecting between various speeds, approaching predicted V _mp, when tending to remain airborne at low cost, but flying at much higher speeds when tending to make best progress at low cost (around predicted speed of maximum range V _mr,). Predictions of air speeds by aerodynamic models proved to be too low for small birds because the models do not account for the gain in speed attained by the reduction in profile drag during bounding flight of small passerines. The models predict excessive speeds for large birds because the power output available for flight seems to decline much more with size than previously assumed.