Evaluation of long-distance orientation in birds on the basis of migration routes recorded by radar and satellite tracking

Predicted flight trajectories differ depending on which orientation cues are used by migrating birds. Results from radar and satellite tracking of migrating birds can be used to test which of the predicted trajectories shows the best fit with observed flight routes, supporting the use of the associated orientation mechanism. Radar studies of bird migration at the Northeast Passage and the Northwest Passage support the occurrence of migration along sun-compass routes in these polar regions. In contrast, satellite tracking of Brent geese (Branta bernicla) migrating from Iceland across Greenland and from Northwest Europe to Siberia show routes that conform most closely with geographic loxodromes, but which are also profoundly influenced by large-scale topography. These evaluations are discussed in relation to the adaptive values of different routes in different parts of the world. Sun compass routes are favourable mainly for east-west migration at high latitudes. For east-west migration at mid and high latitudes magnetic loxodromes are more favourable than geographic loxodromes in certain regions while the reverse holds in other regions. The geometry of migration routes, as recorded by radar and satellite tracking, may be important for understanding the evolution of the complexity of birds' orientation systems, and for providing clues about the orientation mechanisms guiding the birds on their global journeys

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 Eight 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 bring 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' Right power margin

Harmonic oscillatory orientation relative to the wind in nocturnal roosting flights of the swift Apus apus.

Swifts regularly spend the night flying at high altitude. From previous studies based on tracking radar observations, we know that they stay airborne during the night and prefer to orient themselves into the wind direction with an increased angular concentration with increasing wind speed. In this study, we investigated the orientation relative to the wind of individual swifts by frequency (discrete Fourier transform) and autocorrelation analysis based on time series (10s intervals) of the angle between the swifts' heading and the wind direction for radar trackings of long duration (9-60 min). The swifts often showed a significant harmonic oscillation of their heading direction relative to the wind, with a frequency mostly in the range 1-17 mHz, corresponding to cycle periods of 1-16 min. The swifts also sometimes performed circling flights at low wind speeds. Wind speed ranged from 1.3 to 14.8 m s(-1), and we expected to find different patterns of orientation at different wind speeds, assuming that the swifts adapt their orientation to avoid substantial displacement during their nocturnal flights. However, oscillatory orientation was found at all wind speeds with variable frequencies/periods that did not show any consistent relationship with wind speed. It remains to be shown whether cyclic heading changes are a regular feature of bird orientation

Migration Patterns of Tundra Birds: Tracking Radar Observations along the Northeast Passage

Bird migration was recorded by tracking radar and visual observations at 15 study sites, situated between 50°E and 170°E along the Northeast Passage, during a ship-based expedition in July and August 1994. A total of 1087 radar tracks (average duration 220 s) of bird flocks on postbreeding migration were recorded. Migration was dominated by waders and to a certain degree also skuas (especially pomarine skua Stercorarius pomarinus). Terns, gulls, ducks, and geese were also among the migrants tracked by radar. The radar data revealed a major migratory divide at about 100°E (Taymyr Peninsula), with mainly eastbound migration to the east of this divide, and mainly westbound migration to the west of it. The main stream of eastbound migration was directed toward the sector 90-120° and that of westbound migration toward the sector 240-270°; these directions are broadly in parallel with the coasts of the Arctic Ocean east and west of the Taymyr Peninsula, respectively. There was also important ENE migration, which provided strong indications of long-distance flights along orthodrome-like routes directly between Siberia and North America, across vast expanses of the Arctic Ocean pack ice. Analysis of flight directions in relation to wind indicated complete compensation for wind drift. Mean flight altitude was 1.3 km, and the birds regularly travelled at high altitudes above 3 km (9% of the tracks) up to a maximum height of 4.8 km. They preferred to migrate on occasions and at altitudes with following winds; such conditions provided an average gain in speed of 4.6 m/s. There were also recurrent cases of birds migrating in tailwinds of gale force, between 18 and 24 m/s. The birds' airspeed varied between 8 and 22 m/s, with a mean of 14 m/s. Airspeed was significantly correlated with altitude, wind, and vertical speed and seemed to be intermediate between the speeds for minimum power and maximum range predicted by aerodynamic theory.Durant une expédition ayant pour base un navire, réalisée en juillet et août 1994, on a enregistré la migration des oiseaux à l'aide d'un radar de poursuite et d'observations visuelles dans 15 zones d'étude situées entre 50 et 170° de longit. E., le long du passage du Nord-Ouest. On a enregistré un total de 1087 poursuites radar (d'une durée moyenne de 220 s) de volées d'oiseaux en migration après la nidification. La migration était dominée par les échassiers et, à un certain degré également, par les labbes (en particulier le labbe pomarin Stercorarius pomarinus). Les sternes, goélands, canards et oies étaient aussi au nombre des migrants suivis au radar. Les données de radar révèlent une scission migratoire majeure à environ 100° de longit. E. (presqu'île de Taïmyr), avec surtout une migration vers l'est, à l'est de cette division, et une migration vers l'ouest, à l'ouest de cette division. Le courant principal de la migration en direction de l'est était dirigé vers le secteur allant de 90 à 120° et celui de la migration en direction de l'ouest, vers le secteur allant de 240 à 270°; ces directions sont en gros parallèles aux rivages de l'océan Arctique à l'est et à l'ouest de la presqu'île de Taïmyr respectivement. Il y avait aussi une importante migration E.-N.-E., qui offrait une indication assez précise de vols de longue distance suivant un tracé ressemblant à la ligne orthodromique directe entre la Sibérie et l'Amérique du Nord, au travers de vastes étendues de la banquise de l'océan Arctique. L'analyse des directions de vol par rapport aux vents révèle une compensation totale pour la dérive due au vent. L'altitude moyenne de vol était de 1,3 km, et les oiseaux voyageaient régulièrement à altitude élevée, allant de 3 km (pour 9 p. cent des poursuites) à une hauteur maximale de 4,8 km. Les oiseaux préféraient parfois effectuer leur migration à haute altitude avec des vents arrière, de telles conditions offrant un gain de vitesse moyen de 4,6 m/s. On a observé des cas répétés d'oiseaux qui migraient avec un vent arrière soufflant en tempête, entre 18 et 24 m/s. La vitesse relative des oiseaux variait entre 8 et 22 m/s, avec une moyenne de 14 m/s. Cette vitesse relative était corrélée fortement avec l'altitude, le vent et la vitesse verticale, et semblait être intermédiaire entre les vitesses prédites par la théorie aérodynamique pour la puissance minimale et la portée maximale

Age-dependent migration strategy in honey buzzards Pernis apivorus tracked by satellite

Six adult and three juvenile honey buzzards Pernis apivorus were radio-tracked by satellite during autumn migration from southwestern Sweden. All adults crossed the Mediterranean Sea at the Strait of Gibraltar and continued across the Sahara desert to winter in West Africa, from Sierra Leone to Cameroon. Analysing three main steps of the migration, (1) from the breeding site to the southern Mediterranean region, (2) across the Sahara and (3) from the southern Sahara to the wintering sites, the adults changed direction significantly between these steps, and migrated along a distinct large-scale detour. In contrast, the juveniles travelled in more southerly directions, crossed the Mediterranean Sea at various places, but still ended up in the same wintering areas as the adults. Average speeds maintained on travelling days were similar for the two age groups, about 170 km/day in Europe, 270 km/day across Sahara and 125 km/day in Africa south of Sahara. However, as the adults used fewer stopover days en route, they maintained higher mean overall speeds and completed migration in a shorter time (42 days) than the juveniles (64 days). Although the juveniles set out on more direct courses towards the wintering grounds, they did not cover significantly shorter distances than the adults, as they tended to show a larger directional scatter between shorter flight segments. The results corroborate previous suggestions that adult and juvenile honey buzzards follow different routes during autumn migration, and that the birds change migration strategy during their lifetime. While juveniles may use individual vector orientation, social influences and learning may be of great importance for the detour migration of adults. The remarkable and distinct age-dependent shift in migratory route and orientation of the honey buzzard provides a challenging evolutionary problem

Compass orientation and possible migration routes of passerine birds at high arctic latitudes

The use of celestial or geomagnetic orientation cues can lead migratory birds along different migration routes during the migratory journeys, e.g. great circle routes (approximate), geographic or magnetic loxodromes. Orientation cage experiments have indicated that migrating birds are capable of detecting magnetic compass information at high northern latitudes even at very steep angles of inclination. However, starting a migratory journey at high latitudes and following a constant magnetic course often leads towards the North Magnetic Pole, which means that the usefulness of magnetic compass orientation at high latitudes may be questioned. Here, we compare possible long-distance migration routes of three species of passerine migrants breeding at high northern latitudes. The initial directions were based on orientation cage experiments performed under clear skies and simulated overcast and from release experiments under natural overcast skies. For each species we simulated possible migration routes (geographic loxodrome, magnetic loxodrome and sun compass route) by extrapolating from the initial directions and assessing a fixed orientation according to different compass mechanisms in order to investigate what orientation cues the birds most likely use when migrating southward in autumn. Our calculations show that none of the compass mechanisms (assuming fixed orientation) can explain the migration routes followed by night-migrating birds from their high Nearctic breeding areas to the wintering sites further south. This demonstrates that orientation along the migratory routes of arctic birds (and possibly other birds as well) must be a complex process, involving different orientation mechanisms as well as changing compass courses. We propose that birds use a combination of several compass mechanisms during a migratory journey with each of them being of a greater or smaller importance in different parts of the journey, depending on environmental conditions. We discuss reasons why birds developed the capability to use magnetic compass information at high northern latitudes even though following these magnetic courses for any longer distance will lead them along totally wrong routes. Frequent changes and recalibrations of the magnetic compass direction during the migratory journey are suggested as a possible solution

Are flight paths of nocturnal songbird migrants influenced by local coastlines at a peninsula?

By recording nocturnally migrating passerines with tracking radar we have investigated how coastlines affect the migrants’ flight paths. Birds could use coastlines as an orientation aid or as a reference cue to compensate for wind drift while migrating. However, on the small scale of Falsterbo Peninsula in southern Sweden, we found very little effect of coastlines on migrants flight paths, irrespective of altitude. We tracked 2 930 migrants in three autumn and two spring seasons, at altitudes from 60 up to 3 000 meters. We compared tracks of migrants flying in three different areas, which correspond to the three main coastlines, and can demonstrate that the orientation of the tracks did not differ in a way consistent with the coastlines between the areas in autumn, and showed only a slight effect in spring. This is in accordance with earlier infrared device monitoring in Falsterbo, but contrary to earlier visual observations. It supports the view of nocturnally migrating passerines as mainly broad-front migrants. Even though the coastlines on the scale of the peninsula affected the flight paths very little, it is possible that the coastline has an effect on a larger regional scale, by migrants avoiding long sea crossings and thereby being funneled towards the peninsula, but this remains to be investigated

Can vector summation describe the orientation system of juvenile ospreys and honey buzzards? - An analysis of ring recoveries and satellite tracking

Juvenile bird migrants are generally believed to use a clock-and-compass migratory orientation strategy. According to such a strategy migrants accomplish their migration by flying a number of successive flight steps with direction and number of steps controlled by an endogenous programme. One powerful way of testing this is by comparing predictions from a model of such a strategy with observed patterns. We used data from ringing and satellite-based radio telemetry to investigate the orientation system of juvenile ospreys (Pandion haliaetus) and honey buzzards (Pernis apivorus) migrating from Sweden to tropical west Africa. The ring recoveries showed a much larger scatter in the orientation of ospreys than of honey buzzards, but there was only a slight such difference in the satellite tracks. These tracks of individuals of both species were rather straight with a high directional concentration per step. The honey buzzard. data showed a close fit to a simple vector summation model, which is expected if birds follow a clock-and-compass strategy. However, the osprey data did not fit such a simple model, as ring recoveries showed a significantly greater deviation at short distances than predicted on the basis of long distance data. Satellite tracking also indicated less concentrated orientation on short distances. The pattern observed for the osprey can generally be explained by an extended vector summation model, including an important element of pre-migration dispersal. The existence of extensive dispersal in the osprey stands in contrast to the apparent absence of such dispersal in the honey buzzard. The explanation for this difference between the species is unclear. The model of orientation by vector summation is very sensitive to the existence of differences in mean direction between individuals. Assuming such differences, as tentatively indicated by the satellite tracking data, makes simple compass orientation by vector summation inconsistent with the distribution of ring recoveries at long distances, with a high proportion of misoriented birds falling outside the normal winter range

Migration Along Orthodromic Sun Compass Routes by Arctic Birds

Flight directions of birds migrating at high geographic and magnetic latitudes can be used to test bird orientation by celestial or geomagnetic compass systems under polar conditions. Migration patterns of arctic shorebirds, revealed by tracking radar studies during an icebreaker expedition along the Northwest Passage in 1999, support predicted sun compass trajectories but cannot be reconciled with orientation along either geographic or magnetic loxodromes (rhumb lines). Sun compass routes are similar to orthodromes (great circle routes) at high latitudes, showing changing geographic courses as the birds traverse longitudes and their internal clock gets out of phase with local time. These routes bring the shorebirds from high arctic Canada to the east coast of North America, from which they make transoceanic flights to South America. The observations are also consistent with a migration link between Siberia and the Beaufort Sea region by way of sun compass routes across the Arctic Ocean

Dark-bellied Brent Geese Branta bernicla bernicla, as recorded by satellite telemetry, do not minimize flight distance during spring migration

Nine Dark-bellied Brent Geese Branta bernicla bernicla were equipped with satellite transmitters during spring staging in the Dutch Wadden Sea in 1998 and 1999. The transmitters (in all cases less than 3% of body mass) were attached to the back by a flexible elastic harness. One juvenile female was tracked to the Yamal peninsula in 1998. Eight adult males were selected from a single catch of 75 to span the range of body mass observed on the date of capture (11 May 1999) and all but the lightest individual completed the first lap of the migratory flight to the White Sea, Russia, according to the time schedule normal for this species. Six birds were successfully tracked to Taymyr for a total distance averaging 5004 km (range 4577-5164) but judging from later movements none bred (although 1999 was breeding year). Although the routes chosen during spring migration were closely similar; none of the tagged birds migrated together. On average the geese used 16 flights to reach their summer destinations on Taymyr. The longest uninterrupted flights during the first half of the journey (Wadden Sea to Kanin) covered 1056 km (mean of seven adult males, range 768-1331), while the corresponding value for the second half of the migration (Kanin-Taymyr) was only 555 km (mean of six adult males). Only 7% of total time during spring migration was spent in active flight, as contrasted to c. 80% at long-term stopovers. Overall average travelling speed was 118 km/day (range 97-148). Including fattening prior to departure the rate of travel falls to 62 km/day (range 49-70), in keeping with theoretical predictions. Routes followed deviated from the great circle route, adding at least 700 km (16%) to the journey from Wadden Sea to Taymyr, and we conclude that the coastal route is chosen to facilitate feeding, drinking and resting en route instead of minimizing total flight distance