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Many large marine vertebrates are today threatened by human activities and it is therefore crucial to obtain information on their distribution and behaviour at sea. In particular little is known about the time necessary for juveniles to acquire the foraging skills of adults. We tracked 13 juvenile wandering albatrosses Diomedea exulans by satellite telemetry during their first year at sea. They covered an average distance of 184,000 km during the first year and restricted their dispersal to the unproductive waters of the subtropical Indian Ocean and Tasman Sea. This region of low wind velocities does not overlap with the foraging areas used by adults. After an innate phase of rapid dispersal with a fixed flight direction, young birds progressively increased their daily flight distances and attained adult flight efficiency within their first six months at sea. The complete overlap of the juveniles' foraging ranges with major long-line fisheries in the subtropical waters constitutes a major threat that could jeopardize the long term recovery ability of populations of the endangered wandering albatross in the Indian Ocean.

Albatrosses are known for their extreme navigation performance enabling them to locate isolated breeding islands after long-distance migrations across open seas. Little is known about the migration of young albatrosses and how they reach the adults' navigation and foraging skills during the period of immaturity lasting several years and spent permanently flying across the open ocean. We tracked by satellite telemetry the dispersal and migration of 13 juvenile wandering albatrosses from the Crozet Islands during their first year at sea. The young albatrosses covered an average distance of 184,000 km during the first year, restricting their dispersal movement to the unproductive and low wind subtropical Indian Ocean and Tasman Sea. The juveniles initiated the migration by an innate phase of rapid dispersal encoded as a fixed flight direction assisted by southerly winds towards north and northeast. Thereafter each individual restricted its movement to a particular zone of the ocean that will possibly be used until they start breeding 7-10 years later and return in contact with breeding adults. This dispersal in young birds corresponds well with movements observed for adult non-breeding wandering albatrosses. The results show clearly an inherited ability to navigate back to already visited areas in young wandering albatrosses. The juvenile dispersal behaviour and migration at sea suggest a genetically based migration program, encoding navigation to a destination area used throughout the life.

Long distance migration has evolved in many organisms moving through different media and using various modes of locomotion and transport. Migration continues to evolve or become suppressed as shown by ongoing dynamic and rapid changes of migration patterns. This great evolutionary flexibility may seem surprising for such a complex attribute as migration. Even if migration in most cases has evolved basically as a strategy to maximise fitness in a seasonal environment, its occurrence and extent depend on a multitude of factors. We give a brief overview of different factors (e.g. physical, geographical, historical, ecological) likely to facilitate and/or constrain the evolution of long distance migration and discuss how they are likely to affect migration. The basic driving forces for migration are ecological and biogeographic factors like seasonality, spatiotemporal distributions of resources, habitats, predation and competition. The benefit of increased resource availability will be balanced by costs associated with the migratory process in terms of time (incl. losses of prior occupancy advantages), energy and mortality (incl. increased exposure to parasites). Furthermore, migration requires genetic instructions (allowing substantial room for learning in some of the traits) about timing, duration and distance of migration as well as about behavioural and physiological adaptations (fuelling, organ flexibility, locomotion, use of environmental transport etc) and control of orientation and navigation. To what degree these costs and requirements put constraints on migration often depends on body size according to different scaling relationships. From this expos it is clear that research on migration warrants a multitude of techniques and approaches for a complete as possible understanding of a very complex evolutionary syndrome. In addition, we also present examples of migratory distances in a variety of taxons. In recent years new techniques, especially satellite radio

Sea turtles are known to perform long-distance, oceanic migrations between disparate feeding areas and breeding sites, some of them located on isolated oceanic islands. These migrations demonstrate impressive navigational abilities, but the sensory mechanisms used are still largely unknown. Green turtles breeding at Ascension Island perform long oceanic migrations (>2200 km) between foraging areas along the Brazilian coast and the isolated island. By performing displacement experiments of female green turtles tracked by satellite telemetry in the waters around Ascension Island we investigated which strategies most probably are used by the turtles in locating the island. In the present paper we analysed the search trajectories in relation to alternative navigation strategies including the use of global geomagnetic cues, ocean currents, celestial cues and wind. The results suggest that the turtles did not use chemical information transported with ocean currents. Neither did the results indicate that the turtles use true bi-coordinate geomagnetic navigation nor did they use indirect navigation with respect to any of the available magnetic gradients (total field intensity, horizontal field intensity, vertical field intensity, inclination and declination) or celestial cues. The female green turtles successfully locating Ascension Island seemed to use a combination of searching followed by beaconing, since they searched for sensory contact with the island until they reached positions NW and N of the Island and from there presumably used cues transported by wind to locate the island during the final stages of the search.

Green turtles (Chelonia mydas) swim from foraging grounds along the Brazilian coast to Ascension Island to nest, over 2200 km distant in the middle of the equatorial Atlantic. To test the hypothesis that turtles use wind-borne cues to locate Ascension Island we found turtles that had just completed nesting and then moved three individuals 50 km northwest (downwind) of the island and three individuals 50 km southeast (upwind). Their subsequent movements were tracked by satellite. Turtles released downwind returned to Ascension Island within 1, 2 and 4 days, respectively. By contrast, those released upwind had far more difficulty in relocating Ascension Island, two eventually returning after 10 and 27 days and the third heading back to Brazil after failing to find its way back to the island. These findings strongly support the hypothesis that wind-borne cues are used by turtles to locate Ascension Island.

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

Magnetic compass orientation in birds has been shown to be light dependent. Results from behavioural studies indicate that magnetoreception capabilities are disrupted under light of peak wavelengths longer than 565 nm, and shifts in orientation have been observed at higher light intensities (43-44x1015 quanta s-1 m-2). To investigate further the function of the avian magnetic compass with respect to wavelength and intensity of light, we carried out orientation cage experiments with juvenile European robins, caught during their first autumn migration, exposed to light of 560.5 nm (green), 567.5 nm (green-yellow) and 617 nm (red) wavelengths at three different intensities (1 mW m-2, 5 mW m-2 and 10 mW m-2). We used monochromatic light of a narrow wavelength range (half bandwidth of 9-11 nm, compared with half bandwidths ranging between 30 nm and 70 nm used in other studies) and were thereby able to examine the magnetoreception mechanism in the expected transition zone between oriented and disoriented behaviour around 565 nm in more detail. We show (1) that European robins show seasonally appropriate migratory directions under 560.5 nm light, (2) that they are completely disoriented under 567.5 nm light under a broad range of intensities, (3) that they are able to orient under 617 nm light of lower intensities, although into a direction shifted relative to the expected migratory one, and (4) that magnetoreception is intensity dependent, leading to disorientation under higher intensities. Our results support the hypothesis that birds possess a light-dependent magnetoreception system based on magnetically sensitive, antagonistically interacting spectral mechanisms, with at least one high-sensitive short-wavelength mechanism and one low-sensitive long-wavelength mechanism.

Central-place foraging insects such as desert ants of the genus Cataglyphis use both path integration and landmarks to navigate during foraging excursions. The use of landmark information and a celestial system of reference for nest location was investigated by training desert ants returning from an artificial feeder to find the nest at one of four alternative positions located asymmetrically inside a four-cylinder landmark array. The cylindrical landmarks were all of the same size and arranged in a square, with the nest located in the southeast corner. When released from the compass direction experienced during training (southeast), the ants searched most intensely at the fictive nest position. When instead released from any of the three alternative directions of approach (southwest, northwest or northeast), the same individuals instead searched at two of the four alternative positions by initiating their search at the position closest to the direction of approach when entering the landmark square and then returning to the position at which snapshot, current landmark image and celestial reference information were in register. The results show that, in the ants visual snapshot memory, a memorized landmark scene can temporarily be decoupled from a memorized celestial system of reference.

We used radio-telemetry to study autumn migratory flight initiation and orientation in relation to wind and air pressure in a nocturnal passerine migrant, the reed warbler Acrocephalus scirpaceus at Falsterbo, southwest Sweden. The majority of the reed warblers departed in the expected migratory direction towards south of southwest, while a low number of the birds took off in reverse directions between north and east. Flight directions at departure correlated with wind directions. These correlations were particularly prominent at higher wind speeds but were absent at wind speeds below 4 m/s. Birds departing in the expected migratory direction compensated completely for wind drift. The reed warblers preferred to depart during nights with tailwinds and when air pressure was increasing suggesting that reed warblers are sensitive to winds and air pressure and select favourable wind conditions for their migratory flights. Since air pressure as well as velocity and direction of the wind are correlated with the passage of cyclones, a combination of these weather variables is presumably important for the birds' decision to migrate and should therefore be considered in optimal migration models.

We investigated the patterns of nocturnal bird migration in autumn 1998 at a coastal site on the Falsterbo peninsula in south-western Sweden, by means of a passive infrared device. In total 17 411 flight paths, including track direction and altitude, of migrating birds were recorded for 68 nights from August to October. Mean migratory traffic rate per night varied between 6 and 6618 birds km1 h1, with an average of 1319 birds km1 h1. Migration at Falsterbo showed a similar seasonal pattern to that reported for central Europe, with pronounced peaks of migration and intermittent periods with relatively low migratory intensities. Weather factors explained two thirds of the variance in the intensity of bird migration. During nights with intense migration, associated with weak winds, the mean track direction was close to that in central western Europe (225). Birds usually maintained a constant heading independent of wind directions and, in consequence, were drifted by the wind. The mean orientation clearly differed from that of the nearest coastline, suggesting that the birds did not use the topography below to compensate for wind drift.

Albatrosses and sea turtles are known to perform extremely long-distance journeys between disparate feeding areas and breeding sites located on small, isolated, oceanic islands or at specific coastal sites. These oceanic journeys, performed mainly over or through apparently featureless mediums, indicate impressive navigational abilities, and the sensory mechanisms used are still largely unknown. This research used three different approaches to investigate whether bi-coordinate navigation based on magnetic field gradients is likely to explain the navigational performance of wandering albatrosses in the South Atlantic and Indian Oceans and of green turtles breeding on Ascension Island in the South Atlantic Ocean. The possibility that magnetic field parameters can potentially be used in a bi-coordinate magnetic map by wandering albatrosses in their foraging area was investigated by analysing satellite telemetry data published in the literature. The possibilities for using bi-coordinate magnetic navigation varied widely between different areas of the Southern Oceans, indicating that a common mechanism, based on a bi-coordinate geomagnetic map alone, was unlikely for navigation in these areas. In the second approach, satellite telemetry was used to investigate whether Ascension Island green turtles use magnetic information for navigation during migration from their breeding island to foraging areas in Brazilian coastal waters. Disturbing magnets were applied to the heads and carapaces of the turtles, but these appeared to have little effect on their ability to navigate. The only possible effect observed was that some of the turtles with magnets attached were heading for foraging areas slightly south of the control turtles along the Brazilian coast. In the third approach, breeding female green turtles were deliberately displaced in the waters around Ascension Island to investigate which cues these turtles might use to locate and return to the island; the results suggested

The Earths magnetic field and celestial cues provide animals with compass information during migration.Inherited magnetic compass courses are selected based on the angle of inclination, making itdifficult to orient in the near vertical fields found at high geomagnetic latitudes. Orientation cage experimentswere performed at different sites in high Arctic Canada with adult and young white-crownedsparrows (Zonotrichia leucophrys gambelii) in order to investigate birds ability to use the Earths magneticfield and celestial cues for orientation in naturally very steep magnetic fields at and close to the magneticNorth Pole. Experiments were performed during the natural period of migration at night in the localgeomagnetic field under natural clear skies and under simulated total overcast conditions. The experimentalbirds failed to select a meaningful magnetic compass course under overcast conditions at themagnetic North Pole, but could do so in geomagnetic fields deviating less than 38 from the vertical.Migratory orientation was successful at all sites when celestial cues were available.

Satellite telemetry was used to record the submergence duration of green turtles (Chelonia mydas) as they migrated from Ascension Island to Brazil (N=12 individuals) while time/depth recorders (TDRs) were used to examine the depth distribution and dive profiles of individuals returning to Ascension Island to nest after experimental displacement (N=5 individuals). Satellite telemetry revealed that most submergences were short (<5 min) but that some submergences were longer (>20 min), particularly at night. TDRs revealed that much of the time was spent conducting short (24 min), shallow (approximately 0.91.5 m) dives, consistent with predictions for optimisation of near-surface travelling, while long (typically 2030 min), deep (typically 1020 m) dives had a distinctive profile found in other marine reptiles. These results suggest that green turtles crossing the Atlantic do not behave invariantly, but instead alternate between periods of travelling just beneath the surface and diving deeper. These deep dives may have evolved to reduce silhouetting against the surface, which would make turtles more susceptible to visual predators such as large sharks.

The C and N isotopes of feathers from two subspecies of willow warblers (Phylloscopus trochilus trochilus and Phylloscopus trochilus acredula) are isotopically distinct. Our analysis of 138 adult males from 14 sites distributed across Sweden shows that the mean delta(15)N and delta(13)C values of subspecies acredula (from latitudes above 63 degrees N) were significantly higher than the mean delta(15)N and delta(13)C values of subspecies trochilus (from latitudes below 61 degrees N). The analysed willow warbler feathers had been moulted in the winter quarters and the observed isotopic signatures should thus reflect the isotopic pattern of food assimilated in Africa. The isotopic data observed in Sweden match the dine in morphology, both showing abrupt changes around 62 degrees N. This result agrees with data from ringing recoveries indicating that the two subspecies occupy geographically and isotopically distinct wintering grounds in Africa. Our isotopic data suggest that analysis of stable isotopes of C and N is a promising method to track wintering quarters of European birds that migrate to Africa.

Green turtles (Chelonia mydas) that shuttle between their Brazilian feeding grounds and nesting beaches at Ascension Island in the middle of the Atlantic Ocean are a paradigmatic case of long-distance oceanic migrants. It has been suggested that they calculate their position and the direction of their target areas by using the inclination and intensity of the earths magnetic field. To test this hypothesis, we tracked, by satellite, green turtles during their postnesting migration from Ascension Island to the Brazilian coast more than 2000 km away. Seven turtles were each fitted with six powerful static magnets attached in such a way as to produce variable artificial fields around the turtle that made reliance on a geomagnetic map impossible. The reconstructed courses were very similar to those of eight turtles without magnets that were tracked over the same period and in the previous year, and no differences between magnetically disrupted and untreated turtles were found as regards navigational performance and course straightness. These findings show that magnetic cues are not essential to turtles making the return trip to the Brazilian coast. The navigational mechanisms used by these turtles remain enigmatic.

Among birds, falcons are high-performance flyers, in many cases adapted for aerial hunting and hence suitable targets for investigating limits to flight performance. Using an optical range finder, we measured flight tracks of Eleonoras falcon (Falco eleonorae), a species breeding in the Mediterranean region and specialised for hunting autumn passage bird migrants, when commuting between their nesting colony and offshore hunting areas (straight transportation flight) and when searching for prey (transecting and searching flight). Airspeed during searching flight was significantly slower than during straight transportation and transecting flight, but there was no significant difference in airspeed between the latter two flight modes. Straight transportation flight was significantly faster than predicted minimum power speed. Also, during straight transportation flight, the falcons responded to head- and tailwinds by increasing their airspeed when flying into the wind. However, they did not show any significant airspeed adjustments with respect to the angle between the track and the heading, as would be expected in birds trying to maintain a constant track direction. Mean sustainable climb rate (during (greater than or equal to) 240 s) was 1.4+/-0.31 m s-1 (mean +/- s.d., N=13), which is rather a high rate for a bird the size of an Eleonoras falcon. The climb rate was used to calculate maximum load-carrying capacity and maximum sustained horizontal flapping flight speed. The mean wingbeat frequency during powered climbing flight was 4.68 Hz, which was used to estimate the mass-specific muscle work. When falcons were leaving the colony for offshore hunting, they gained altitude by slope-soaring when there was an onshore wind. We formulated a simple criterion for the required gliding-flight rate of climb during an initial slope-soaring episode when minimizing the energy cost of reaching a certain altitude far out over the sea (which is where the prey is to be found). This