Orientation experiments with displaced juvenile barn swallows (Hirundo rustica) during autumn migratory season

In a previous paper, we reported that juvenile barn swallows (Hirundo rustica) trapped during the autumn migratory season and tested in modified Emlen funnels under overcast conditions can use magnetic information for compass orientation, although they did not orient toward the expected migratory direction. To further investigate this behaviour, in the present paper, we report the results of two series of displacement experiments performed on juvenile barn swallows belonging to two different roost sites [Massaciuccoli Lake (ML) and Terzo Cavone (TC)] and tested using the same protocol as that previously adopted. On the whole, the outcome of the ML and TC experiments was quite different with respect to (a) the degree of bird activity within the funnel (substantially lower for TC) and (b) the effect of the experimental displacement on the directional preferences of the tested subjects (not detectable in TC, while evident in ML swallows). Taken together, these results suggest that migrating swallows could react differently to the experimental tests depending on the roost site where they were caught. Indeed, TC swallows, trapped in southern Italy, close to the seacoast, and characterized by fairly large amounts of fat stores, mainly broke off their migratory activity or reoriented their directional preferences toward inland sites, possibly in search of more profitable habitats. On the other hand, ML swallows, trapped about 660 km to the north and characterized by lower amounts of fat stores, apparently tried to compensate for the experimental displacement.

Compensatory behaviour after displacement in migratory birds

Using a meta-analysis approach we re-analysed orientation cage experiments with displaced migrants found in the literature. A rather large proportion of the orientation experiments showed directional shifts after displacements, indicating ability for birds to detect and react on such displacements. There was a clear difference between overcast and experiments where birds had a view of the starry sky. In experiments under a starry sky, the birds compensated the displacements, whilst under overcast unaltered or reverse orientations were generally displayed. This indicates a role for the stars to be involved in detection of the changes in position. Such a role of celestial cues is further stressed by the results of several studies manipulating a planetarium sky. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Sunset orientation of robins,Erithacus rubecula,with different fields of sky vision

Summary Migratory orientation of robins (Erithacus rubecula) at sunset was recorded using orientation cages, under clear autumn skies. The aim of the experiments was to examine the importance of different visible sky sections for the orientation of robins. I obtained the following results: (1) Robins tested with the visible sky section limited to 90° around zenith (45° above the horizon) showed a mean orientation that coincided with the average sunset azimuth, with little scatter around the mean angle (Fig. 2). (2) When the birds were allowed a more extensive field of sky vision (maximum 160°), they chose headings on an approximate north-south axis, significantly different from tests with a restricted view of the sky (Fig. 3). (3) Experiments were also performed in which the response of robins to a mirror deflection (about 120° counterclockwise) of visual cues in the lower parts of the sunset sky was examined. The outcome indicated that visual information in the lower part of the sky may be critical for the orientation of robins (Fig. 5). These results, together with recent findings that robins captured and tested at two nearby sites show distinctly different orientation behavior in relation to experimental manipulations of the magnetic field, suggest that priorities among orientation cues may differ depending on the migratory situation encountered.     

Migratory orientation of red-eyed vireos,Vireo olivaceus,in relation to energetic condition and ecological context

How and when migrants integrate directional information from different sources may depend not only on the bird’s internal state, including fat stores, but also on the ecological context during passage. We designed experiments to (1) examine the influence of stored fat on the decision to migrate and on the choice of migratory direction and (2) investigate how the integration of orientation cue information is tied to energetic status in relation to migration across an ecological barrier. Migratory orientation of red-eyed vireos (Vireo olivaceus) at twilight was recorded using two different techniques, orientation cage experiments and free-flight release tests, during both fall and spring migration. During fall migration, the amount of stored fat proved decisive for directional selections of the vireos. Fat birds chose directions in accordance with migration across the Gulf of Mexico. Lean birds oriented either parallel to the coast line (cage tests) or moved inland (free-flight releases). Whereas only fat birds showed significant responses to experimental deflections of the geomagnetic field, lean birds displayed a tendency to shift their activity in the expected direction, making it difficult to evaluate the prediction that use of the magnetic compass is context dependent. Fat loads also had a significant effect on the decision to migrate, i.e., fat individuals were more likely to embark on migration than were lean birds (true for both cage and release experiments). During spring migration, a majority of experimental subjects were classified as lean, following their arrival after crossing the Gulf of Mexico, and oriented in seasonally appropriate directions. The vireos also showed significant responses to experimental deflections of the geomagnetic field regardless of their energetic status. Free-flight release experiments during spring migration revealed a significant difference in mean directions between clear sky and overcast tests. The difference may indicate a compensatory response to wind drift or possibly a need for celestial cues to calibrate the magnetic compass. Finally, this is the first demonstration of magnetic compass orientation in a North American vireo. Received: 15 December 1995/Accepted after revision: 24 March 1996     

Night sky orientation of migratory pied flycatchers raised in different magnetic fields

Summary To test whether the initial night sky orientation response of migratory pied flycatchers (Ficedula hypoleuca) is calibrated from the ambient magnetic field experienced by birds during their first summer, three groups of pied flycatchers were hand-reared and then held under different magnetic field conditions during the course of the summer. All groups were held outdoors and given full exposure to the day and night sky. One group was exposed to the local earth's magnetic field. A second group was exposed to a magnetic field of local earth strength, local earth inclination shifted 105° counter-clockwise relative to the local earth's field. The last group was exposed to a vertical, and thus nondirectional magnetic field.In autumn, the birds were tested for their orientation under the night sky in the absence of a directional magnetic field. When tested, all three groups were oriented with mean directions varying from south to southeast. No statistical differences emerged in any between group comparisons. The data indicate that earth's magnetism does not serve as a calibrating reference in the development of a pied flycatcher's initial orientation response to the night sky.     

Magnetic compass mediates nocturnal homing by the alpine newt, Triturus alpestris

Experiments were carried out to investigate the use of magnetic compass cues in the nocturnal homing orientation of the alpine newt Triturus alpestris. Tests were carried out at a site 9 km to the east–northeast of the breeding pond. Newts were tested at night in an outdoor circular arena that provided an unimpeded view of celestial cues, in one of four symmetrical alignments of an earth-strength magnetic field. In tests carried out under partly cloudy skies newts exhibited homeward magnetic compass orientation. Because the moon was visible in some trials, but obscured by clouds in others, we investigated whether the presence of the moon contributed to the scatter in the distribution of magnetic bearings. When the moon was visible, the distribution of magnetic bearings was more scattered than when the moon was obscured by clouds, although in neither case was the distribution significant due, in part, to the small sample sizes. Moreover, when the moon was visible, newts oriented along a bimodal axis perpendicular to the moon azimuth, suggesting that the presence of the moon may have affected the newts behavior. To provide a more rigorous test of the role of magnetic compass cues when celestial cues were unavailable, nocturnal tests were carried out during the following migratory season under total overcast. In the absence of celestial compass cues, the distribution of magnetic bearings exhibited highly significant orientation in the homeward direction. These findings indicate that newts are able to orient in the homeward direction at night using the magnetic compass as the sole source of directional information. Moon light altered the newts behavior. However, this apparently resulted from the asymmetrical distribution of moon light in the testing arena, rather than the use of an alternative compass.     

Clock-shift experiments with Savannah sparrows, Passerculus sandwichensis, at high northern latitudes

Orientation can be difficult for nocturnal bird migrants at high northern latitudes because of the large changes of magnetic declinations, rapid longitudinal time-shifts experienced during a long-distance flight and the invisibility of stars during the polar summer. Both sunset cues as well as geomagnetic cues have been shown to be of great importance in the orientation system of Savannah sparrows, Passerculus sandwichensis. We used clock-shift experiments to investigate whether geomagnetic and sunset cues were used for migratory orientation by wild-caught young Savannah sparrows at high geomagnetic latitudes in Northern Canada. We exposed birds to a 4-h slow clock-shift, expecting a 60° clock-wise shift in orientation after the treatment. Under natural clear skies in the local geomagnetic field, the birds responded by showing a significant axial mean orientation directed towards the position of the setting sun in the NW and towards their preferred migratory direction in the SE. After exposure to the clock-shift for 6 days and nights the birds showed a clear response to the treatment and shifted significantly towards NNE. Birds that first oriented towards NW in the experiments before clock-shift tended to shift clock-wise, thus reacted to the clock-shift in the expected way. The reaction of the individual birds that originally oriented towards SE seems to vary. In summary, our birds did not select a constant angle (menotaxis) in relation to the sun's position during the experiments, but presumably were affected by the sun showing phototaxis or followed their magnetic compass. Possible explanations of the unexpected experimental results are discussed. Electronic Publication     

Magnetic compass orientation in the yellow-faced honeyeater,Lichenostomus chrysops,a day migrating bird from Australia

Summary In Australia, the southern populations of the yellow-faced honeyeater, Lichenostomus chrysops (Meliphagidae), perform annual migrations, with routes following the eastern coastline. In order to assess the role of magnetic cues in the migratory orientation of this diurnal migrant, its directional behaviour was recorded in recording cages under natural and experimentally manipulated magnetic-field conditions. During autumn the birds tested indoors in the local geomagnetic field showed a directional change from north initially to northwest later in the season (Fig. 1 a, b), which corresponds well with the general pattern of movement of this species in the field. Deflecting magnetic north to ESE resulted in a clockwise shift of the mean direction by 77° and 71°, respectively (Fig. 1 c, d), while no significant directional tendencies were observed in a magnetic field with a compensated horizontal component (Fig. 1 e, f; see Table 1). In outdoor tests in spring, the birds preferred southerly directions when tested in the local geo-magnetic field. In a magnetic field with a reversed vertical component (i.e. with an inclination pointing down instead of upwards) the birds reversed their directional tendencies and oriented northward (Fig. 2, Table 2). These results clearly show: (1) that yellow-faced honeyeaters can use the magnetic field for direction finding, and (2) that their magnetic compass functions as an inclination compass, as has been shown for several holarctic migrants.Correspondence to: W. Wiltschko     

Influence of body condition and weather on departures of first-year European robins, <Emphasis Type="Italic">Erithacus rubecula</Emphasis>, from an autumn migratory stopover site

How migratory birds decide when to leave a stopover site is important to the understanding of bird migration strategies. Our study looks at how body condition and the weather affect the decision to depart on nocturnal migratory flight. During two autumn migration seasons (2002–2003), we radio tracked 51 first-year European robins, Erithacus rubecula, at a stopover site on the Courish Spit (Eastern Baltic) from the first day after landing until their migratory departure. The tagged robins stopped over for 1–14 days. There was no clear relationship between stopover duration and energetic condition on arrival. Weather conditions (wind, precipitation, and cloud cover) on departure differed measurably between years. In 2002, robins took off mainly under following winds and clear skies. In 2003, there were mainly light head winds and partially cloudy or overcast skies. This could be explained by the year-specific role of weather factors in making the decision to depart. In both years, robins making short (1–2 days) stopovers took off in more varied weather situations than those individuals with long stopovers. This suggests that robins from the former group were more inclined to continue with migration than longer-stay birds that, apart from re-fuelling, could be waiting for favourable weather. The lack of a relationship between stopover duration and body condition and some departures under unfavourable weather conditions suggest that endogenous spatiotemporal programmes may play an important role in controlling stopover duration in robins.     

The role of daytime cues in the development of magnetic orientation in a night-migrating bird

To assess the role of celestial rotation during daytime in the development of the magnetic compass course, pied flycatchers (Ficedula hypoleuca Pallas, Muscicapidae) were handraised in Latvia under various celestial and magnetic conditions. Tests were performed during autumn migration in the local geomagnetic field (50 000 nT, 73° inclination) in the absence of celestial cues. A group of birds that had never seen the sky showed a bimodal preference for the migratory southwest-northeast axis, whereas a second group that had been exposed to the natural sky from sunrise to sunset in the local geomagnetic field showed a unimodal preference for the seasonally appropriate southwesterly direction. A third group that had also been exposed to the daytime sky, but in the absence of magnetic compass information, also oriented bimodally along a southwest-northeast axis. These findings demonstrate that observing celestial rotation during daytime enables birds to choose the right end of the migratory axis for autumn migration at the Latvian test location. This transformation of axial behavior into appropriate migratory orientation, however, requires the birds to have simultaneous access to information on both celestial rotation and the geomagnetic field. Received: 19 September 1997 / Accepted after revision: 22 November 1997     

Magnetic orientation by hatchling loggerhead sea turtles (Caretta caretta) from the Gulf of Mexico

Loggerhead sea turtle hatchlings emerge from nests on either the east or west coast of the South Florida peninsula and then migrate offshore in opposite directions. Under laboratory conditions, magnetic cues induce east coast hatchlings to swim in directions that promote their transport by oceanic surface currents, such as the North Atlantic gyre. However, the surface currents used by west coast hatchlings are unknown. We examined the responses of west (Sarasota) hatchlings to magnetic cues in the Gulf of Mexico, the Florida Straits, and the Gulf Stream to determine their (1) likely migratory routes (2) orientation where currents lead into the Atlantic Ocean, and (3) orientation adjacent to Florida’s east coast. The results suggest that migration inside Gulf waters may be circuitous, that the turtles respond appropriately to enter Atlantic waters, and that orientation along Florida’s east coast probably promotes transport by the Gulf Stream into the North Atlantic gyre.     

Interaction between solar orientation and landscape visibility in Talitrus saltator (Crustacea: Amphipoda)

A series of experiments on the littoral amphipod Talitrus saltator (Montagu) was carried out between April and September, 1978–1981, both under natural conditions and inland with artificial landscapes of different heights on the horizon, in order to assess the visual importance of the landscape in zonal orientation in populations from the Mediterranean coast, and to determine interactions between solar orientation and orientation based on the landscape. Inland, orientation in controls (permitting only vision of the sky and the sun) was compared to that of the experimental individuals, who could see a simulated landscape positioned landwards to them and seawards. In nature, sandhoppers released in a level arena with the landscape screened from view were compared with others released in an unscreened arena and on the sand in absolutely natural conditions. Situations where solar orientation contradicted local cues were produced both by releasing the sandhoppers on a shore diversely orientated to their own, and by shifting their internal clock by nine hours. Results show that T. saltator uses the landscape as a cue in its orientation towards the sea, in conjunction with solar orientation: the latter being the principal factor involved even when the sandhoppers are separated from the local optical factor in the sky (in trials at a distance from the sea). In fact, with the artificial landscape set seawards, none of the populations we studied showed any orientation based on the landscape comparable in accuracy to orientation based exclusively on the sun. From the experiments carried out in natural conditions, it is possible to deduce that conflicting conditions gave rise to two types of results: deviation from the mean direction and an increase in dispersion, up to total dispersion when the local factors were in total contrast to solar orientation (clock-shifted sandhoppers released on the sand).     

Orientation in captive migratory and sedentary Australian silvereyes <Emphasis Type="Italic">Zosterops lateralis</Emphasis> (Zosteropidae)

The orientation of two closely related subspecies of Australian silvereye Zosterops lateralis was studied in captivity over 14 months. Migratory silvereyes Z. lateralis lateralis showed significant directional preferences during the spring and autumn migration periods and also displayed orientated behaviour during the breeding period. In contrast, the non-migratory subspecies Z. lateralis familiaris did not display any significant directional preferences at any time. This is the first time that the orientation behaviour of a migratory and non-migratory subspecies has been compared over the duration of an annual cycle, both during and outside the migratory periods. The results suggest that migratory silvereyes possess an endogenous program determining the timing and direction of autumn and spring migration and that this program is unique to the migratory subspecies. This is also the first comparison of this nature on southern hemisphere birds and demonstrates that the migratory behaviour of southern hemisphere migrants may be more similar to that of northern hemisphere migrants than previously thought.Submitted to Behavioral Ecology and Sociobiology: 23 Jan 2006.     

Sunset and the orientation of a nocturnal bird migrant: A mirror experiment

Summary If savannah sparrows, (Passerculus sandwichensis), a North American night migrant, select a migratory heading based upon the setting sun, a shift in the position of that cue should produce a predictable shift in the migrant's nocturnal orientation. I tested this hypothesis by shifting the sunst position with mirrors and by recording the bird's orientation in Emlen funnels. The control group displayed directionality appropriate for spring migration (=342°). The mean heading of experimentals (=272°), which were exposed to a cue-shifted situation, was in the expected westerly direction relative to the control mean (P<0.05, V-test). The setting sun appears to be a sufficient source of directional information for this avian migrant.     

Corticosterone levels in relation to migratory readiness in red-eyed vireos (<Emphasis Type="Italic">Vireo olivaceus</Emphasis>)

We examined the relationship between plasma levels of corticosterone and the migratory activity and directional preference of red-eyed vireos during fall migration at the northern coast of the Gulf of Mexico. Corticosterone is thought to play a role in physiological and behavioural processes before, during, and after long-distance migratory flights. An increase in corticosterone at the onset of migratory flights can be expected in birds that are energetically prepared to migrate in a seasonally appropriate southerly direction. Red-eyed vireos ( Vireo olivaceus) were tested in orientation cages under clear twilight skies. Just prior to the orientation experiments, blood was sampled to assay baseline corticosterone levels. Average corticosterone level for all birds was 22.8 ng/ml. Red-eyed vireos with higher than average baseline levels of corticosterone were significantly more active in orientation cages compared to birds with lower levels of corticosterone. Moreover, birds with higher than average levels oriented in a southwesterly direction, which is consistent with a trans-Gulf flight, whereas individuals with levels below average showed a NNW mean direction. Although there was no significant difference in baseline levels of corticosterone between fat and lean birds, individual mass loss between capture and test was negatively correlated with corticosterone levels. Results from this study clearly demonstrate that corticosterone influences departure decisions and the choice of direction during migration.     

Orientation of the pied flycatcher Ficedula hypoleuca: cue-conflict experiments during spring migration

Hierarchical relationships among different compass systems in long-distance migrants are still a matter for discussion because different studies have led to highly variable and apparently contradictory results. We carried out cue-conflict experiments during spring migration on pied flycatchers Ficedula hypoleuca (Passeriformes, Muscicapidae). Birds were exposed to a conflict between celestial and magnetic information by altering the polarized light pattern or magnetic field. The polarization pattern was shifted (±90°) with filters, whereas the magnetic field was altered (+90°) through Helmholtz coils. Birds were tested in modified Emlen funnels both before and after the cue conflict; during the tests, only the natural magnetic field was available. This protocol was designed to test whether the experimental birds recalibrated their magnetic compass on the directional information derived from the light polarization pattern when the region near the horizon was visible during the conflict. Contrary to this expectation, we did not record any significant shift in magnetic orientation after one or repeated exposures to the cue conflict. Our results support earlier studies, which suggest that the magnetic field is the primary compass cue during the migratory period.     

The magnetic compass mechanism of birds and its possible association with the shifting course directions of migrants

Summary Many birds of the northern hemisphere shift their migratory course to more southerly directions when moving from northern to southern latitudes. Birds from Central Europe, for example, change their course from SW to S or from SE to S respectively (Fig. 1). This also seems to apply to some other animals.The hypothesis presented here explains the observed shifts in migratory direction on the basis of changes in the parameters of the earth's magnetic field and hence would make a genetic fixation of shifts in the migratory direction unnecessary.To determine the direction of migration birds do not refer to the polarity of the magnetic field but to its dip (=). According to the hypothesis presented here, the birds, however, do not refer to the direction of dip as previously believed but to the individual apparent angle of dip (=), this angle changes depending on the heading of the bird (see Fig. 3 and Eq. 1). Maintaining a species specific or population specific the bird will move in its predetermined migratory direction. Changes in the dip of the earth's magnetic field correspond to changes in latitude. According to the hypothesis with fixed, the migratory direction will change when the dip changes. Given the hypothesis and the parameters of the earth's field theoretical migratory paths of birds between summer and winter quarters may be calculated (Figs. 8–11). The calculated tracks and the actually observed migratory routes agree well. This is also confirmed by radar and other observations of migratory directions in areas of different dip angles (Fig. 13). Displacing migrating birds to areas of smaller dip angles (= lower magnetic latitudes) results in predeterminable shifts in the birds migratory direction (Figs. 5, 6). The hypothesis also accounts for the so far unexplained orientation behaviour of transequatorial migrants under the magnetic equator.A very simple model of this hypothetical compass mechanism may be based on the assumption of the sensor axis is supposed to correspond to the apparent angle of dip when moving in the migratory direction. In this position the difference between the apparent angle of dip and the angle of the sensor is zero. Any change in the direction of movement, however, will result in a difference leading to a response of an assumed receptor. When maintaining the zero difference the bird invariably sticks to its migratory course. The proposed mechanism is a null instrument unaffected by changes in field intensity and not depending on the measurement of absolute values.     

Multiple regression analysis of the sources of variation in orientation of two sympatric sandhoppers, <Emphasis Type="Italic">Talitrus saltator</Emphasis> and <Emphasis Type="Italic">Talorchestia brito</Emphasis>, from an exposed Mediterranean beach

Two sympatric species from an exposed sandy beach in north-western Tunisia, the sandhoppers Talitrus saltator (Montagu) and Talorchestia brito Stebbing, were compared as regards orientation under natural conditions. Sets of experiments were carried out on the beach using two experimental arenas, one permitting the view of both the sky and landscape and the other of the sky only. Replicates were made at different hours of the day, in different days and period of the year and with various climatic conditions. Multiple regression models were fitted to the angular data obtained. These models allow an estimate of the trends of variation depending on a combination of factors. The simultaneous effects of environmental factors influencing orientation were quantified for each species. The sun was confirmed as the major orientation cue in these populations, but the concentration of directional choices was enhanced by the landscape vision. The climatic factors affected orientation in different ways depending on the time of day and year. When the two species were compared, Talitrus saltator showed a higher flexibility of response than Talorchestia brito under the same environmental conditions, supporting the hypothesis of a higher level of terrestrialization of the former species.     

Migratory orientation of juvenile yellow-rumped warblers (Dendroica coronata) following stopover: sources of variation and the importance of geographic origins

Birds migrating along coastlines may be at increased risk if displacement occurs toward open-ocean. Eastern North America experiences prevailing northwesterly winds during autumn, which could compel some migrants to drift eastward. Therefore, migrants at stop-over sites along this route may be a mixture of on- and off-course individuals. We assessed whether orientation behavior of juvenile yellow-rumped warblers (Dendroica coronata) captured at a stop-over site in southern Nova Scotia was related to where they originated from (i.e., likely on/off-course). We hypothesized three scenarios after displacement: 1) continued orientation in the migratory direction selected before displacement, 2) orientation from the new location toward the previous destination, or 3) correction to regain the original pathway. Using stable isotopes, we determined that stop-over migrants originated from nearby areas (and assumed on-course) and as far away as western Hudson Bay, over 1,600 km northwest (and assumed off-course) of the site. We used video-based orientation registration cages to determine an individuals’ migratory orientation. Because numerous factors influence migratory orientation (e.g., fuel reserves, celestial cues), we simultaneously assessed the influence of body condition and cloud cover, in addition to geographic origin, on orientation behavior. Individuals that originated closer to the site tended to orient more southwesterly. Orientation directions became increasingly more west-northwesterly the further away an individual originated from the site (i.e., the more likely it was to have been displaced). The result is most consistent with scenario three: individuals from northwest origins likely respond to easterly displacement by orienting westerly to reestablish their previous migratory route.     

Migratory and reproductive movements of male leatherback turtles (<Emphasis Type="Italic">Dermochelys coriacea</Emphasis>)

The biology of the endangered leatherback turtle (Dermochelys coriacea) at sea is poorly understood. As research has been almost exclusively limited to studies of nesting females, the movements of male turtles and the timing and location of mating activity have remained unknown. We report on the first deployments of satellite tags on male leatherbacks. Male migration to and residency in waters adjacent low-latitude nesting beaches in the western Atlantic suggest that this is where mating occurs, and return migration to these areas reveals male fidelity for breeding sites. Rate of travel decreased markedly after arriving in coastal breeding areas, where males remained for up to 96 days before assuming northward migrations. The initiation of these northward migrations coincided with peak nesting activity in adjacent nesting colonies. Data from satellite-linked time-depth recorders attached to two males revealed diel dive patterns in breeding areas and marked differences in diving behaviour between migratory and breeding periods in one turtle. When male turtles were in waters adjacent nesting colonies, their movements differed from those reported for nesting females, with females ranging farther from shore. Our results suggest that male leatherbacks may be vulnerable to entanglement in coastal fishing gear in waters adjacent nesting beaches.Communicated by R.J. Thompson, St. Johns