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     

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.     

Orientation of juvenile barn swallows (<Emphasis Type="Italic">Hirundo rustica</Emphasis>) tested in Emlen funnels during autumn migration

Although hirundines have been used extensively in homing experiments, to date no investigation of their migratory orientation has been carried out, despite the well-known migratory habits of many species of this family. This paper reports on a study of the orientation of the barn swallow (Hirundo rustica), a typical diurnal trans-Saharan migrant. Modified Emlen funnels were used to verify the suitability of this species for cage experiments and investigate the role of visual and magnetic cues during the birds first migratory journey. Juvenile swallows were mist-netted at a roost site in central Italy and then tested in a site 19 km apart. Orientation experiments were performed under four experimental conditions: natural clear sky and simulated overcast, in both local and shifted magnetic fields (magnetic North=geographical West). Under clear sky, the swallows tended to orient phototactically toward the best-lit part of the funnel and failed to respond to the magnetic field shift. Under overcast conditions, they oriented northward and modified their directional choices as expected in response to the shifted magnetic North. On the whole, our data indicate that swallows can use magnetic information for compass orientation. Possible explanations for the northward orientation of birds tested under overcast conditions are discussed.Communicated by W. Wiltschko     

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.     

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     

Re-orientation in clock-shifted homing pigeons subjected to a magnetic disturbance: a study with GPS data loggers

Some authors have proposed that homing pigeons are able to correct the error in orientation following a phase-shift treatment by using the magnetic compass reference. They reported that clock-shifted pigeons bearing magnets display a greater deflection compared to magnetically unmanipulated clock-shifted birds. However, this hypothesis tested by recording pigeons’ vanishing bearings has led to contradictory results. The present study reports pigeons’ tracks recorded with a GPS and shows that clock-shifted pigeons bearing magnets displayed a greater deviation through the whole route compared to the magnetically unmanipulated shifted pigeons. Moreover, the analysis of the tracks shows that the birds belonging to both experimental groups stop in coincidence with their subjective night. When re-starting their journey, the birds corrected the clock-shift induced error in orientation, but the magnetically manipulated pigeons were less efficient in doing so. Our results are consistent with the hypothesis that homing pigeons released from unfamiliar location re-orient after clock shift by using the magnetic compass.     

Experimental evidence of reciprocal altruism in the pied flycatcher

Although human behaviour abounds with reciprocal altruism, few examples exist documenting reciprocal altruism in animals. Recent non-experimental evidence suggests that reciprocal altruism may be more common in nature than previously documented. Here we present experimental evidence of mobbing behaviour, the joint assault on a predator in an attempt to drive it away, as reciprocal altruism in the breeding pied flycatcher (Ficedula hypoleuca). Given a choice, pied flycatchers assisted in mobbing initiated by co-operating neighbours and did not join in mobbing when initiated by conspecific neighbours which had defected from necessary assistance 1 h before. The results suggest the birds followed a ‘tit-for-tat’-like strategy and that mobbing behaviour of breeding birds may be explained in terms of reciprocal altruism.     

Prey size increases with nestling age: are provisioning parents programmed or responding to cues from offspring?

As offspring grow, parents often feed them with different sizes or taxa of prey to suit the changing nutritional or energetic demands. We investigated whether such changes in prey types were innate and inflexible or whether they were based on the proximate cue of offspring size. We created experimental broods in which parent blue tits Cyanistes caeruleus and pied flycatchers Ficedula hypoleuca received either older nestlings simulating rapid development or younger nestlings simulating delayed development. The size of prey increased over time in all brood types, suggesting a strong programmed pattern of foraging by parents. However, delayed broods were provisioned with smaller prey than control or advanced broods indicating some plasticity in the response by the parents to the size of nestlings. Although female birds brought smaller prey than male birds, both sexes showed the same rate of increase of prey size with time and brought similar types of prey items. The proportion of soft, digestible larva prey in the nestling diet decreased over time in pied flycatchers but increased in blue tits. Counter to the hypothesis that spiders provide unique and preferred nutrients for young nestlings, the proportion of arachnids in the diet did not change with nestling age for either species. The lack of treatment effect on the taxa of prey delivered suggests that temporal shifts in diet composition are not driven by the proximate cues of nestling age or size but that the feeding patterns are fairly innate and fixed in these altricial birds.     

Orientation of plaice (Pleuronectes platessa) in the open sea: evidence for the use of external directional clues

We studied the orientation of plaice (Pleuronectes platessa L.) in the open sea (Southern Bight of the North Sea) during May and June of 1979 and 1980, using continuous, on-line data from 6 fish equipped with transponding acoustic compass tags. During the day, fish swimming in midwater did not maintain their heading. At night, the same fish maintained their heading in a way that was consistent with active orientation to some external directional clue, but inconsistent with the use of an inertial guidance system. We believe this to be the first analysis to demonstrate the use of external guidance clues by fish in the open sea. Because fish were able to maintain a consistent heading in midwater by night, in the apparent absence of both visual and tactile clues, the external reference is likely to be geophysical. Possible clues include the electrical field generated by the flow of sea water through the geomagnetic field and the earth's magnetic field itself.     

Geomagnetic field affects spring migratory direction in a long distance migrant

Night-migrating song birds travel to and from their wintering and breeding areas often separated thousands of kilometers apart and are clearly capable of finding intended goal areas from a distant location. Displacement experiments provide a useful way to highlight orientation and navigational skills in migrants. To investigate which cues birds actually use to compensate for displacement and the exact mechanism of each cue, experiments with manipulation of single cues are required. We conducted a simulated displacement of lesser whitethroats Sylvia curruca on spring migration. Birds were displaced not geographically but in geomagnetic space only, north and south of their breeding area to test whether they incorporate information from the geomagnetic field to find their breeding area. Lesser whitethroats held in southeast Sweden but experiencing a simulated displacement north of their breeding area (Norway) failed to show a consistent direction of orientation, whereas birds displaced south of their breeding area (Czech Republic) exhibited consistent northerly orientation, close to the expected seasonally appropriate direction, after displacement toward the trapping location. The absence of a clear compensatory direction in birds displaced north might be due to unfamiliar magnetic information or lack of sufficient information such as a magnetic gradient when moving around. By isolating one orientation cue, the geomagnetic field, we have been able to show that lesser whitethroats might incorporate geomagnetic field information to determine latitude during spring migration.     

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     

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.     

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     

Temporal peaks in social information: prospectors investigate conspecific nests after a simulated predator visit

Individuals of many taxa gather social information in order to make informed decisions with lowered unpredictability. Social information may show temporal periods of higher information value during certain events. However, the value of information is expected to decrease with time since the event and individuals accessing this information are expected to do so quickly. In birds, prospectors visit the nests of other individuals to gather information about reproductive decisions and breeding success. Individuals are known to prospect even during their own breeding to access this information. We investigated prospecting behavior of pied flycatchers (Ficedula hypoleuca) at conspecific nests within a short time period following a simulated predator visit. We performed predator and control presentations at flycatcher nests, and recorded all prospectors (foreign conspecific individuals) visiting these nests within 45-min periods before and after the presentations. We found a significant increase in conspecific prospectors visiting focal nests after the presentation of a predator near the nest. The prospectors in our data comprised mainly pied flycatchers actively breeding in the neighborhood of the focal nest. We also found that the increase in prospecting at these nests is of short-duration, and that prospecting rates decreased back to background prospecting levels relatively quickly. Our results find support for the prediction that individuals may respond to temporally peaking value of information after certain observable events. This result suggests that prospecting may be especially aimed at specific periods of high information value.     

Low frequency of extra-pair paternity in pied flycatchers revealed by DNA fingerprinting

Summary Genetic parentage of 135 nestlings from 27 broods of polygynous and monogamous pied flycatchers Ficedula hypoleuca was analyzed by means of multilocus DNA fingerprinting. The minisatellite probe alpha-globin 3HVR detected approximately 12 scorable bands per fingerprint, and the proportion of bands shared between presumably unrelated adults averaged 0.22+0.08 SD. The fingerprints of 125 of the 135 nestlings made a complete match to those of their putative parents. In 4 nestlings a single mismatched band occurred, but since band sharing with both putative parents was high, the single mismatches were assumed to be caused by mutation. The 6 remaining nestlings had 5 or more mismatched bands each, low band-sharing proportions with their putative father and high band-sharing proportions with their putative mother. We thus conclude that they were all sired through extra-pair copulations (EPCs). Hence, only 4% of nestlings were sired through EPCs, and none resulted from intraspecific brood parasitism. One of the cuckolding males was identified, explaining all 5 mismatched bands in the nestling's fingerprint. Three of the illegitimate nestlings were from primary nests of polygynous males; 3 were from nests of monogamous males. The fact that many males in this study started to advertise for a second female in a distant territory several days before their first mate began egglaying, and still managed to secure almost exclusive paternity in their first brood, suggests that male polyterritoriality is not costly in terms of lost paternity. Common anti-cuckoddry tactics performed by male birds, like high rate of within-pair copulation and continuous mate-guarding thoughout the female's fertilizable period, do not seem to be important in pied flycatchers.Offprint requests to: J.T. Lifjeld     

Reduced costs of mixed-species pairings in flycatchers: by-product or female strategy?

Heterospecific matings are generally assumed to be unconditionally disadvantageous due to reduced viability or fertility of hybrid offspring. For female collared flycatchers (Ficedula albicollis) mated to male pied flycatchers (Ficedula hypoleuca), the cost of heterospecific pair formation is reduced due to high levels of conspecific extra-pair paternity and a male-biased offspring sex ratio. In order to investigate whether these cost-reducing mechanisms are the result of female mating strategies, rather than being a by-product of species incompatibilities, we manipulated the plumage of male collared flycatchers before pair formation to make them resemble male pied flycatchers. Since species incompatibilities are absent in this design, any systematic effect of manipulation on sex ratio or paternity would indicate a role of female mating strategy. Paternity was determined by means of a likelihood approach that controls the errors made in assigning a chick to be ‘within-pair’ or ‘extra-pair’. Neither the sex ratio nor the male share of paternity was affected by the manipulation in a systematic manner. We therefore conclude that our experimental data provide no support for the suggestion that female behavioural strategies are markedly adjusted in response to formation of mixed-species pairs. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Northern magnetic displacements trigger endogenous fuelling responses in a naive bird migrant

In a previous study, we found that juvenile northern wheatears (Oenanthe oenanthe) exposed to a magnetic displacement to the west of their natural migration route increased their body mass. The total intensity and inclination used for the western displacement may also have been interpreted as northern compared to the experimental site (stronger total field intensity and steeper inclination angle). In order to investigate whether the fuelling response was a response to an unexpected magnetic field or specific to the northern magnetic field, we conducted a new experiment. Juvenile wheatears from the same study population were magnetically displaced to southwestern magnetic fields, exposing the birds to unexpected magnetic combinations, but eliminating the possible effect of a northern magnetic field. A control group was kept in the local geomagnetic field in Sweden for comparison. There was no difference in body mass increase between treatments, suggesting that the fuelling response previously found was not a simple response to an unexpected magnetic field, but rather a specific response to the northern magnetic field. Juvenile wheatears may have developed a fuelling response to northern magnetic fields in order to enable a successful flight towards the migration goal.     

Deceptive behavior in pied flycatchers

Summary One explanation for polygyny in pied flycatchers (Ficedula hypoleuca) is the deception hypothesis, which proposes that females mate with already-mated males only because they are unaware that such males already have a mate on another territory. Recently this hypothesis has been criticized on the grounds that already-mated and unmated males differ sufficiently in their behavior that human observers can discriminate between the two classes fairly easily. Here we test whether male pied flycatchers change their behavior when visited by a female so as to make this discrimination more difficult. In our experiments we presented a caged female near the nestbox of an advertising male, in order to mimic the situation in which a female investigates the male and his nest site. During control periods, when only an empty cage was presented, we found that already-mated males were present on their secondary territories significantly less and sang significantly less than unmated males on their primary territories, confirming the earlier results. When we presented the stimulus female, all behaviors we measured changed significantly relative to control periods for both mated and unmated males: both classes of males increased the time spent on the territory, decreased singing rates, and increased various courtship behaviors. During the experimental periods there were no significant differences between mated and unmated males on any of the behavioral measures. Discriminant analysis was more successful in classifying males as to mating status using data from the control periods (without female) than using data from the experimental periods (with female). We conclude that when a female is present male pied flycatchers change their behavior in ways that make it more difficult to discriminate mated from unmated males.Offprint requests to: W.A. Searcy     

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.     

Orientation of homing pigeons at magnetic anomalies

Summary Young homing pigeons released at a site on the edge of a magnetic anomaly and then in the center of the anomaly show better orientation at the anomalous site than birds released there for the first time. To test the possibility that this improvement is the result of birds learning to obtain navigational information at magnetic anomalies, several groups of pigeons were trained at a series of different anomalies, in different directions from their home loft. When these birds were than tested at an unfamiliar anomaly they were disoriented. They showed no evidence of having learned to obtain navigational information at magnetic anomalies. It is suggested that the disorientation seen at anomalies may be due to a disturbance of position-fixing information at the release site.