Pigeon homing: Olfactory orientation—a paradox

Summary In order to find out whether the different ways that pigeons are raised and maintained at the various lofts affect their orientation behavior, especially the selection of navigational factors, a group of birds was raised according to the procedures of our Italian colleagues in a wind-exposed loft on the roof. The behavior of these R-birds was then compared with that of G-birds living in a garden loft, raised and trained according to the normal Frankfurt procedure. When R-birds were made anosmic by closing the nostril with cotton during transportation and a local anesthetic was used at release, their reaction was similar to that of Italian pigeons: the deviation of their vanishing bearings from the home direction increased significantly, leading to a marked decrease in homeward orientation. In contrast, the orientation of the anosmic G-birds did not differ from that of their controls; their directional selections agreed with those of the controls of the R-group. These data indicate that the conditions of raising and maintaining homing pigeons may be of crucial importance in determining the pigeons' attitude toward olfactory input. Finally, olfactory orientation is discussed; the paradoxical finding that the G-birds, not using olfaction, oriented like the controls of the R-group that did use olfactory input, leads to the question of whether olfactory input really conveys navigational information to the birds.     

Short-term residence in deflector lofts alters initial orientation of homing pigeons

Summary A modification of the deflector-loft technique first outlined by Baldaccini et al. (1975) is presented in which experienced homing pigeons that do not permanently reside in deflector lofts were housed in them for periods of 7–20 days. Upon release these birds consistently exhibited a deflection of mean vanishing bearings in the directions predicted by the olfactory hypothesis of pigeon homing. Two potential explanations for this short-term deflector-loft effect are suggested. One is that the olfactory map sense of homing pigeons is very flexible and capable of accurate readjustment in as short a period as seven days. Alternatively, it may be that nonolfactory cues are being altered by the deflector lofts in such a way as to result in behavior by pigeons that is consistent with the olfactory hypotheses. The short-term technique has the practical benefit of making it possible to conduct far more experiments in a single field season than was possible with the original deflector-loft method.     

Can pigeons be fooled about the actual release site position by presenting them information from another site?

Summary The aim of the experiment was to test the hypothesis that pigeons depend on route- and/or site-specific airborne parameters to establish their position relative to the loft. Pigeons were transported to the release site with free access to the environmental air. They were then enclosed in large airtight containers filled with air from the release site and either transported via the loft to a release site in the opposite direction with respect to the loft (experimental birds) or via the loft back to the same site (control birds). Before release the pigeons (still inside the containers) were made anosmic by a local anesthetic applied to the olfactory membranes through the nostrils. Vanishing bearings of experimental birds were on the average in a direction opposite to home while the vanishing bearings of control birds were homeward oriented. For this initial orientation the pigeons seem to rely on airborne cues obtained at a site where they last had access to ambient air or cues obtained from the air inside the sealed containers. Irrespective of the treatment the bearings pooled with respect to north still show a residual orientation to the NE. There was no difference either in the homing speeds or in the homing times of anosmic control and experimental pigeons. We therefore have to assume further mechanisms guiding the pigeons home in addition to a possibly olfactory one.     

Pigeon homing: Different effects of olfactory deprivation in different countries

Summary This study compares the orientation of untreated pigeons and pigeons subjected to olfactory deprivation at two lofts near Pisa, Italy, at a loft at Ithaca, New York, USA, and at a loft at Frankfurt a.M., FRG. The experimental birds were rendered anosmic by nasal plugs until Gingicain, a local anaesthetic, was applied shortly before release. The Italian and American control pigeons appeared to orient towards home equally well, while the control pigeons in Germany frequently preferred directions that deviated significantly from the home direction. The effect of olfactory deprivation was small in the USA and in Germany; it was significantly larger in Italy, indicating that Italian pigeons depend on olfactory information to a much greater extent. These findings suggest that there are important regional differences in the strategies and cues pigeons use to navigate. The varied roles of olfactory information, and the reasons for these differences are discussed.     

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.     

Correct and false olfactory orientation of homing pigeons as depending on geographical relationships between release site and home site

Summary Pigeons from two German home sites were released at a site near Mantua in northern Italy. The home sites, Andechs and Würzburg, are 303 and 508 km north of the release site, respectively. Not only the initial bearings but still more the distributions of recoveries after a longer flight distance (median 65 km) were very different in pigeons from these two lofts. While the majority of the Wurzburg birds were found north of the release site, almost all birds from Andechs were found south of it (Fig. 1). Pigeons from both lofts, if made anosmic by sectioning the olfactory nerves, showed no average tendency towards change of latitude. These findings strongly suggest that both correct and false positional information were deduced by the birds from olfactory inputs. A coherent (though very hypothetical) interpretation of these and earlier results is based on regularly varying proportions of chemical tract compounds in the atmospheric boundary layer over the Alps and adjacent regions (Fig. 4).     

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.     

Spatio-temporal use of the urban habitat by feral pigeons (Columba livia )

Feral pigeons are descendants of wild rock pigeons that have adapted to the urban habitat. They have partially conserved the foraging behaviour of their wild ancestors (flights to agricultural areas) but have also developed new habits. Previous studies on the foraging strategies of feral pigeons have given various results, e.g. maximum distances reached by the pigeons (measured in a straight line from the resting places) differed between 0.3–0.5 km and 18–25 km. This study focuses on the spatio-temporal activity of feral pigeons in the urban habitat. We equipped 80 free-living feral pigeons from Basel, Switzerland with GPS receivers. We found three different foraging strategies for pigeons in Basel: (1) in the streets, squares and parks near the home loft, (2) in agricultural areas surrounding the city, (3) on docks and railway lines in harbours. The maximum distance reached by a pigeon was 5.29 km. More than 32% of the pigeons remained within 0.3 km of the home lofts and only 7.5% flew distances of more than 2 km. Females covered significantly longer distances than males, preferring to fly to more abundant and predictable food sources. Temporal activity patterns showed to be influenced by sex, breeding state and season. In contrast to wild rock pigeons and to feral pigeons in other cities, pigeons in Basel showed a clear bimodal activity pattern for breeding birds only. The differences between our results and those of other studies seem to be partly method-dependent, as the GPS-technique allows to record the pigeons’ localisations continuously in contrast to other methods. Other differences might be due to different kinds of food supply in the various cities. Our study shows that feral pigeons have individual foraging strategies and are flexible enough to adapt to different urban environments.Electronic supplementary material Supplementary material is available for this article at     

Pigeon homing in relation to geomagnetic,gravitational, topographical,and meteorological conditions

Summary Experienced homing pigeons were released at sites unfamiliar to them and with magnetic and gravity anomalies as well as in areas with rather normal fields throughout the FRG (41 releases when sunny, 14 when overcast; Figs. 1–3). The second-order release data were subjected to both univariate and multivariate statistical analysis (stepwise regression, factor analysis). The magnetic field strength and its gradients within the 1-km circle around the release site were determined from aeromagnetic maps of the anomalies of magnetic total intensity. Analogous variables were derived from gravity anomaly maps. It was tested whether the pigeons fly along that gradient to minimize the difference between the magnetic or gravity field at the release site and the loft at maximum rate. Further independent variables described magnetic K index, day-to-day variations of the magnetic components, topography, meteorological conditions, the number of the releases the pigeons had done, and the distance.Over magnetic anomalies widely varying in strength (departing – 250 nT to 300 nT from normal 600 m above ground), extent, and distance from loft, the pigeons vanished with less deviation from the homeward direction and faster than they did in areas with less irregular fields under sunny conditions; this is in contrast to other studies on magnetic anomalies, except one. At sites of gravity anomalies (15–49 mgal), the pigeons were significantly less homeward oriented and homed slower than at less anomalous sites (–9 to 14 mgal).Variables related to gravity were best predictors in 8 and and second predictors in 3 out of 15 regression analyses of the navigational parameters for the releases under sun. Six times the (absolute) amount of the gravity difference between the release site and the loft was selected first (Figs. 7B, C, 8B, C). The results suggest gravity to be involved in navigation as the pigeons' distance measure. Homeward directedness declined with increasing amount of the gravity gradient in the first 12 releases under sunny skies as well as when overcast (Fig. 8A). A preferred compass direction towards north-northeast was determined, being closest to the grand mean vector of the ascending gravity gradient (Fig. 6). The analyses failed to show directional preferences as assumed by the hypotheses tested. Temperature and degree of cloud cover provided some information for predicting mean vector lengths and mean vanishing times, respectively. The surface wind component in the homeward direction was correlated with median homing performances.     

Navigational strategies of homing pigeons at familiar sites: do landmarks reduce the deflections induced by clock-shifting?

To analyze the navigational strategy of homing pigeons at familiar sites in view of a possible role of local landmarks, two groups of pigeons—one familiar to the release site, the other unfamiliar—were released with their internal clock shifted 6 h fast, with untreated birds of both groups serving as controls. The two groups showed median deflections of 67% and 57%, respectively, of the expected size, with no consistent difference in the size of the deflection between familiar and unfamiliar birds. This clearly shows that familiarity with the release site and with the local landscape features does not affect the size of the deflections induced by clock-shifting. Obviously, pigeons familiar with the release site do not change their navigational strategy, but still continue to determine their home course solely as a compass course. General problems with orientation by landmarks are discussed; however, landmarks may help birds to recognize a site and recall the respective course.

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Geographical and temporal variability in pigeon homing

Summary Young homing pigeons from the same German stocks were housed in two lofts, one in southern Germany, near Munich, and one in Italy, near Pisa. In the course of 1 year, two synchronized releases at sites 22–25 km NNW and SSE from each of the lofts were conducted every month. The pigeons that returned were released a second time at a site about 75 km east of home.Both initial homeward orientation and homing success were considerably better in Italy than in Germany. Annual cycles, with maxima in summer and minima in winter, were observed in both countries. They were most pronounced in initial orientation in Italy and in homing performance in Germany. Correlations between homing parameters and ambient temperature correspond to the annual cycles, but they do not indicate that the geographical and seasonal differences in homing behaviour are directly caused by actual temperature at the time of release.Our findings (together with earlier ones) suggest that environmental conditions may be variably conducive to the homeward orientation of pigeons according to spatial and temporal variations in the climate.     

The air speed of migrating birds and its relationship to the wind

Summary This paper deals with the flight speeds of migrating birds observed by radar over a Swiss Alpine pass. The distributions of air speeds for different classes of birds and the influence of the wind on air speeds were investigated. Our findings differ greatly from observations of bird migration over the North Atlantic Ocean and the North American continent. Our data reveal: (1) that the air speeds of the bulk of migrating birds were in the range of 8–18 m/s and that the amount of slow flyers (air speed below 5 m/s) was less than 5%; (2) that there was an obvious influence of the wind on air speeds and, moreover, the data showed a distinctly variable compensatory behavior among different bird classes; (3) that the wind component in the direction of the bird's heading was a better predictor for air speed than the wind direction. Although we do not speculate about possibilities and mechanisms of wind detection, a simple argument shows how birds could estimate wind directions if they do use the moving surface as a reference system.     

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 behavior of homing pigeons at the Gernsheim anomaly

Pigeons were released at four release sites within the Gernsheim anomaly, a magnetic 'hill' with a peak 199 nT above the regional reference field and gentle 'slopes' to all sides, situated 44 km south of the Frankfurt loft. Local magnetic conditions at the sites differed in total intensity and in direction and steepness of the intensity gradient. At all sites, the pigeons were well oriented, showing counterclockwise deviations from the home directions that were most pronounced in the western part of the anomaly. There was no systematic difference in orientation behavior or homing performance between the sites within the anomaly and a control site outside. No effect of the local gradient direction was found, nor did the difference in intensity between home loft and the release site affect behavior. This argues against the use of magnetic navigational factors. However, pigeons released for the first time within the anomaly tended to have longer mean vectors with increasingly steeper gradients, which could mean that the birds might somehow have realized the anomalous nature of the local magnetic conditions and ignored them, relying on non-magnetic cues instead.Communicated by R. Gibson     

Homing behaviour in the sleepy lizard (Tiliqua rugosa): the role of visual cues and the parietal eye

To investigate how visual cues are integrated into a navigational strategy for homing in the Australian sleepy lizard (Tiliqua rugosa), lizards were displaced beyond their home range, either with full access to visual cues or with no access to visual cues during the displacement. Homeward orientation was significantly worse when lizards were denied visual cues during the displacement than when they were not. However when lizards were displaced with their field of view restricted to the sky, their homeward orientation was equally as good as that of lizards displaced with no visual restriction. These experiments suggest that sleepy lizards use celestial cues to determine the compass bearing of the outward journey, and reverse this bearing to orient in the homeward direction (course reversal). In a subsequent experiment, lizards oriented randomly with respect to home when the parietal eye was entirely covered with a patch during the displacement and return, while control lizards fitted with a sham parietal eye patch were well oriented towards home. In both groups, the lateral eyes were unobstructed and had complete access to visual cues including celestial cues and landmarks. These results suggest that the parietal eye plays a highly significant role in sleepy lizard homing, perhaps mediating a sky polarization compass sense.     

Wind selectivity of migratory flight departures in birds

Optimal migration theory predicts that birds minimizing the overall time of migration should adjust stopover duration with respect to the rate of fuel accumulation. Recent theoretical developments also take into account the wind situation and predict that there is a time window (a set of days) during which birds should depart when assisted by winds but will not do so if there are head winds. There is also a final day when birds will depart irrespective of wind conditions. Hence, the wind model of optimal migration theory predicts that birds should be sensitive to winds and that there should be a correlation between departures and winds blowing towards the intended migration direction. We tested this assumption by tracking the departures of radio-tagged passerines during autumn migration in southern Sweden. Our birds were moderately to very fat when released and therefore energetically ready for departure. There was a significant correlation between direction of departure and wind direction. We also found that during days when birds departed there was a significantly larger tail wind component than during days when birds were present but did not depart. Our results show that passerines do take the current wind situation into account when departing on migratory flights. We also briefly discuss possible clues that birds use when estimating wind direction and strength. The inclusion of wind is an important amendment to optimal migration theory of birds and should be explored further. Received: 1 March 1999 / Received in revised form: 4 October 1999 / Accepted: 16 October 1999     

The development of sun compass orientation in young homing pigeons

Summary A series of clock-shift experiments with young homing pigeons of various ages was performed to determine at what age they normally learn sun compass orientation. The response of untrained pigeons to shifting of their internal clock seems to depend on their age. When the clock-shifted birds were tested at an age of 11 weeks and younger, their departure bearings did not differ significantly from those of controls (Fig. 1, diagrams on the right); in tests with birds 12 weeks and older the characteristic deviation indicating the use of the sun compass was observed (Figs. 2 and 3). Birds that had participated in a short training program, however, used the sun compass at 8 weeks, the earliest age tested (Fig. 1, diagrams on the left). These findings show that the time of development of the sun compass strongly depends on flying experience. Within the first months of a bird's life, it seems to take place after the bird has been confronted with the need to orient, either spontaneously during extended exercise flights around its loft or imposed by training releases.The departure bearings of the very young, inexperienced birds that did not rely on the sun compass, however, were already oriented homeward. This indicates that the ability to navigate develops independently of the sun compass, before the sun compass is learned.Dedicated to Prof. Dr. F.W. Merkel for his 70th birthday     

Detour experiments with homing pigeons: information obtained during the outward journey is included in the navigational process

Summary To test the hypothesis that information on the route of the outward journey is involved in the orientation of displaced homing pigeons, we compared the behavior of control pigeons that had been displaced by the most direct route with that of experimental pigeons that had been transported along detours to the same release sites. At distances of 40 km we found no consistent effect. At distances between 75 and 130 km, however, deviations to the left of the direct route induced deflections to the left, while deviations to the right induced deflections to the right, i.e. the deflections of the vanishing bearings tended to compensate for the initial detour of the outward journey. The deflections were smaller than the deviations of the routes; they were not related to the routes themselves or the location of the release sites. A significant correlation emerged with the vector length of the controls, as longer vectors were associated with smaller deflections. This suggests that information on the route of the outward journey is used together with local map information in the navigational process, the significance of the route-specific information apparently depending on quality and reliability of the available local information. The nature of factors controlling the detour effect is still open.Correspondence to: R. Wiltschko     

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.     

Contribution of olfactory navigation and non-olfactory pilotage to pigeon homing

Summary By means of training flights (mostly flock releases), two groups of young homing pigeons were made familar with a larger area asymmetrically extending from the loft toward SW and NE, respectively. Thereafter, birds of both groups were released within each of the training areas with which one sort of birds was familiar (F⁺), the other unfamiliar (F^-). Even the F⁺ pigeons had never been released at the test site itself (nearest previous release 10 km apart). Half of each group was allowed to smell environmental odors (O⁺); the other half breathed charcoal-filtered air during transportation and at the release site until a few minutes before release when they were deprived of olfaction by intranasal application of Xylocain (O^-). The two test sites were 53 km distant from home. There was little difference in initial orientation as well as in homing performance between pigeons that were allowed to smell natural air and were familiar with the area (F⁺O⁺) and those that were privileged in only one respect (F⁺O^- and F^-O⁺). Yet if none of the preconditions was met (F^-O^-), performances were drastically reduced. The findings show that pigeons make use of two independent homing methods, olfactory navigation (presumably based on a navigational map) and non-olfactory pilotage (presumably based on a topographical map). The latter method is restricted to a more or less familiar area determined by individual experience. Its boundaries are poorly defined and can be estimated by the experimenter only in rough approximation. Within this area, the homing system takes advantage of more or less redundant inputs. Outside of it, olfactory information seems indispensable.