Do bearing magnets affect the extent of deflection in clock-shifted homing pigeons?

When released after clock-shift, homing pigeons fail to orient towards the home direction but display a consistent deflection of their initial orientation due to the difference between the real sun azimuth and the computed azimuth according to the subjective time of each single bird. It has been reported that the size of the observed deflection is frequently smaller than expected and a discussion on the possible factors affecting the size of deflection has emerged. Some authors have proposed that the major factor in reducing the deflection after clock-shift is the simultaneous use of both the magnetic and the sun compasses, giving true and erroneous information, respectively, about the home direction. Therefore, a magnetic disturbance, by impeding the use of the geomagnetic information in determining the home direction, is presumed to increase the size of the deflection up to the levels of the expectation. To test this hypothesis, we released three groups of clock-shifted birds from unfamiliar locations (unmanipulated pigeons, pigeons bearing magnets on their head, and pigeons bearing magnets on their back) together with a group of unshifted control birds. As no difference in the orientation of the three groups emerged, we were not able to confirm the hypothesis of the role of the magnetic compass in reducing the expected deflection after clock-shift.Communicated by W. Wiltschko     

When does bearing magnets affect the size of deflection in clock-shifted homing pigeons?

Pigeons whose internal clock is shifted by 6 h show deflections from the direction of untreated controls, yet these deflections are often smaller than predicted. Magnets temporarily disabling the magnetic compass increased these the deflections significantly (R. Wiltschko and Wiltschko 2001), indicating a compromise between sun compass and magnetic compass. – Recently, Ioalé et al. (2006) claim that they could not replicate our findings. The reason lies in a difference in the behavior of the clock-shifted pigeons without magnets: in the study of Ioalè et al. (2006), their deflections was already almost as large as that of our pigeons carrying magnets. This difference is probably caused by the limited experience of the pigeons of Ioalè et al. (2006): Their birds, in contrast to ours, had not used their sun’ compass during extended homing flights at various times of the year and, not having been faced with the necessity to compensate the saisonal changes of the sun’s arc, gave the sun compass more weight than our birds did.A comment to the paper by Ioalè, Odetti and Gagliardo (2006) Behav Ecol Sociobiol 60: 516–521.     

Pigeon homing: the effect of a clock-shift is often smaller than predicted

This analysis is based on 103 releases with 6-h clock-shifted pigeons of various ages and experiences. Resetting the internal clock normally leads to a significant change in initial orientation; however, in half of the cases, the induced deflections are significantly smaller than predicted by the sun compass hypothesis. The relative size of the deflections decreases with increasing age and experience (Fig. 3). Only young pigeons with limited experience respond as expected, while old birds show deflections which are, on the average, only slightly more than half of the predicted size, except at extremely familiar sites (Table 2). There is no difference between fast and slow shifts (Fig. 4). It is not possible to clearly specify under what circumstances smaller deflections occur; previous clock-shifts (Fig. 5), familiarity with the release site (Table 4) and duration of the shifting procedure (Table 5) do not seem to be the reasons. Clock-shifting also tends to decrease the vector lengths and has a marked effect on homing performance (Table 7). Nevertheless, considerable numbers of clock-shifted birds return on the day of release before their internal clock has begun to be reset back to normal. The general role of the sun compass in bird orientation is considered and theoretical implications of our findings are discussed in view of the map and compass-model and the possibility that an alternative, non-time-compensating compass is used in parallel with the sun compass.     

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     

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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Pigeon homing: sun compass use in the southern hemisphere

Although the sun compass of birds is based on learning the sun's arc during development, it was unclear whether birds can use the sun when its apparent movement is reversed, in particular, whether northern birds that have been introduced into the southern hemisphere can use the southern sun. To answer this question, clock-shift experiments were performed with local homing pigeons in Auckland, New Zealand (37°S). In three fast-shift tests and two slow-shift tests, the experimental birds showed deflections from the untreated controls that were the mirror images of those observed in the northern hemisphere. These results clearly show that homing pigeons in New Zealand use a sun compass that is adapted to the situation in the southern hemisphere. The learning processes establishing the compensation mechanisms thus appear to be free of constraints concerning the direction of the sun's movement. Differences from recent findings with migratory birds, where the direction of celestial rotation proved of crucial importance for establishing the migratory direction, are discussed: the differences may arise from the different orientation tasks, in particular, from the involvement of innate information in establishing the migratory direction. Received: 13 November 1997 / Accepted after revision: 28 February 1998     

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.     

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.     

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     

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     

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     

Pigeon homing: a comparison between groups raised under different conditions

Summary The homing ability of a group of pigeons raised under conditions of continuous exposure to windborne information was compared to that of another group subjected to exposure to winds only during the time spent in pre-test training flights. The results of release experiments at sites that were unfamiliar to the pigeons show that both bird groups, which had been made temporarily anosmic, had impaired initial homeward orientation and homing ability. These results, which do not entirely concur with the recent findings of Wiltschko and his coworkers, show that the raising and training procedures we used do not influence the nature of the cues used in pigeon homing. If the exposure of birds to wind-borne information is reduced, this does not bring into play any effective non-olfactory mechanisms of navigation; it simply hinders the development of the olfactory map and impairs the homing ability. Offprint requests to: S. Benvenuti     

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     

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.     

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.     

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.     

Initial orientation of homing pigeons at the magnetic equator with and without sun compass

Summary To test the present hypotheses concerning the functioning of the bird's magnetic compass, pigeons reared near the magnetic and geographic equator (Fortaleza, NE Brasil) were released 300 km NW of their home in the horizontal field at the magnetic equator. Pigeons released in the morning and in the afternoon were roughly homeward oriented whereas pigeons released at noon with the sun near the zenith vanished close to magnetic north. According to the Wiltschko model of the magnetic compass they should not be able to pick up specific directions. A considerable number of young and inexperienced pigeons returned home against a continuously blowing trade wind. This result contradicts the hypothesis of olfactory navigation as currently discussed.     

Bird orientation: Single homing pigeons compared with small flocks

Summary The hypothesis that bird flocks orient more accurately than single individuals was tested on homing pigeons. Birds were released both singly and in flocks of three to six. Vanishing bearings were recorded and it was found that flocks were less scattered around the mean direction than singly released birds. Homing times were found to be shorter for flocks as compared to singles. This suggests that the average homing pigeon can gain in directional accuracy and save energy by joining other pigeons heading for the same goal.     

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.