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     

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.     

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.     

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     

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.     

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.     

Further experiments on olfactory navigation and non-olfactory pilotage by homing pigeons

Summary Supplementary to a previous investigation (Wallraff and Neumann 1989), further experiments were conducted with homing pigeons that were either familiar of unfamiliar with the release area, and that had or lacked olfactory access to environmental odours. All four possible pairwise combinations of these factors were tested. The previous results were confirmed, showing that in an unfamiliar area olfactory inputs are necessary for home-related orientation, while in a familiar area both olfactory and non-olfactory information can be utilized and each is more or less redundant as long as the other kind of information is also available. The degree of redundancy of olfactory inputs varies, obviously depending on the pattern of individual local experience. Correspondence to: H.G. Wallraff     

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.     

The effect of a “permanent” clock-shift on the orientation of experienced homing pigeons

Summary A group of experienced homing pigeons vas subjected to a 6 h slow shift of their internal clock and kept under these conditions for more than 2 months. During the overlap time between the natural and artificial photoperiods they were released for training flights to familiarize them with an area while living in a permanent shift.Tested outside the permanent shift training range, the experimentals always deviated about 30° clockwise from the mean of their controls, markedly less than in a regular 6 h slow shift. Inside the permanent shift training range, however, they oriented like the controls (Fig. 2). When their internal clock was returned to normal, the birds showed a larger counterclockwise deflection on their first flight, which was roughly comparable to the effect of a regular 6 h fast shift (Fig. 3). On later flights after normalization, this large shift was no longer found; instead we observed a roughly 30° counterclockwise deflection when they were released inside the permanent shift training range in the morning. This deflection did not seem to occur in the afternoon or outside the permanent shift training range (Figs. 4, 5), and it disappeared when the birds were repeatedly released from the same site (Fig. 6).The occurrence or non-occurrence of the deflection was independent of the duration of the shift or the time passed after normalization; it seemed to depend solely on whether the birds had become familiar with a given site in the situation of the permanent shift. This argues against an effect based on the sun compass. We tend to assume that the still unknown navigational map is involved. In this case, however, as the deflection is independent of the home direction and the type of release site bias, the factors in question would act very differently from the gradients on which the traditional concepts of the navigational map are based. The processes establishing and updating the map and their possible differences are discussed.Died on August 17, 1980     

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     

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).     

Orientation of sandhoppers at different points along a dynamic shoreline in southern Tuscany

Orientation experiments were carried out on Talitrus saltator (Crustacea Amphipoda) at four points along 3 km on a dynamic sandy beach inside the Maremma Regional Park (Grosseto, Italy) to highlight behavioural variation related to distance from a river mouth, to erosion or accretion of shoreline, and to human trampling on the beach. Tests were performed using circular transparent Plexiglas arenas, contemporaneously at the four points. Replicates were made in 2 different months (September 2002 and May 2003), on 2–3 successive days, in the morning and afternoon. The distributions of the angles of orientation were compared for the different points and seasons, and multiple regression analysis was performed to test the effects of environmental and intrinsic variables on orientation. Sandhoppers showed the highest scatter at the eroded shoreline, intermediate scatter at the accreting beach most distant from the river mouth, and consistent orientation seaward at the least disturbed point. Orientation of sandhoppers was significantly affected by season, global radiation, time of day, distance from the river mouth, and human trampling. Sex and air humidity were of minor significance in the multiple regression model. The results, on the one hand, confirm plasticity in orientation of sandhoppers living on a dynamic shoreline, and on the other hand, show that variation in orientation could potentially be used as a bioindicator of shoreline changes.Communicated by R. Cattanevo-Vietti, Genova     

Pigeon homing: A repetition of the deflector loft experiment

Summary Pigeons brought up in closed cages deflecting the winds to the left or to the right show, on release, a bias of the mean vanishing directions to the left or right of the mean direction of untreated control birds. The results are in full accord with the findings of Baldaccini et al. (1975).Supported by the Deutsche Forschungsgemeinschaft     

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.