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.     

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.     

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     

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.     

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.     

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.     

Pigeon navigation: Charcoal filter removes relevant information from environmental air

Summary Homing pigeons were displaced and kept until they were released in airtight containers ventilated with environmental air that could be passed through: (a) a filter made of fiberglass paper retaining large portions of the solid and liquid aerosol particles, (b) an additional filter consisting of activated charcoal, or (c) no filter (controls). Before its release, each bird was taken out of the container, and its olfactory epithelium was immediately anesthetized by lidocaine (Xylocaine). Thus, neither experimentals nor controls were able to smell while their initial orientation behavior was being observed.The controls' initial bearings were better homeward-oriented than those of pigeons ventilated with charcoal-filtered air in 14 of 17 releases conducted in Italy and Germany, at distances of 24–155 km. In the final analysis, the bearings of the charcoal-filter birds did not show any relation to the direction toward home, whereas those of the controls did. Pigeons ventilated with air that had passed through only the paper filter did not behave noticeably differently from control pigeons.It is concluded that the material bases of olfactory navigation in pigeons are substances dispersed in the atmosphere, most probably in a molecular state.     

Pigeon homing: new airbag experiments to assess the role of olfactory information for pigeon navigation

Summary In three series of experiments we assessed the effects of olfactory and non-olfactory information collected en route or at the release site on the initial orientation of homing pigeons. In the first experiment, pigeons were transported in open crates to two sites located in opposite directions from the home loft. They were left at the site for 1 h, then put into airtight containers filled with air from that site and brought back to the loft. From there, controls were transported back to the original site. Experimentals were transported to the opposite site. Upon arrival at the site, the olfactory mucosae of both groups were anesthetized with Gingicain. Thus in this experiment, control and experimental pigeons were exposed to different olfactory as well as to different non-olfactory information during displacement and at the site. In the second series, controls and experimentals were treated as in the first experiment, except that they were enclosed in the airtight containers at the very beginning of the experiment and were ventilated with synthetic air until arrival at the final release site. This treatment excluded the possibility to perceive olfactory information en route or at the site. In this series, the two groups differed only with respect to non-olfactory information perceived during displacement and/or at the release site. In the third series, we exposed pigeons at the loft to air collected either at the later release site (controls) or to air collected at a site located in opposite direction of the home loft (experimentals). Here the two groups differed only with respect to their exposure to air of different origin. In all three series, the pooled controls showed a directional preference that was statistically indistinguishable from the home direction. All three experimental groups were disoriented. In the first two series, the differences in the initial orientation of control and experimental pigeons were highly significant. In the third experiment, there was only very weak statistical evidence for a difference between controls and experimentals. These results suggest that more than one factor is involved in the pigeons' navigation system. According to the present experiment with synthetic air, pigeons probably gather and process non-olfactory information during the first part of their transport from the home loft to the release site. Thus, some kind of, in this case non-olfactory, route reversal seems to be involved in the homing process. In addition, the experiment involving only manipulations of airborne information indicated some olfactory component. Previous experiments at our loft did not result in disorientation of the pigeons if either only the access to airborne information had been removed or when otherwise unmanipulated pigeons had been transported in detours to the final release site. Therefore, we assume that our pigeons' navigation system relies on several cues. Deletion of one cue can be compensated by other information. Often the initial orientation of our pigeons is disturbed only when at least one cue is removed and another one provides false information.     

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.     

Homing behavior of wood thrushes (Hylocichla mustelina)

Summary Using the wood thrush (Hylocichla mustelina) as an example of a typical nocturnal migrant, we employed radio telemetry to follow breeding birds during the course of homing movements following displacements. Seven thrushes were displaced over distances of 6.5 to 17.3 km in a variety of directions from their nesting territories. The thrushes moved in a series of short flights (mean = 2.1 km) performed primarily at dawn. Consequently, the birds took several days to home from even these relatively short displacements. Thrushes flew under clear and solid overcast skies and even in light rain. The pooled individual flights of the birds were significantly oriented in the homeward direction (Fig. 2). Their orientation relative to home did not improve significantly as they progressed toward the goal. Three of the birds were documented to have returned to their home territories. The detailed tracks of the birds preclude the possibility that they homed by random search.     

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.     

Postfledging Dispersal of White-crowned Pigeons: Implications for Conservation of Deciduous Seasonal Forests in the Florida Keys

From 1988 to 1991, we studied the postfledging dispersal of 31 radio-tagged White-crowned Pigeons ( Columba leucocephala ) from three natal keys in Florida Bay. Immature birds dispersed from the natal keys at 26–45 days after batching, and most young dispersed more than 20 km during the first 10 days postdispersal. Dispersing birds flew either north to the Florida mainland or east to northeast to the mainline Florida Keys. On the mainland, immature birds fed nearly exclusively within Everglades National Park or an adjacent state wildlife management area. On the mainline keys, White-crowned Pigeons selectively used 5.01–20 ha forest fragments (p < 0.10) during the first 72 hours postdispersal. After this period, dispersing birds showed no preference among fragment size classes but used deciduous seasonal forests more frequently than suburban habitat(p < 0.10). The spatial pattern of dispersal on the mainline keys suggests that, during the first 72 hours postdispersal. White-crowned Pigeons are not able to reach northern Key Largo, where 69% of the deciduous seasonal forests are protected in state or federal ownership. Protection of large forest fragments, especially on southern Key Largo, should be a priority for maintaining populations of White-crowned Pigeons. These forests provide a series of "stepping stones" that enable dispersing immature White-crowned Pigeons to fly to more distant areas where habitat availability is less restricted. This species is threatened in Florida and may play an important role in maintaining plant species diversity in the seasonal deciduous forests of south Florida by dispersing seeds of at least 37 species of trees and shrubs. Protection of sufficient habitat to allow successful postfledging dispersal of this important seed disperser will also protect the ecosystem's biodiversity.     

Colony choice in birds: models based on temporally invariant site quality

We propose two stochastic models to explain how birds choose colonies. In the resource choice model, birds settle at each site at a rate proportional to the total resources the site contains. In the reduced resource choice model, a smaller cohort of birds enters sites at a rate determined by the total resources at each, and the remaining individuals enter sites at a rate that is linearly proportional to the total number of birds already nesting at each site. Thus, a fraction of birds chooses sites based on the resources present, and the remainder are attracted to a site by the presence of other birds. Colony site quality is assumed not to vary between years. Both models result, on average, in an ideal free distribution of colony sizes if the birds' settlement rate is linearly related to the resources in a site, if resources are distributed equally among individuals within sites, and if individuals with equal resources have equal fitness. We applied these models to long-term data on colony sizes and site usage of cliff swallows in south-western Nebraska. A test of the resource choice model suggested that the swallow population as a whole did not choose sites based strictly on site quality or the total resources contained at each site. However, a test of the reduced resource choice model suggested that a smaller fraction of the individuals in each colony may have based their choice of site on local resource availability, with the remaining birds aggregating at those sites based on the number of birds already settled there. Tests of these models may provide insight into how individuals choose colony sites and why colonies vary in size.     

Strategies of homing behavior in the red squirrel,Tamiasciurus hudsonicus

Summary Seventeen red squirrels (Tamiasciurus hudsonicus) were displaced and released about 600 or 2000 m from their home site, and the routes they followed during several hours after release were recorded. The analysis of these routes shows that, in their first attempts to leave the release site area, the squirrels' orientation is non-random with respect to the home direction (Fig. 2a). However, after a few hundred meters in this initial direction, the animals tend to go back to the release site area, and then to perform a series of radial forays centered on the latter.The results suggest that, in the conditions of the experiment, the squirrels have some information on the direction, but not on the distance to the home site. Their behavior appears to be governed by two complementary (but occasionally conflicting) strategies, one being to use the directional information available, the other not to proceed further than a distance which corresponds to the distance normally covered in spontaneous exploratory trips. It is argued that this double strategy is appropriate for spontaneous homing trips consecutive to an exploratory foray.     

Natal dispersal in Belding's ground squirrels (Spermophilus beldingi)

Summary Dispersal from the natal site was documented in two populations of Belding's ground squirrels (Spermophilus beldingi) living at different altitudes in the Sierra Nevada of North America. Distance dispersed and age at dispersal were monitored by a combination of observation, trapping, radio telemetry, and examination of road kills. Dispersal was sexually dimorphic in both populations (Tables 1 and 2). All surviving males emigrated before they were 55 weeks of age, with most dispersing midway through the juvenile summer (Fig. 1). By contrast, most females remained within the boundaries of their mothers' home range (Fig. 2 and 3). Those very few females that did emigrate moved distances from their natal burrows similar to those travelled by dispersing males (200–450 m; Table 2), but females tended to disperse at a slightly older age. Significant differences between the study populations were found in distances moved by juveniles of both sexes (Fig. 2). Body weights of juvenile male dispersers were significantly greater than were those of juvenile males of equivilent ages that had not yet dispersed (Fig. 4). The results were considered in light of Shields' (1982) discussion of dispersal and inbreeding. I concluded that if dispersal is related to reproductive success, then dispersal distances selected for use in tests of evolutionary hypotheses should be measured just prior to the subject animals' first reproduction. Finally, although not tested in the present study, the evolutionary hypothesis most consistent with my data suggests that dispersal in S. beldingi may function to minimize nuclear family incest.     

Loft features influence the processing of navigational information by pigeons

The present study evaluates the impact of lofts on the development of the navigational system of pigeons. A series of results obtained between 2000 and 2004 were analyzed. The H-loft was placed 10 m above the ground, with a ring format and seven windows all around it as well as three external aviaries that allowed birds to have a wide view of the outdoors and of the sky and knowledge of the wind dynamics. The L-loft was placed at ground level, with a single window transformed into an external aviary. The L-loft was protected from the wind by surrounding trees, reducing the ability of the pigeons to access the wind dynamics and also restricting their access to information regarding the outside area and the sky. In a preliminary analysis, we found the parameter of “vanishing time” to differ significantly between lofts in two of the five age classes. A thorough analysis of this parameter throughout the ontogenetic development of the birds demonstrated a pronounced improvement in the information-processing system with age, with adults vanishing quicker than younger birds, although an impairment of development occurred during the third month of age, perhaps due to a reorganization of the system. This result was obtained when the effect of lofts was considered in the different age classes. Results suggest that loft features are an important factor in the ability of the young pigeons to exploit navigational cues.     

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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Satellite tracking of migrating loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>) displaced in the open sea

Loggerhead turtles (Caretta caretta) are known to migrate towards fixed, individually-specific residential feeding grounds. To study their spatial behaviour and their navigational ability, five loggerheads nesting in South Africa were captured when about to start their postnesting migration and tracked by satellite after having been displaced from their usual migratory route. The first turtle, released south of Madagascar about 1,148 km from the capture site, moved west up to mainland Africa and then reached her feeding grounds by following the coast. A second turtle, released farther away (2,140 km) close to La Réunion Island, stopped for some time on the Madagascar east coast, then turned southwards to round the island and regain the African mainland in the northwest, without however allowing us to establish the location of her residential grounds. Three other turtles were released off the Tanzanian coast, 2,193 km north of their nesting area, at the northern edge of the distribution of the feeding grounds along the African coast. All of them headed north, and one turtle found her residential grounds located north of the release site. The other two females started long-distance oceanic wanderings in which they crossed nearly the entire Indian Ocean, apparently being transported by the sea currents of the region. We conclude that adult loggerhead turtles are apparently unable to compensate for the displacement and can return to a pelagic life style characteristic of juvenile turtles. These findings suggest that South African loggerheads rely on simple orientation mechanisms, such as the use of the coastline, as a guide, and compass orientation, possibly integrated by spatiotemporal programmes and/or acquired maps of familiar sites.Communicated by R. Cattaneo-Vietti, Genova     

Intermittent breeding as a cost of site fidelity

Intermittent breeding (skipping a breeding season) can be the result of an adaptive decision by a focal individual, trading current reproductive success in favour of future reproductive success (residual reproductive value hypothesis). In contrast, an individual can also be forced by conspecifics to abandon the familiar breeding site and refrain from breeding due to lack of suitable alternative breeding sites or mates (competition hypothesis). I studied intermittent breeding in the territorial and site-faithful Eurasian oystercatcher Haematopus ostralegus, using a dataset covering 20 years. Intermittent breeding (in total 86 cases) occurred among breeders that formerly bred in high- as well as low-quality territories. The main factor associated with intermittent breeding in high-quality sites was death of a mate, while in low quality sites divorce was the most prominent factor. In 93% of the cases birds were forced to cease breeding due to pressure from conspecifics consistent with the competition hypothesis. There was no association between intermittent breeding and promotion to a territory of better quality. Instead, oystercatchers returned to breeding habitat of similar quality and at a very close distance (median distance 128 m) from the previous breeding location. Breeding absences lasted on average 2.4 years, with a maximum of 9 years, and the quality of the territory obtained after the absence varied with the duration of it. Birds who re-bred in a high-quality territory acquired this on average faster than those that re-bred in a low-quality territory, indicating that birds in high-quality sites are better competitors.     

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