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     

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.     

Homing pigeons do extract directional information from olfactory stimuli

Summary Two groups of pigeons were kept from fledging time in two cages fully exposed to winds. From time to time, the cage containing experimentals was additionally exposed to an artificial air current coming from a specific direction and carrying a scent of benzaldehyde. When both groups were exposed to benzaldehyde scent during transportation and at the release site, the control birds flew homeward, whereas the experimentals oriented in the direction roughly opposite that from which they were used to perceiving the benzaldehyde at the loft. When benzaldehyde was not applied, experimental pigeons were homeward oriented like controls.     

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 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.     

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     

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.     

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.     

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.     

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 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.     

Homeward paths of displaced juvenile alligators as determined by radiotelemetry

Summary To test their ability to home without searching or wandering, 19 juvenile alligators were radiotracked after displacement to areas 1–10 home range diameters beyond their areas of familiarity. Movement rates varied inversely with vegetation density, and in some cases habitat barriers appeared to deflect an alligator's homeward path. However, failures to home occurred only in those cases where major habitat barriers separated an alligator from its home site. In all other cases the alligators headed directly homeward; at least 10 of these completed their homeward journeys. The orienting ability of the alligators was neither dependent on the conditions under which they were displaced, nor the environmental conditions prevailing after release. These observations support the hypothesis that alligators possess a true navigational ability.     

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.     

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     

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     

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.     

Sex differences in partner preference in mated pairs of saddle-back tamarins (Saguinus fuscicollis)

Summary The attraction of permanently cohabiting males and females to their pair mate was assessed in a preference situation in which a subject was given a choice between its pair mate and a stranger of the mate's sex. Animals were tested in a choice apparatus, consisting of 1 cage housing the subject, 1 cage housing its pair mate, and 1 cage housing the strange stimulus animal. Wire mesh tunnels connected the subject's cage with that of each stimulus animal. Each tunnel was divided into 5 sections of equal length. During a series of 30-min choice tests, the subjects' location was scored every 10 sec as being in the home cage or in one of the sections of the tunnels leading to the cages of the other animals. In addition to these location scores, being at Zero Distance from a stimulus animal, as well as a number of behaviors directed at both stimulus animals, were recorded. Males were in closer proximity to their females than to strange females (Fig. 2), approached their females more frequently and directed more Tongue Displays and Sniffs at their mates (Figs. 3–5). Females did not prefer their males in terms of these behaviors (Figs. 2–5). Moreover, they directed a higher number of Gazes at strange males (Fig. 6). Both males and females were more frequently at Zero Distance from their pair mates than from strange stimulus animals and had more Nonaggressive Contacts with their pair mates (Figs. 7–8). These results suggest that males show a strong attachment to their female pair mates while females are attracted to their mates as well as to strange males.     

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     

春季不同天气城市街头绿地内PM2.5质量浓度分布特征研究

于2012年春季,采用水平分布监测法监测了3种天气下距交通污染源不同距离的街头绿地中的PM2.5质量浓度,研究了不同天气下PM2.5的日变化、水平分布规律及绿地对PM2.5的净化效应,为城市街头绿地、城市公园建设及市民合理选择休闲锻炼时间和地点提供理论依据。结果表明：1）晴天和多云时,PM2.5质量浓度上午高于下午,7：00浓度最高,15：00最低;雨后阴天基本保持上升趋势。2）PM2.5日均质量浓度为晴天（61.67μg·m-3）〈多云（187.98μg·m-3）〈雨后阴天（291.48μg·m-3）。晴天除5：00和7：00外,其他时刻均达到国家二级标准,且13：00—15：00达到国家一类功能区空气质量要求;多云和雨后阴天PM2.5质量浓度分别超过国家二级浓度限值150.6%和288.6%。3）观测时段内,无论哪种天气,5：00、7：00、11：00和15：00绿地的净化功能较强,19：00净化功能均最差,所有监测点无一例外的表现为负效应。4）3种天气下,PM2.5质量浓度在距离道路10~25 m最高,绿地的净化效应最差,55 m外基本可以形成稳定的森林内环境。5）在南方高湿环境下,空气相对湿度是影响PM2.5质量浓度的主要因素,晴天和多云天气PM2.5浓度与相对湿度呈显著正相关,而雨后阴天二者呈负相关关系。6）在一定阈值内,街头绿地能够缓解PM2.5污染,为居民提供良好的休闲环境。从游憩时间来看,市民可以选择晴天进入街头绿地休闲,多云和雨后阴天尽量减少外出;从活动最佳地段来看,距离污染源55 m以上适宜休闲锻炼;从街头绿地的规划面积来看,半径以不小于55 m为宜。