Pigeon homing: The influence of topographical features in successive releases at the same site

A new device, the direction-recorder, offered the possibility to extend earlier studies of homing behaviour of pigeons when relevant topographical elements (mountains and large tracts of water) interpose between the release site and the home loft. Three series of experiments were carried out at three different sites to investigate intraindividual and interindividual variability in subsequent tosses from the same locality. Two release sites were chosen behind a mountain chain with respect to home; at the third site homeward directed route crosses the sea. From our results it turns out that homing pigeons may adopt different strategies. Moreover, a wide intraindividual variability was observed in repeated tosses at the same site; some pigeons remained faithful to the first route, whereas other birds tried successive new routes which, in most cases, were significantly shorter than previous ones. This result indicates that pigeons try, and are actually able, to improve their performance in subsequent releases from the same site.     

Does Familiarity with the Release Site Influence the Initial Orientation of Homing Pigeons? Experiments with Clock-shifted Birds

Experiments were performed to test whether the familiarity with the release site plays a role in the initial orientation of homing pigeons. Repeated releases of 6 h clock-shifted birds from the same site during the shifting time produced an improvement of their initial orientation: the shift effect decreased progressively. Since in subsequent releases from unfamiliar sites the shift effect reappears, the course correction observed at the familiar site is attributable to local stimuli and not to a general recalibration of the sun compass.     

Learning multiple routes in homing pigeons

The aerial lifestyle of central-place foraging birds allows wide-ranging movements, raising fundamental questions about their remarkable navigation and memory systems. For example, we know that pigeons (Columba livia), long-standing models for avian navigation, rely on individually distinct routes when homing from familiar sites. But it remains unknown how they cope with the task of learning several routes in parallel. Here, we examined how learning multiple routes influences homing in pigeons. We subjected groups of pigeons to different training protocols, defined by the sequence in which they were repeatedly released from three different sites, either sequentially, in rotation or randomly. We observed that pigeons from all groups successfully developed and applied memories of the different release sites (RSs), irrespective of the training protocol, and that learning several routes in parallel did not impair their capacity to quickly improve their homing efficiency over multiple releases. Our data also indicated that they coped with increasing RS uncertainty by adjusting both their initial behaviour upon release and subsequent homing efficiency. The results of our study broaden our understanding of avian route following and open new possibilities for studying learning and memory in free-flying animals.     

Olfaction and the homing ability of pigeons raised in a tropical area in Brazil

Several workers have investigated the effect of anosmia on pigeon navigation in different geographical locations because it has been suggested that homing behavior is based on different cues, such as olfactory cues, the Earth's magnetic field or infrasound, and that in the absence of one cue another would be used. In this situation, no cue is universally indispensable, including olfactory ones. In order to extend such observations to a novel biome, we observed the behaviour of 192 young inexperienced birds raised in southeastern Brazil, a tropical area where olfactory tests had never been run before. The birds were released from eight symmetrically distributed sites 17 to 44 km from the loft. Half of these birds (experimentals) had been made temporarily anosmic by washing their olfactory mucosae with 4% solution of ZnSO4 the day before release, while controls were treated with Ringer solution. The results of release tests showed that anosmia totally impaired the navigational performance of experimental birds, which were unable to home from sites at relatively short distances from home (34-44 km) and whose pooled initial bearings produced a (negative) homeward component not significantly different from 0. Homing performance of controls was significantly better, and their pooled vanishing bearings had a significant homeward component, in spite of much scatter in individual releases. We conclude that pigeon homing in the study area depends on olfactory information, even though local environmental conditions in the interior of the State of Sao Paulo, as in several other parts of the world, do not appear to be as favorable as Italy for the development of efficient olfactory navigation.     

What happens to translocated game birds that 'disappear'?

The ultimate goal of most translocation efforts is to create a self-sustaining wild population of a species deliberately moved from one part of their range to another. As follow-up of a translocation attempt is often difficult, causes for failure are relatively unknown. Dispersal away from the release site is one potential source of failure because it decreases the likelihood of the released population establishing itself post-translocation. In this study, we used chukar Alectoris chukar as a surrogate for translocated game birds in order to conduct a large-scale experimental study. We observed that these desert-adapted birds demonstrate a strong fidelity for specific water sources. We also report the propensity for the translocated individuals to either disperse and return to their original water source site or remain at the release site. During two field seasons, we observed opposing behaviors such that the proportion of individuals returning to the capture site, versus those remaining at the release site, shifted between years. We analyzed this change between the years as well as within the years to assess the potential underlying causes such as translocated distance, differences in rainfall between seasons and water source type. We concluded that homing behavior was strong in this non-migratory bird species and that strength of this homing behavior varied, potentially due to conditions surrounding the limiting resource, water availability. The large-scale, original data presented here may help to explain why some releases result in a successfully established population while other releases result in widely dispersed individuals.     

Multivariate analysis of initial orientation in homing pigeons: Is there a “preferred compass direction”?

The mean vanishing direction of homing pigeons of a given release often deviates from the home direction considerably. Any kind of geographical systematic in these deviations might give us a hint on the distribution of the information involved in the homing process. Analysis of releases arranged symmetrically around the home loft with methods of circular statistics could not resolve the pending question of site-specificity vs. directional effects in the initial orientation of homing pigeons unambiguously. In part, this ambiguity comes about by the facts that

Does pigeon homing depend on stimuli perceived during displacement?

Summary In order to test whether stimuli perceived during passive displacement are important for the subsequent homing, pigeons were transported in an apparatus designed to prevent them from receiving relevant information: The experimental birds were continuously rotated quite rapidly (1.5 cps, radial acceleration about 4 g); in addition, they were exposed to an artificial magnetic field and supplied with bottled air. Control birds were transported in open-air cages on top of the van with free view to all sides.Five pairs of releases from equal distances in opposite directions were conducted. Experienced birds were released at distances of about 15, 90, and 300 km from the loft, inexperienced birds at distances of about 180km. In each pair of releases both groups of pigeons were significantly homeward oriented. Neither in initial orientation nor in homing performance nor in the distribution of recoveries were the experimental birds inferior to the controls or in any perceptable way different from them.It is concluded that homing of passively displaced pigeons is not primarily based on information gathered during the outward journey.Abbreviations EP experimental pigeon(s) - CP control pigeon(s) The possibility to maintain our pigeon loft in a building that belongs to the Zoological Institute (Prof. M. Lindauer) of the University of Würzburg is gratefully acknowledged.     

Acetylcholine as sensory transmitter in crustacea

Summary In an attempt to learn more about the so-called map component in pigeon navigation, nine series of tests comprising 34 test releases were performed at a release site, 89 miles NNE of Ithaca, New York, where Cornell pigeons regularly depart nonrandomly but with a large clockwise deviation from the true home direction. The tests included releases of: (1) experienced pigeons new to the site, under sun; (2) pigeons with previous experience at the site, under sun; (3) experienced pigeons new to the site, under total overcast; (4) pigeons with previous experience at the site, under total overcast; (5) first-flight youngsters, under sun; (6) directionally trained pigeons; (7) pigeons from two other Ithaca-area lofts; (8) pigeons from two more distant lofts; (9) Bank Swallows from an Ithaca colony; (10) clock-shifted pigeons; (11) radio- and airplane-tracked pigeons.The results of these tests indicate that the directional bias of the Cornell pigeons is, in general, not dependent on weather conditions or on the previous experience of the birds. Moreover, a similar bias is shown by both pigeons from other Ithacaarea lofts and Bank Swallows from Ithaca. And a similar bias from their respective home directions is shown by birds from other areas. Clock-shifted pigeons departing more directly toward home have poorer homing success than controls. It is concluded that some environmental factor basic to the avian homing process is rotated clockwise at this release site, that biologically the birds are not making an error but are probably reading correctly a distorted map.I thank Irene Brown and Timothy Larkin for their aid in all phases of this project; Donald Windsor for help in two releases; Charles Walcott and Martin Michener for aid in airplane tracking; J. Downhower for collaboration in the radio tracking and swallow releases; Andre Gobert, Howard French, A. E. Newton, and Ludwig Karl for permitting me to use their pigeons, George Yerdon, Forest Ranger, and Andy Misura, District Ranger, New York State Department of Environmental Conservation, for their cooperation in making the facilities of the Castor Hill Fire Tower available to us; Monica Howland for preparing the drawings; and Bertha Blaker for typing. Research supported in part by NSF Grants GB-13046X and GB-35199X, and in part by Federal Hatch funds.     

Light-dependent information: influence of loft conditions on young pigeon’s navigational system

This study compares the initial orientation and homing performance of young inexperienced pigeons following their transportation to near and distant places in total darkness (treatment) and their subsequent release. The birds were housed in two lofts at the Lisbon Zoo. Each loft had its own specific features: the H-loft was exposed to prevailing winds and allowed an unhindered view of the surrounding landscape; the L-loft was protected from the wind and allowed only a partial view of the surroundings. Pigeons used in the release tests were between 6 and 7 weeks old. We found that, in general, the initial orientation of the pigeons was affected by the treatment: following release at near places, there was an increase in the scatter and a decrease the homeward component, suggesting that light-dependent information collected en route was used by young pigeons. The effect of the treatment was only temporary based on the observation that the homing performance was not affected. However, the distance of the release site strongly influenced the homing performances as pigeons appeared to be unable to home when released at locations distant from the loft. Based on the scatter or the homeward component, inter-loft differences were apparent with respect to different median vanishing intervals and the reactions of specific pigeons when subjected to the same treatment (transport in darkness) following release at near and distant places. These findings suggest that light-dependent information collected en route is a component of the young pigeon’s navigational system but that, at the young age of the birds tested here, it is preferentially used in familiar areas. In addition, the importance of the light-dependent information appears to depend upon prior experience obtained in the lofts.     

The Orientational Consequences of Flocking Behaviour in Homing Pigeons,Columba livia

Two experiments are described which investigate the orientational consequences of flocking in homing pigeons Columba livia. Previous experiments have shown that homing pigeons placed inside a clear-sided release box for 5 min before release from a familiar site have enhanced ground homing speed compared with those placed in an opaque-sided box. It is assumed that previewing the surrounding landscape allows for faster homing since a bird denied this information must accumulate the knowledge on release. In experiment 1, using the same technique developed in these experiments but releasing the birds in pairs we showed that within familiar areas, homing pigeons can exploit a partner that has acquired more information, allowing them to home more quickly. In experiment 2 we attempted to test three potential strategies which may occur during homing flights. The results do not conclusively distinguish between these three mechanisms but suggest that orientation of the pairs of birds is most likely to have resulted from a compromise of individual tendencies, or from following the best homer, but not from following a ‘governing leader’. The consequence of these mechanisms is discussed.     

Activity patterns of insectivorous bats and birds in northern Scandinavia (69° N), during continuous midsummer daylight

Previous studies suggest that many species of insectivorous bats are nocturnal, despite the relatively low availability of their insect prey at night, because of the risk of predation by diurnal predatory birds. We hypothesised that if this was the case bats living above the arctic circle would alter their feeding behaviour during midsummer because there would no longer be any benefit to restricting their activity to the period when their prey are least abundant. Alternatively, if bats were more influenced by competition from aerial insectivorous birds they would continue to feed at 'night' to avoid such competition. In northern Norway (69°  N), during continuous midsummer daylight, insectivorous sand martins ( Riparia riparia ) concentrated their aerial feeding activity when aerial insects were most abundant. The birds stopped feeding between 23:00 and 07:00 when aerial insects were least abundant. In contrast, northern bats ( Eptesicus nilssonii ), fed mostly between 22:00 and 02:00, coinciding with the lowest aerial insect availability, and with the period when light levels were lowest (ca 1000 lux). Bat activity patterns were closest to those predicted by the avian competition hypothesis. The low densities of both sand martins and Northern bats in the study area, however, were less consistent with this hypothesis. Possibly populations of both species were higher historically and the observed patterns reflected historical competition. Bat activity was most closely correlated to ambient light levels. This raised two alternative explanations that we could not eliminate. Perhaps there was differential predation risk, between the brightest and darkest parts of the day, because the visual capacities of falcons are strongly dependent on luminance. Alternatively the bats may have been entrained to emerge at given light levels by their behaviour at other times of year.     

Patterns of success in game bird introductions in the United States

Better predictions of the success of species’ introductions require careful evaluation of the relative importance of at least three kinds of factors: species characteristics, characteristics of the site of introduction, and event-level factors such as the numbers of individuals released. (Henceforth, we call this propagule pressure.) The 1644 introductions of 17 Phasianid species released in various US states during the Foreign Game Investigation Program provides a particularly rich source of data to test these ideas. An examination of these records indicates that 13 of these 17 species always failed, despite generally numerous individual releases and large numbers of individuals in each release. Moreover, ten of these species have been successfully introduced elsewhere. Only four of the 17 species were successful in at least one state. Some 20 sets of releases of three of these four species always failed in some states, again given generally numerous individual releases of large numbers of individuals in each release. Simply, the combination of site and species factors explain the lack of successes. This leaves a combination of 18 states where one or more of the four species succeeded. For these, there are significant differences in the numbers of birds introduced from state to state. But only for two species Alectoris chukar and; Tetraogallus himalayensis are there significant differences that show a greater chance of success when more individuals are introduced. These results support the conclusion that the number of individuals released, meaning propagule pressure, is not as important as characteristics of the species and the location to where its introduction occurred.     

Swimming behaviour and dispersal patterns of headstarted loggerhead turtles Caretta caretta

Swimming behaviour and dispersal patterns were studied in headstarted loggerhead turtles Caretta caretta which were released at three different sites on the Caribbean island of Curaçao (Netherlands Antilles) and at one site on the neighbouring island of Klein Curaçao, after 1–2.5 yrs of captivity. Turtles were tagged and followed up to a distance of 6125 m offshore, using a boat with a Global Positioning Unit. The released turtles reverted to typical hatchling behaviour and showed an offshore migration almost perpendicular to the coastline. No significant differences were found in directional swimming among the four sites. The turtles swam almost continuously about 30 cm under the water surface; their mean overall swimming speed was higher than in adult wild loggerheads suggesting a 'frenzy'-like swimming stage. The turtles exhibited diving behaviour, and the dive frequency and duration was comparable to that of similar-sized (wild) turtles. The present study demonstrates that upon release the headstarted loggerheads behave naturally and show dispersal patterns similar to wild hatchling turtles. The fact that the released turtles were still able to show offshore directional swimming suggests that the headstarting did not affect their short-term orientation abilities.     

Discovering the migration and non‐breeding areas of sand martins and house martins breeding in the Pannonian basin (central‐eastern Europe)

The central‐eastern European populations of sand martin and house martin have declined in the last decades. The drivers for this decline cannot be identified as long as the whereabouts of these long distance migrants remain unknown outside the breeding season. Ringing recoveries of sand martins from central‐eastern Europe are widely scattered in the Mediterranean basin and in Africa, suggesting various migration routes and a broad non‐breeding range. The European populations of house martins are assumed to be longitudinally separated across their non‐breeding range and thus narrow population‐specific non‐breeding areas are expected. By using geolocators, we identified for the first time, the migration routes and non‐breeding areas of sand martins (n = 4) and house martins (n = 5) breeding in central‐eastern Europe. In autumn, the Carpathian Bend and northern parts of the Balkan Peninsula serve as important pre‐migration areas for both species. All individuals crossed the Mediterranean Sea from Greece to Libya. Sand martins spent the non‐breeding season in northern Cameroon and the Lake Chad Basin, within less than a 700 km radius, while house martins were widely scattered in three distinct regions in central, eastern, and southern Africa. Thus, for both species, the expected strength of migratory connectivity could not be confirmed. House martins, but not sand martins, migrated about twice as fast in spring compared to autumn. The spring migration started with a net average speed of > 400 km d^–1 for sand martins, and > 800 km d^–1 for house martins. However, both species used several stopover sites for 0.5–4 d and were stationary for nearly half of their spring migration. Arrival at breeding grounds was mainly related to departure from the last sub‐Saharan non‐breeding site rather than distance, route, or stopovers. We assume a strong carry‐over effect on timing in spring.     

Use of trace elements in feathers of sand martin Riparia riparia for identifying moulting areas

We investigated whether trace elements in tail feathers of an insectivorous and long-distance migratory bird species could be used to identify moulting areas and hence migratory pathways. We analysed tail feathers from birds of different age and sex collected from a range of different breeding sites across Europe. The site of moult had a large effect on elemental composition of feathers of birds, both at the European and African moulting sites. Analysis of feathers of nestlings with known origin suggested that the elemental composition of feathers depended largely upon the micro-geographical location of the colony. The distance between moulting areas could not explain the level of differences in trace elements. Analysis of feathers grown by the same individuals on the African wintering grounds and in the following breeding season in Europe showed a large difference in composition indicating that moulting site affects elemental composition. Tail feathers moulted in winter in Africa by adults breeding in different European regions differed markedly in elemental composition, indicating that they used different moulting areas. Analysis of tail feathers of the same adult individuals in two consecutive years showed that sand martins in their first and second wintering season grew feathers with largely similar elemental composition, although the amounts of several elements in tail feathers of the older birds was lower. There was no difference between the sexes in the elemental composition of their feathers grown in Africa. Investigation of the trace element composition of feathers could be a useful method for studying similarity among groups of individuals in their use of moulting areas.     

Pigeon navigation: Time course of olfactory signal processing and dependence on access to fresh environmental air

Summary 16 releases, centrally symmetrical by pairs and involving distances of displacement from 25 to 172 km, were conducted with homing pigeons pre-treated in different ways: FIL birds were, until few minutes before release, confined to containers ventilated with ambient air that had passed through filters consisting of activated charcoal. NOFIL birds were confined to containers ventilated with unfiltered air, either from departure at the loft onwards (4 experiments) or for at least 4 h at the release site (after transport under FIL conditions; 12 experiments). The olfactory epithelia of XYL birds were locally anaesthetized a few minutes before release, while NOXYL birds were not treated with xylocain. FIL/NOFIL conditions were combined with XYL/NOXYL conditions, resulting in 4 types of experimental treatment.On average, the untreated pigeons (NOFILNOXYL) were best homeward oriented and the double-treated pigeons (FIL-XYL) poorest. More importantly, the effect of olfactory deprivation during initial flight alone (NOFIL-XYL) was small, whereas long-term filtration of environmental air was quite effective even if the pigeons could smell during release time (FIL-NOXYL) (Fig. 5).These findings indicate that pigeons usually need to be exposed to local odorous air for more than only few minutes in order to utilize information extricated from this air for site localization.Additional experiments showed that homeward orientation is also prevented, if the pigeons, although breathing natural air, are ventilated with restricted volumes of fresh air.Our results are discussed with regard to the homing mechanism of pigeons as well as to their methodological consequences.Abbreviations FIL breathing air filtered by charcoal - NOFIL breathing air unfiltered - XYL xylocain applied to pigeons' nostrils - NOXYL no xylocain applied (see p. 140)     

Orientation by pigeons deprived of olfaction by nasal tubes

Summary In an effort to avoid the trauma and other nonolfactory effects produced by surgical sectioning of pigeons' olfactory nerves, and to avoid the interference with breathing produced by nostril plugs, a way of making pigeons anosmic by inserting plastic tubes in their nostrils was developed. A total of 16 experimental releases were conducted from unfamiliar sites to compare the homing behavior of birds wearing a tube in each nostril with controls wearing a tube in only one nostril. In five short-distance releases (less than 25 km), no convincing differences in initial orientation, vanishing intervals, or homing success were observed. In eleven releases from longer distances (more than 76 km), the experimental birds were random in three cases and the controls were random in two. In no case were the differences in the distributions of the bearings of experimental and control birds statistically significant, nor were there ever significant differences in vanishing intervals. However, experimental birds had much poorer homing success from these long-distance releases. It is concluded, in view of the anosmic pigeons' good orientation at distant unfamiliar sites, that olfaction is not necessary for homeward orientation and hence that it cannot be the basis of the birds' navigational map. Poor homing success from long distances is probably a consequence of the physical irritation and interference with breathing unfortunately produced by the nasal tubes.We thank our colleagues, Irene Brown, Timothy Larkin, and André Gobert for their help in conducting the releases. This research was supported by Grant BMS 75 18905 AO2 from the National Science Foundation.     

Strong homing does not predict high site fidelity in juvenile reef fishes

After being displaced, juvenile reef fishes are able to return home over large distances. This strong homing behaviour is extraordinary and may allow insights into the longer-term spatial ecology of fish communities. For example, it appears intuitive that strong homing behaviour should be indicative of long-term site fidelity. However, this connection has rarely been tested. We quantified the site fidelity of juvenile fishes of four species after returning home following displacement. Two species, parrotfishes and Pomacentrus moluccensis, showed significantly reduced site fidelity after returning home. On average, they disappeared from their home sites almost 3 d earlier than expected. Mortality or competitive exclusion does not seem to be the main reasons for their disappearance. Rather, we suggest an increased propensity to relocate after encountering alternative reef locations while homing. It appears that some juvenile fishes may have a higher innate spatial flexibility than their strict homing drive suggests.

     

High-speed railways are not barriers to Pyronia tithonus butterfly movements

Linear infrastructures such as railways and roads can be barriers to the movements of individuals and, hence, may have strong impacts on populations. We tested the barrier effect of a high-speed railway for Pyronia tithonus, a butterfly species showing homing behaviour when displaced. We captured, marked and displaced 152?individuals in two different locations. One-third of the butterflies were released at a capture plot, one-third on the other side of the railway (in a similar habitat) and one-third on the same side but 100?m away from the capture plot. We obtained recapture rates of 40 and 23?% per location. Many (31?%) butterflies crossed the railway, showing homing behaviour. Thus, contrary to wide, busy roads, high-speed railways do not seem to be barriers for these butterflies. We suggest that in an intensive agrarian landscape, railway verges can play a substitution habitat role for grassland butterflies.     

Pigeon homing: Does initial orientation include a ‘preferred compass direction’?

Summary To find out whether the initial orientation of pigeons is affected by a spontaneous directional tendency, as postulated by Wallraff's (1974, 1978, 1982) hypothesis, experienced birds were released at test sites distributed symmetrically around their loft. — The length of the mean vectors of the single releases, the deviations from the home direction, the homeward components as well as the homing speed did not show a correlation with the geographic position of the home direction. Summarizing four sites each on 21 experimental circles, we frequently obtained significant compass vectors, but they varied in direction between 115° ESE and 351° N, depending on what sites had been used, and did not indicate a uniform trend.A preferred compass direction as an integrated part of pigeon navigation, being the reason for the frequently observed deviations from the home direction, could not be confirmed. — The problematic nature of simply pooling the data of several symmetrically distributed test sites and calling any resulting significant vector a preferred compass direction is discussed, together with other possible reasons for asymmetrical distributions of release site biases.Abbreviation PCD preferred compass direction