Homing behaviour in the sleepy lizard (Tiliqua rugosa): the role of visual cues and the parietal eye

To investigate how visual cues are integrated into a navigational strategy for homing in the Australian sleepy lizard (Tiliqua rugosa), lizards were displaced beyond their home range, either with full access to visual cues or with no access to visual cues during the displacement. Homeward orientation was significantly worse when lizards were denied visual cues during the displacement than when they were not. However when lizards were displaced with their field of view restricted to the sky, their homeward orientation was equally as good as that of lizards displaced with no visual restriction. These experiments suggest that sleepy lizards use celestial cues to determine the compass bearing of the outward journey, and reverse this bearing to orient in the homeward direction (course reversal). In a subsequent experiment, lizards oriented randomly with respect to home when the parietal eye was entirely covered with a patch during the displacement and return, while control lizards fitted with a sham parietal eye patch were well oriented towards home. In both groups, the lateral eyes were unobstructed and had complete access to visual cues including celestial cues and landmarks. These results suggest that the parietal eye plays a highly significant role in sleepy lizard homing, perhaps mediating a sky polarization compass sense.     

Orientation of the pied flycatcher Ficedula hypoleuca: cue-conflict experiments during spring migration

Hierarchical relationships among different compass systems in long-distance migrants are still a matter for discussion because different studies have led to highly variable and apparently contradictory results. We carried out cue-conflict experiments during spring migration on pied flycatchers Ficedula hypoleuca (Passeriformes, Muscicapidae). Birds were exposed to a conflict between celestial and magnetic information by altering the polarized light pattern or magnetic field. The polarization pattern was shifted (±90°) with filters, whereas the magnetic field was altered (+90°) through Helmholtz coils. Birds were tested in modified Emlen funnels both before and after the cue conflict; during the tests, only the natural magnetic field was available. This protocol was designed to test whether the experimental birds recalibrated their magnetic compass on the directional information derived from the light polarization pattern when the region near the horizon was visible during the conflict. Contrary to this expectation, we did not record any significant shift in magnetic orientation after one or repeated exposures to the cue conflict. Our results support earlier studies, which suggest that the magnetic field is the primary compass cue during the migratory period.     

Heterospecific call recognition and phonotaxis in the orientation behavior of the marbled newt,Triturus marmoratus

The role of acoustic cues as reference cues for orientation by amphibians has been demonstrated in anurans, the only amphibian group that engages in acoustic communication, but not in urodeles. Orientation responses of marbled newts, Triturus marmoratus, were studied to determine whether heterospecific calls elicited positive phonotaxis. The orientation tests consisted in presenting either a familiar acoustic stimulus, the advertisement calls of natterjack toads (Bufo calamita), or a control stimulus, the advertisement calls of European green toads (Bufo viridis) that the newts would not be expected to recognize. Marbled newts and natterjack toads occur in simpatry, but T. marmoratus and B. viridis are allopatric species. Thus, T. marmoratus is distributed over the northern half of the Iberian Peninsula, whereas B. viridis occurs in the Balearic Islands, but not over the Iberian Peninsula. Newts were released in a circular arena while a recorded chorus of natterjack toads or European green toads played outside the arena to determine whether they displayed positive phonotactic orientation. Our results show that marbled newts performed positive phonotaxis when exposed to the breeding calls of natterjack toads, but not to those of European green toads. Newts chose a compass course in the direction of the advertisement calls of B. calamita. Acoustic information might improve orientation accuracy. This study is the first to provide evidence of heterospecific call recognition and positive phonotactic response in urodeles.Communicated by W. Wiltschko     

Acoustic orientation in the palmate newt, Lissotriton helveticus

Experiments reported here were carried out to investigate the use of acoustic cues by palmate newts (Lissotriton helveticus) for orientation and to study whether this behavior is learned, or whether two populations of palmate newts that cohabit with different frog species (Iberian green frog, Rana perezi, and European common brown frog, Rana temporaria) show different phonotactic preferences. The orientation tests consisted of presenting a control stimulus (white noise), a sympatric acoustic stimulus (calls of R. perezi or R. temporaria, depending on the origin of newts), or an allopatric stimulus (calls of natterjack toads, Bufo calamita, or R. perezi). Newts were released in a circular arena, while the acoustic stimuli were presented outside of the circular arena in four different compass orientation directions (0, 90, 180 and 270°). In this study, we show that L. helveticus performed positive phonotaxis toward the calls of R. perezi only when both species shared habitat, orienting randomly when R. perezi was absent from the newt’s natal population. Newts from both populations oriented randomly when exposed to the allopatric and control acoustic stimuli. These results suggest, for the first time, that recognition of the sympatric heterospecific calls could be learned. However, newts sharing the breeding pond with a population of R. temporaria oriented randomly when exposed to the calls of this species. The fact that the breeding seasons of R. temporaria and L. helveticus do not overlap in time does not allow the use of R. temporaria calls as a guidance mechanism for migrating individuals of L. helveticus.     

The role of daytime cues in the development of magnetic orientation in a night-migrating bird

To assess the role of celestial rotation during daytime in the development of the magnetic compass course, pied flycatchers (Ficedula hypoleuca Pallas, Muscicapidae) were handraised in Latvia under various celestial and magnetic conditions. Tests were performed during autumn migration in the local geomagnetic field (50 000 nT, 73° inclination) in the absence of celestial cues. A group of birds that had never seen the sky showed a bimodal preference for the migratory southwest-northeast axis, whereas a second group that had been exposed to the natural sky from sunrise to sunset in the local geomagnetic field showed a unimodal preference for the seasonally appropriate southwesterly direction. A third group that had also been exposed to the daytime sky, but in the absence of magnetic compass information, also oriented bimodally along a southwest-northeast axis. These findings demonstrate that observing celestial rotation during daytime enables birds to choose the right end of the migratory axis for autumn migration at the Latvian test location. This transformation of axial behavior into appropriate migratory orientation, however, requires the birds to have simultaneous access to information on both celestial rotation and the geomagnetic field. Received: 19 September 1997 / Accepted after revision: 22 November 1997     

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     

Do leaf-cutter ants Atta colombica obtain their magnetic sensors from soil?

How animals sense, process, and use magnetic information remains elusive. In insects, magnetic particles are candidates for a magnetic sensor. Recent studies suggest that the ant Pachycondyla marginata incorporates iron-containing particles from soil. We used leaf-cutter ants Atta colombica to test whether soil contact is necessary for developing a functional magnetic compass. A. colombica is the only invertebrate known to calculate a path-integrated home vector using a magnetic compass. Here, we show that A. colombica requires contact with soil to incorporate magnetic particles that can be used as a magnetic compass; yet, we also show that ants can biosynthesize magnetic particles. Workers from a soil-free colony ignored a 90° shift in the horizontal component of the geomagnetic field, yet oriented homeward despite the occlusion of any geocentric cues. In contrast, workers from a soil-exposed colony oriented to an intermediate direction between their true and subjective home in the shifted field. Homeward orientations under shifted fields suggest that ants calculated a path-integrated vector using proprioceptive information. Strikingly, ants from the soil-free colony also had magnetic particles; yet, as observed by ferromagnetic resonance, these particles differed from those in soil-exposed ants and were not associated with a magnetic compass sensitive to this experimental manipulation.     

Calibration of the sun compass by sunset polarized light patterns in a migratory bird

Summary Migrating birds derive compass information from the sun, stars, geomagnetic field and polarized light, but relatively little is known about how these multiple sources of directional information are integrated into a functional orientation system. We found that migratory warblers exposed to a rotated polarized light pattern at sunset oriented at a constant angle to the axis of polarization. When polarized light cues were eliminated, this shifted orientation was maintained relative to the setting sun. Polarized light patterns, thus, appear to provide a calibration reference for the sun compass in nocturnal migrants, and may also play a role in calibrating other compass systems. Correspondence to: J.B. Phillips     

Orientation of juvenile barn swallows (<Emphasis Type="Italic">Hirundo rustica</Emphasis>) tested in Emlen funnels during autumn migration

Although hirundines have been used extensively in homing experiments, to date no investigation of their migratory orientation has been carried out, despite the well-known migratory habits of many species of this family. This paper reports on a study of the orientation of the barn swallow (Hirundo rustica), a typical diurnal trans-Saharan migrant. Modified Emlen funnels were used to verify the suitability of this species for cage experiments and investigate the role of visual and magnetic cues during the birds first migratory journey. Juvenile swallows were mist-netted at a roost site in central Italy and then tested in a site 19 km apart. Orientation experiments were performed under four experimental conditions: natural clear sky and simulated overcast, in both local and shifted magnetic fields (magnetic North=geographical West). Under clear sky, the swallows tended to orient phototactically toward the best-lit part of the funnel and failed to respond to the magnetic field shift. Under overcast conditions, they oriented northward and modified their directional choices as expected in response to the shifted magnetic North. On the whole, our data indicate that swallows can use magnetic information for compass orientation. Possible explanations for the northward orientation of birds tested under overcast conditions are discussed.Communicated by W. Wiltschko     

Solar and landscape cues as orientation mechanisms in the beach-dwelling beetle Eurynebria complanata (Coleoptera,Carabidae)

The present paper reports on the first orientation experiments conducted on the strand-living beetle Eurynebria complanata (Linnaeus, 1767) (Coleoptera, Carabidae). The experiments were carried out from June to October 1988. Two different populations were used: one inhabiting a Tyrrhenian beach in Italy and the other a beach on the Atlantic coast of France. Response to sun compass cues was demonstrated in each population at the collection site and for the French population after transportation to Italy, where experiments were carried out on a differently oriented beach. Landscape cues were shown to improve the beetle's orientation capacity when these were tested together with a visible sun. Artificial landscapes were also tested under laboratory conditions. Different-sized silhouettes were placed in the four cardinal directions, and these envoked different responses depending on their height. The beetles oriented towards a black silhouette when this subtended an angle of 25°. The results obtained for the two populations are compared and discussed from an eco-ethological point of view.     

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.     

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     

Moon orientation in sandhoppers: effects of lighting treatments on the persistence of orientation ability

The moon orientation rhythm persists in sandhoppers removed from environmental entrainments relative to moon periodicity. In order to investigate the underlying mechanism, the crustacean amphipods Talorchestia capensis and Talitrus saltator were collected at new moon and exposed in the laboratory to an artificial light during the day, but to a variety of lighting treatments during the night: (1) continuous dark, (2) dim light in phase with the natural moon, (3) continuous dim light. The animals were tested under the moon 9–21 days later by using a new type of arena. The ability to orientate in a direction perpendicular to the shore persisted in sandhoppers under treatments 1 and 2, but not under treatment 3. A disturbance due to a phototactic tendency in Talorchestia was also observed in animals captured on the day of the test when exposed to unnatural lighting, but not when they were kept under natural light. The present findings show that the timing mechanism allowing compensation for changes in the moon’s position also persists in animals that have been long removed from entraining factors. The dependence of this orientation ability on nocturnal lighting and the disturbing effect of sudden changes in light intensity support the idea that the phasal lighting of the moon, and perhaps sunrise and sunset act as resetting factors for the moon’s orientation rhythm.     

Clock-shift experiments with Savannah sparrows, Passerculus sandwichensis, at high northern latitudes

Orientation can be difficult for nocturnal bird migrants at high northern latitudes because of the large changes of magnetic declinations, rapid longitudinal time-shifts experienced during a long-distance flight and the invisibility of stars during the polar summer. Both sunset cues as well as geomagnetic cues have been shown to be of great importance in the orientation system of Savannah sparrows, Passerculus sandwichensis. We used clock-shift experiments to investigate whether geomagnetic and sunset cues were used for migratory orientation by wild-caught young Savannah sparrows at high geomagnetic latitudes in Northern Canada. We exposed birds to a 4-h slow clock-shift, expecting a 60° clock-wise shift in orientation after the treatment. Under natural clear skies in the local geomagnetic field, the birds responded by showing a significant axial mean orientation directed towards the position of the setting sun in the NW and towards their preferred migratory direction in the SE. After exposure to the clock-shift for 6 days and nights the birds showed a clear response to the treatment and shifted significantly towards NNE. Birds that first oriented towards NW in the experiments before clock-shift tended to shift clock-wise, thus reacted to the clock-shift in the expected way. The reaction of the individual birds that originally oriented towards SE seems to vary. In summary, our birds did not select a constant angle (menotaxis) in relation to the sun's position during the experiments, but presumably were affected by the sun showing phototaxis or followed their magnetic compass. Possible explanations of the unexpected experimental results are discussed. Electronic Publication     

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.     

Interaction between solar orientation and landscape visibility in Talitrus saltator (Crustacea: Amphipoda)

A series of experiments on the littoral amphipod Talitrus saltator (Montagu) was carried out between April and September, 1978–1981, both under natural conditions and inland with artificial landscapes of different heights on the horizon, in order to assess the visual importance of the landscape in zonal orientation in populations from the Mediterranean coast, and to determine interactions between solar orientation and orientation based on the landscape. Inland, orientation in controls (permitting only vision of the sky and the sun) was compared to that of the experimental individuals, who could see a simulated landscape positioned landwards to them and seawards. In nature, sandhoppers released in a level arena with the landscape screened from view were compared with others released in an unscreened arena and on the sand in absolutely natural conditions. Situations where solar orientation contradicted local cues were produced both by releasing the sandhoppers on a shore diversely orientated to their own, and by shifting their internal clock by nine hours. Results show that T. saltator uses the landscape as a cue in its orientation towards the sea, in conjunction with solar orientation: the latter being the principal factor involved even when the sandhoppers are separated from the local optical factor in the sky (in trials at a distance from the sea). In fact, with the artificial landscape set seawards, none of the populations we studied showed any orientation based on the landscape comparable in accuracy to orientation based exclusively on the sun. From the experiments carried out in natural conditions, it is possible to deduce that conflicting conditions gave rise to two types of results: deviation from the mean direction and an increase in dispersion, up to total dispersion when the local factors were in total contrast to solar orientation (clock-shifted sandhoppers released on the sand).     

Astronomical orientation and learning in the shrimp Palaemonetes antennarius

Adult and young individuals of the shrimp Palaemonetes antennarius (Milne Edwards) collected in the field (experienced), and laboratory-born (inexperienced) young were tested in a transparent bowl screened to hide the surrounding landscape from view. Field individuals were collected in South Tuscany, Italy, and testing was carried out in spring-summer 1987 and 1988. Results confirm that adults are able to assume an ecologically efficient escape direction (towards deep-water) by using celestial cues. They also show that: (1) orienting factors consist of the sun and polarized light in the sky; (2) the compass mechanism is time-compensated; (3) escape direction is not genetically fixed but (4) is learnt during the shrimp's lifetime and allows ample modification. From the results, it is also hypothesized that genetic determination of the escape direction is related to the type of larval development (with or without planktonic phases) as well as to the vagility of the species.     

Sunset orientation of robins,Erithacus rubecula,with different fields of sky vision

Summary Migratory orientation of robins (Erithacus rubecula) at sunset was recorded using orientation cages, under clear autumn skies. The aim of the experiments was to examine the importance of different visible sky sections for the orientation of robins. I obtained the following results: (1) Robins tested with the visible sky section limited to 90° around zenith (45° above the horizon) showed a mean orientation that coincided with the average sunset azimuth, with little scatter around the mean angle (Fig. 2). (2) When the birds were allowed a more extensive field of sky vision (maximum 160°), they chose headings on an approximate north-south axis, significantly different from tests with a restricted view of the sky (Fig. 3). (3) Experiments were also performed in which the response of robins to a mirror deflection (about 120° counterclockwise) of visual cues in the lower parts of the sunset sky was examined. The outcome indicated that visual information in the lower part of the sky may be critical for the orientation of robins (Fig. 5). These results, together with recent findings that robins captured and tested at two nearby sites show distinctly different orientation behavior in relation to experimental manipulations of the magnetic field, suggest that priorities among orientation cues may differ depending on the migratory situation encountered.     

Pigeon homing in relation to geomagnetic,gravitational, topographical,and meteorological conditions

Summary Experienced homing pigeons were released at sites unfamiliar to them and with magnetic and gravity anomalies as well as in areas with rather normal fields throughout the FRG (41 releases when sunny, 14 when overcast; Figs. 1–3). The second-order release data were subjected to both univariate and multivariate statistical analysis (stepwise regression, factor analysis). The magnetic field strength and its gradients within the 1-km circle around the release site were determined from aeromagnetic maps of the anomalies of magnetic total intensity. Analogous variables were derived from gravity anomaly maps. It was tested whether the pigeons fly along that gradient to minimize the difference between the magnetic or gravity field at the release site and the loft at maximum rate. Further independent variables described magnetic K index, day-to-day variations of the magnetic components, topography, meteorological conditions, the number of the releases the pigeons had done, and the distance.Over magnetic anomalies widely varying in strength (departing – 250 nT to 300 nT from normal 600 m above ground), extent, and distance from loft, the pigeons vanished with less deviation from the homeward direction and faster than they did in areas with less irregular fields under sunny conditions; this is in contrast to other studies on magnetic anomalies, except one. At sites of gravity anomalies (15–49 mgal), the pigeons were significantly less homeward oriented and homed slower than at less anomalous sites (–9 to 14 mgal).Variables related to gravity were best predictors in 8 and and second predictors in 3 out of 15 regression analyses of the navigational parameters for the releases under sun. Six times the (absolute) amount of the gravity difference between the release site and the loft was selected first (Figs. 7B, C, 8B, C). The results suggest gravity to be involved in navigation as the pigeons' distance measure. Homeward directedness declined with increasing amount of the gravity gradient in the first 12 releases under sunny skies as well as when overcast (Fig. 8A). A preferred compass direction towards north-northeast was determined, being closest to the grand mean vector of the ascending gravity gradient (Fig. 6). The analyses failed to show directional preferences as assumed by the hypotheses tested. Temperature and degree of cloud cover provided some information for predicting mean vector lengths and mean vanishing times, respectively. The surface wind component in the homeward direction was correlated with median homing performances.