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.     

Genetic determination and plasticity in the sun orientation of natural populations ofTalitrus saltator

As has been previously shown, talitrid crustaceans have an inherited compass which causes them to head in a direction appropriate to their original shoreline (even after several generations in the laboratory), while learnt components can improve the correctness of orientation in natural conditions. In order to extend this analysis to a greater range of populations and to show differences in the determination of direction finding behaviour, seven natural populations ofTalitrus saltator (Montagu) from differently oriented shores of the Ligurian (Marina di Vecchiano, Pisa), Tyrrhenian (Castiglione della Pescaia and Tombolo di Feniglia, Grossetto) and Adriatic (Casal Borsetti, Ravenna; Torre Fantine, Marina di Lesina and Lido di Siponta, Foggia). Italian coasts were compared for their rigidity-plasticity in sun orientation. The study was conducted from 1982 to 1985. Differences were given in relation to the dynamics of the shorelines, the stability of which changes over the years due to natural and artificial causes. Furthermore, in order to reveal something about the genetic determination of the compass, mass-crosses between geographically distant populations (Ligurian-Tyrrhenian on one side and Adriatic on the other) were performed, and the F1 and F2 inexpert offspring tested for sun orientation. The results show a partial or total disruption of the compass in cross offspring which is discussed in the light of oligogenic and polygenic transmission mechanisms.     

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.     

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: the effect of a clock-shift is often smaller than predicted

This analysis is based on 103 releases with 6-h clock-shifted pigeons of various ages and experiences. Resetting the internal clock normally leads to a significant change in initial orientation; however, in half of the cases, the induced deflections are significantly smaller than predicted by the sun compass hypothesis. The relative size of the deflections decreases with increasing age and experience (Fig. 3). Only young pigeons with limited experience respond as expected, while old birds show deflections which are, on the average, only slightly more than half of the predicted size, except at extremely familiar sites (Table 2). There is no difference between fast and slow shifts (Fig. 4). It is not possible to clearly specify under what circumstances smaller deflections occur; previous clock-shifts (Fig. 5), familiarity with the release site (Table 4) and duration of the shifting procedure (Table 5) do not seem to be the reasons. Clock-shifting also tends to decrease the vector lengths and has a marked effect on homing performance (Table 7). Nevertheless, considerable numbers of clock-shifted birds return on the day of release before their internal clock has begun to be reset back to normal. The general role of the sun compass in bird orientation is considered and theoretical implications of our findings are discussed in view of the map and compass-model and the possibility that an alternative, non-time-compensating compass is used in parallel with the sun compass.     

Pigeon homing: sun compass use in the southern hemisphere

Although the sun compass of birds is based on learning the sun's arc during development, it was unclear whether birds can use the sun when its apparent movement is reversed, in particular, whether northern birds that have been introduced into the southern hemisphere can use the southern sun. To answer this question, clock-shift experiments were performed with local homing pigeons in Auckland, New Zealand (37°S). In three fast-shift tests and two slow-shift tests, the experimental birds showed deflections from the untreated controls that were the mirror images of those observed in the northern hemisphere. These results clearly show that homing pigeons in New Zealand use a sun compass that is adapted to the situation in the southern hemisphere. The learning processes establishing the compensation mechanisms thus appear to be free of constraints concerning the direction of the sun's movement. Differences from recent findings with migratory birds, where the direction of celestial rotation proved of crucial importance for establishing the migratory direction, are discussed: the differences may arise from the different orientation tasks, in particular, from the involvement of innate information in establishing the migratory direction. Received: 13 November 1997 / Accepted after revision: 28 February 1998     

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.     

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     

Sun-compass migration by Aurelia aurita (Scyphozoa): population retention and reproduction in Saanich Inlet,British Columbia

The scyphomedusa Aurelia aurita in Saanich Inlet, a north-south oriented fjord in British Columbia, uses a sun compass to migrate in a southeasterly direction during the day. When the sky is overcast and at night, A. aurita orients randomly and is dispersed passively by gentle tidal currents. The net result is daily reaggregation of medusae into enormous swarms along the southeastern shore of the fjord. Observations of spawning A. aurita in these swarms suggest that sun-compass migration and aggregative behavior may have evolved to facilitate reproduction and to maintain the population within this fjord throughout the year.     

When does bearing magnets affect the size of deflection in clock-shifted homing pigeons?

Pigeons whose internal clock is shifted by 6 h show deflections from the direction of untreated controls, yet these deflections are often smaller than predicted. Magnets temporarily disabling the magnetic compass increased these the deflections significantly (R. Wiltschko and Wiltschko 2001), indicating a compromise between sun compass and magnetic compass. – Recently, Ioalé et al. (2006) claim that they could not replicate our findings. The reason lies in a difference in the behavior of the clock-shifted pigeons without magnets: in the study of Ioalè et al. (2006), their deflections was already almost as large as that of our pigeons carrying magnets. This difference is probably caused by the limited experience of the pigeons of Ioalè et al. (2006): Their birds, in contrast to ours, had not used their sun’ compass during extended homing flights at various times of the year and, not having been faced with the necessity to compensate the saisonal changes of the sun’s arc, gave the sun compass more weight than our birds did.A comment to the paper by Ioalè, Odetti and Gagliardo (2006) Behav Ecol Sociobiol 60: 516–521.     

Sunset and the orientation of a nocturnal bird migrant: A mirror experiment

Summary If savannah sparrows, (Passerculus sandwichensis), a North American night migrant, select a migratory heading based upon the setting sun, a shift in the position of that cue should produce a predictable shift in the migrant's nocturnal orientation. I tested this hypothesis by shifting the sunst position with mirrors and by recording the bird's orientation in Emlen funnels. The control group displayed directionality appropriate for spring migration (=342°). The mean heading of experimentals (=272°), which were exposed to a cue-shifted situation, was in the expected westerly direction relative to the control mean (P<0.05, V-test). The setting sun appears to be a sufficient source of directional information for this avian migrant.     

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     

Polarized light sensitivity in the stage I zoea of the mud crab Panopeus herbstii

Stage I zoea larvae of Panopeus herbstii were tested for their orientations while swimming in a glass chamber and exposed to either polarized or unpolarized light. The larvae assumed a primarily vertical orientation of the dorsal spine (body axis horizontal) under all experimental conditions. Sensitivity to polarized light was demonstrated when larvae exposed to a polarization plane tilted off the horizontal shifted their orientations in the direction of the e-vector tilt. In addition, more of the larvae exposed to a horizontal e-vector were oriented to within a few degrees about the vertical direction, and they were less variable in their orientations than the populations of larvae exposed to unpolarized light. The possible ecological implications of such behavior are discussed.     

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     

Do bearing magnets affect the extent of deflection in clock-shifted homing pigeons?

When released after clock-shift, homing pigeons fail to orient towards the home direction but display a consistent deflection of their initial orientation due to the difference between the real sun azimuth and the computed azimuth according to the subjective time of each single bird. It has been reported that the size of the observed deflection is frequently smaller than expected and a discussion on the possible factors affecting the size of deflection has emerged. Some authors have proposed that the major factor in reducing the deflection after clock-shift is the simultaneous use of both the magnetic and the sun compasses, giving true and erroneous information, respectively, about the home direction. Therefore, a magnetic disturbance, by impeding the use of the geomagnetic information in determining the home direction, is presumed to increase the size of the deflection up to the levels of the expectation. To test this hypothesis, we released three groups of clock-shifted birds from unfamiliar locations (unmanipulated pigeons, pigeons bearing magnets on their head, and pigeons bearing magnets on their back) together with a group of unshifted control birds. As no difference in the orientation of the three groups emerged, we were not able to confirm the hypothesis of the role of the magnetic compass in reducing the expected deflection after clock-shift.Communicated by W. Wiltschko     

Bottoms up: great bustards use the sun to maximise signal efficacy

Visual displays are signals that may be selected to increase visibility. Light is a crucial component in the transmission of visual signals, and white colour is very conspicuous when illuminated by sun and exhibited against darker backgrounds. Here we tested the hypothesis that orientation of sexual displays in male great bustard (Otis tarda) depends upon position of the sun, i.e., males direct their uplifted white tails towards the sun in order to maximise signal detectability to distant females. We recorded the orientation of 405 male displays in relation to the sun and to females at seven leks. Great bustard males signalled towards the sun more often than expected by chance in early morning, although this pattern was not obvious at other times of day, when males displayed more towards females. Our hypothesis was further supported by the fact that displays were more directed towards the sun when the sun was most visible. Males were more likely to direct their displays towards females during the most elaborate components of their courtship display and when there were fewer males on the lek. Pointing white plumage to the sun may be a behaviour selected in species living in steppe-like open landscapes if individuals obtain net fitness benefit by increasing the likelihood of mating.     

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.     

Solar orientation in the snail Nerita plicata (Prosobranchia: Neritacea) on a beach near Watamu,Kenya

Investigations carried out under natural conditions suggest that Nerita plicata Linné is capable of orientating to the sun. Behaviour is predominatly photonegative during the morning, and photopositive during the afternoon. Accuracy of orientation improves as the brightness of the sun increases. At Watamu (Kenya), snails tended to orientate away from the sea at all times of the day. Orientatory performance is not affected by the temporal proximity of high or low water, and locomotory speeds seem to be unaffected by the strength of the sun, the time of day, or the state of the tide.     

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