The Journal of Experimental Biology 203, 23412348 (2000) Printed in Great Britain The Company of Biologists Limited 2000

0 JEB2729

2341

PARTIAL EXPERIENCE WITH THE ARC OF THE SUN IS SUFFICIENT FOR ALL-DAY SUN COMPASS ORIENTATION IN HOMING PIGEONS, COLUMBA LIVIA
CHERI A. BUDZYNSKI1,2,*, FRED C. DYER3 AND VERNER P. BINGMAN1,2 of Psychology and 2J. P. Scott Center for Neuroscience, Mind, and Behavior, Bowling Green State University, Bowling Green, OH 43403, USA and 3Department of Zoology, Michigan State University, East Lansing, MI 48824, USA
1Department

*e-mail: cherib@bgnet.bgsu.edu

Accepted 2 May; published on WWW 10 July 2000 Summary orientation cue. The control group and the experimental The ability of animals to learn to use the sun for group, which was exposed to the morning sun for the rst orientation has been explored in numerous species. In time, succeeded in orienting in the training direction birds, there is conicting evidence about the experience during test 1. The orientation of the experimental group needed for sun compass orientation to develop. The was no different from that of the control group, although prevailing hypothesis is that birds need entire daytime the experimental rst trial directional response latencies exposure to the arc of the sun to use the sun as an were greater than the control latencies. Subsequently, we orientation cue. However, there is also some evidence continued training both groups in the afternoon and then indicating that, even with limited exposure to the arc of the tested the pigeons during the morning under complete sun, birds, like insects, can use the sun to orient at any time cloud cover. Both groups displayed random directional of day. We re-examine this issue in a study of compass responses under cloud cover, indicating that the observed orientation in a cue-controlled arena. Two groups of young orientation was based on the visibility of the sun. The data homing pigeons received different exposure to the sun. The indicate that pigeons with limited exposure to the arc of the control group experienced the sun throughout the day; the sun can, like insects, use the sun for orientation at any time experimental group experienced only the apparent descent of day. of the sun. After 8 weeks of sun exposure, we trained both groups in the afternoon to nd food in a specic compass direction in an outdoor arena that provided a view of the sun but not landmarks. We then tested the pigeons in the Key words: pigeon, Columba livia, orientation, homing, sun compass, learning. morning for their ability to use the morning sun as an

Introduction Beginning with the pioneering discoveries of Karl von Frisch with honeybees, Apis mellifera, and Gustav Kramer with starlings, Sturnus vulgaris, it has become clear that a wide variety of species use a time-compensated sun compass in navigation (von Frisch, 1950, 1965; Kramer, 1952a,b). To use the sun as a compass, an organism must compensate for the apparent changes in the azimuth of the sun throughout the day. These compensations are made by relying on an internal sense of time that is integrated with measurements of the changing position of the sun relative to some frame of reference (Braemer, 1960; Dyer and Dickinson, 1994; Hoffmann, 1960; Schmidt-Koenig, 1960, 1990; Schmidt-Koenig et al., 1991). The existence of sun compass orientation is well established in many species. However, in no species do we have a clear understanding of the behavioral strategies used in sun compass orientation (for a review, see Able, 1991). One major unresolved question concerns how much experience with the course of the sun is needed for organisms to learn to use the sun for orientation. Insects do not need complete exposure to the arc of the sun to estimate its position at a time of day when they have never seen the sun (Dyer and Dickinson, 1994; Lindauer, 1957, 1960; Wehner and Mller, 1993). Similarly, Braemer (1960) reported that some species of sh (Lepomis cyanellus, Lepomis gibbosus and Aequidens portalegrensis) were able to compensate correctly for the apparent movement of an articial sun at any time of day even with limited exposure to the natural sun. Both Schmidt-Koenig (1963a) and Hoffmann (1953, 1959, 1960) also provided anecdotal evidence that adult pigeons, Columba livia, and adult and young starlings were able to use the sun compass at a time of day at which they had never seen the sun. However, the prevailing hypothesis in birds, based on the research of Wiltschko and Wiltschko (1980) with young homing pigeons, is that they must have exposure to the entire arc of the sun to develop a sun compass that can be used at any time of day. The conclusions of Wiltschko and Wiltschko (1980) and

2342 C. A. BUDZYNSKI, F. C. DYER AND V. P. BINGMAN

those of Schmidt-Koenig (1963a) and Hoffmann (1953, 1959, 1960) are contradictory. Thus, our study was designed to reexamine the development of the sun compass to determine whether homing pigeons with only partial experience with the course of the sun could use that experience to orient at a time of day when they had never seen the sun. Materials and methods Subjects The subjects were 16 homing pigeons, Columba livia. They were bred at Bowling Green State University and did not see the sun until 5 weeks of age. During the subsequent 8 weeks, we provided the pigeons with food, water and grit ad libitum. During later experimental training, the pigeons were fooddeprived to 85 % of their free-feeding body mass. We provided the pigeons with mixed grain during experimental training. The pigeons were housed in groups at the Bowling Green State University Mercer Road Ecology and Ethology Research Station in Bowling Green, Ohio, USA (8339W, 4123N). Upon self-sufficiency at 5 weeks of age, the pigeons were randomly divided into two groups (for each group, N=8) and placed into two adjacent indoor lofts (2 m2 m3 m each) located within the research station building (ground area 9 m9 m). The six windows (44 cm30 cm) of the research station building were covered with two layers of translucent Plexiglas, the rst 6 mm thick and the second 3 mm thick. The two layers of Plexiglas eliminated direct observation of the suns disk and reduced the penetration of ultraviolet light to less than 1 % of ambient light and the penetration of visible light to less than 10 % of ambient light. Inside the loft, the light levels remained constant throughout the day so it is unlikely that the experimental pigeons could have relied on any direct or indirect information about the path of the sun. Both lofts were connected to outdoor aviaries (3 m2 m2 m each). From the enclosed lofts, we could place the control and experimental pigeons in the outdoor aviaries. The control aviary was situated on the corner of the loft, which provided exposure to the sky from east-southeast to north. The experimental aviary was adjacent to the control aviary, which provided full exposure only to the western half of the sky. Exposure to the course of the sun The control and experimental groups received different exposure to the course of the sun during the 8 weeks before experimental training and during training. The control group lived continuously in the outdoor aviary except in extremely bad weather and could view the sky throughout the day. The experimental group, in contrast, could only view the sky when we placed them each day in the outdoor aviary from local noon (approximately 12:40 h Eastern Standard Time, EST), when the sun was due south, until sunset. During the mornings, the experimental pigeons were held in the indoor loft. Apparatus The testing apparatus was an eight-sided outdoor arena (1 m
Remote manual release line

Start chamber

Food bowl

Fig. 1. A diagram of the apparatus used during experimental training and testing.

in diameter) of a design similar to those successfully employed in studies of sun compass learning in pigeons (Bingman and Jones, 1994; Chappell and Guilford, 1995; Gagliardo et al., 1996; Schmidt-Koenig, 1963a,b; see Fig. 1). Each wall of the arena had a 15 cm2 opening through which a pigeon could reach a metal food cup surrounded by blinders. The ceiling of the arena was clear Plexiglas in which we cut a circular hole in the center. A cylindrical start chamber (30 cm in diameter, 27 cm in height) could be lifted through the hole. The experimenter manually lifted the start chamber with a pulley system to release a pigeon. The arena was level with respect to gravity and was placed on a rotating board so that its orientation could be changed. It was mounted on a table and placed in an open eld that offered the pigeon in the arena no view of surrounding landmarks (Bingman and Jones, 1994). Training We trained both groups between 12:40 h EST and 19:30 h EST from April to October 1998. Each pigeon was trained at different times in the afternoon for 1015 consecutive days when the sun was visible. Once a pigeon had completed its training, we tested it on the morning immediately following its last day of training. The following protocol was similar to the training procedures used by Bingman and Jones (1994). Once fooddeprived, the pigeons were trained in the test arena. We initially placed each pigeon into the chamber for acclimation. Once they had learned to eat from the food cups accessible through the openings on the arena walls, they were trained towards a xed compass direction. We trained four pigeons from each group to receive food reinforcement from the north and four pigeons from each group to receive food reinforcement from the south. During training, we placed food reinforcement into the training direction cup, ensuring that the food was not visible to the pigeon. For each trial, we placed a pigeon into the start chamber and then manually lifted the start

Sun compass orientation in pigeons 2343

chamber. We allowed a pigeon to search the cups until it found the food reinforcement. The experimenter recorded each cup in which the pigeons beak passed through an opening as a directional response. Once the pigeon had obtained reinforcement, it was returned to the start chamber to begin a new trial. We repeated this procedure 20 times per session. After each trial, the experimenter rotated the arena to a new orientation, selected at random, and randomly changed her viewing position. This ensured that the pigeons could not use either landmarks within the arena or the position of the experimenter to learn the training direction. An initial directional response of a trial to the cup in the training direction or to the two adjacent cups was recorded as correct (e.g. if north was the training direction, responses to the northwest, north and northeast cups were all recorded as a correct response). Although the start chamber was designed to minimize variability in the starting position, pigeons occasionally moved within the arena before orienting towards a cup. We recorded the adjacent cups as correct to take into account the variability in starting position from trial to trial. The pigeons were trained until they reached a criterion of 80 % correct responses for ve consecutive sessions in which the rst response of trial 1 was either towards the training direction cup or towards either of the two cups adjacent to the training cup. An experimental session generally lasted approximately 30 min. We trained the pigeons at different times throughout the afternoon to minimize the possibility of them learning to respond to the sun as a xed landmark rather than as a true compass. The group range of the azimuth of the sun during training was 182266 for control pigeons and 184275 for experimental pigeons; hence, the direction of the rewarding food cup relative to the azimuth of the sun varied by as much as 90 from day to day. Test 1: morning orientation After the pigeons had met the performance criterion, we tested them for their ability to use the morning sun to nd the cup in the direction to which they had been trained in the afternoon. The pigeons were tested between 09:00 h EST and 10:00 h EST and received 10 trials without food reinforcement (the group range of the sun azimuth during test 1 was 93124 for control pigeons and 94111 for experimental pigeons). For the data analysis (see below), the rst directional responses of the pigeons were the critical measure of behaviour because, during the non-reinforced test session, a pigeon might change its behaviour after the rst trial because of the absence of reinforcement. However, the pigeons received 10 trials for two reasons. (i) We tried to keep the test session similar to the training sessions, which had multiple trials. (ii) We were curious to determine whether non-reinforced trials would affect the subsequent responses of the pigeons, i.e. would the pigeons change their search strategies because of lack of reinforcement? All other variables were identical to those described for the training sessions. Immediately after a pigeon had chosen a food cup, or after the pigeon had made no response for 5 min, the experimenter returned the pigeon to the start chamber to begin the next trial. Test 2: orientation under overcast skies This experiment was performed to determine whether the results of test 1 were based on sun compass learning or on some other compass mechanism (e.g. magnetic compass or some other directional cue, see Wiltschko and Wiltschko, 1988). After test 1, the pigeons training continued every 4 days during the afternoon as described above. On the rst day of complete cloud cover, we tested the pigeons in the morning as described in test 1. If the orientation in the test arena was based on the use of the sun, then the cloud cover would prevent the birds from searching in the correct compass direction. Data analyses During training and testing, the experimenter recorded the direction of the rst food cup chosen for each trial. The rst trial from each session was used for the initial data analyses. During training, all trials were reinforced. Thus, only the rst trial reected learning across sessions and did not reect within-session learning. Similarly, because the test trials were non-reinforced, the rst trial during a test session was the critical measure of behaviour. During the test session, a pigeon might tend to change its behaviour after the rst trial as a result of the lack of reinforcement. Three types of analysis were undertaken. (i) We examined the directional responses recorded on the rst trial during the last 5 days of training. For each pigeon, a mean vector was computed from the ve directional responses. We calculated a group mean vector for each group using the direction of the mean vector of the individual pigeons. (ii) For test 1, we analyzed the directional response of the rst trial. We generated a mean vector for each group using the directional response of the rst trial of the individual pigeons. (iii) For test 2, we analyzed the directional response for each bird on the rst trial of the test session. We generated a mean vector for each group using the directional response from the rst trial of the individual pigeons. For all three analyses, we used a Rayleigh test to determine whether the distribution of angles for each group differed from uniform, and we also performed between-group comparisons of directional responses using a Watson U2-test (Batschelet, 1981). Ninety-nine per cent condence intervals were computed for each group (Batschelet, 1981) to determine whether the direction of the mean vector of each group differed signicantly from either the training direction or a xed angle direction based on the last day of training. Because a xed angle direction is based on the angle between the training direction and the sun on the last training day (which was different for each bird), the rst trial directional response of each bird was normalized so that condence intervals could be computed (i.e. for each pigeon, the expected xed angle response was normalized to 360 ). Additional analyses on the rst ve test trials of each bird for test 1 and test 2 were computed to determine whether

2344 C. A. BUDZYNSKI, F. C. DYER AND V. P. BINGMAN

continued testing with non-reinforced trials would affect the response of the pigeons. However, one must be cautious in the interpretation of these results because of the absence of reinforcement in these trials. For these analyses, we again used a Rayleigh test to determine whether the distribution of angles for each group differed from uniform, and we also performed between-group comparisons of directional responses using a Watson U2-test (Batschelet, 1981). On the last day of training and during test 1, using a stopwatch, we recorded the latency to the rst directional response for each bird as an additional way of monitoring possible behavioural differences between the control and experimental pigeons during training and testing. With the latency to the directional response as the dependent variable, we used a two-factor within-group analysis of variance (ANOVA) with group (control and experimental) and day (last training day and test 1) as the independent variables to identify any possible differences. Results Training The two groups learned the task equally well. The mean number of training sessions for the control group was 15.500.63 sessions and for the experimental group was 13.380.80 sessions (means S.E.M., t14=2.09, not signicant). The mean directional responses of the rst trials during the last 5 days of training are presented in Fig. 2 and Fig. 3A,B. The rst trial directional responses of the control and experimental pigeons on the last 5 training days were well oriented in the trained compass direction indicating that, from day to day, the birds changed their orientation relative to the sun and their orientation was signicantly different from uniform. We noted no differences between groups in directional responses on the last 5 days of training. Test 1: morning orientation During the morning test session, we expected the control pigeons to select the training compass direction because they had previously received exposure to the course of the sun in the morning. Hence, as shown originally by Kramer (1952a,b), the control pigeons should have been able to compensate for the change in the solar azimuth from afternoon to morning. However, for the experimental pigeons to orient towards the training direction, they would have to estimate the course of the sun in the morning based only on their knowledge of its apparent movement in the afternoon in a manner similar to that of insects (Dyer and Dickinson, 1994; Wehner and Mller, 1993). The directional responses of each individual for the rst trial are presented in Fig. 2 and Fig. 3C,D. Both the experimental and control groups were non-randomly oriented, and the distribution of mean vectors did not differ. To test whether the mean vector of each group was signicantly different from either the normalized training direction or the normalized xed-angle direction, a 95 % condence interval for each
180 135 90 45 0 Angle relative to the sun (degrees) -45 -90 -135 -180 06:00 180 135 90 45 0 -45 -90 -135 -180 06:00 08:00 10:00 12:00 14:00 Time (h) 16:00 18:00 08:00 10:00 12:00 14:00 16:00 18:00

Fig. 2. The initial directional response on the rst trial for the last 5 days of training and test 1 for (A) control pigeons and (B) experimental pigeons. Each directional choice is represented as an angle relative to the sun at the time of training in the afternoon or testing in the morning. Each individual is represented by a different symbol. The solid line in A is the sun azimuth on 30 August 1998 and represents the median sun azimuth experienced by the control pigeons during training and testing. The solid line in B is the sun azimuth on 8 August 1998 and represents the median sun azimuth experienced by the experimental pigeons during training and testing. The error bars represent the predicted xed angle response with the 95 % condence intervals based on the last day of training for control and experimental pigeons. The dashed line indicates the earliest time of training in the afternoon (12:40 h).

group was computed. (i) The mean vectors of the control group and the experimental group did not differ signicantly from the normalized training compass direction (360 ). The condence intervals for the control group was 32640 and that for the experimental group was 832 . (ii) In contrast, the mean vectors of both groups differed signicantly from the xed angle direction (normalized to 360 ). The condence intervals for the control group was 8545 and that for the experimental group was 18532 . For the ve test trial analysis, the mean vector for the control group was 312 and the mean vector length was 0.90 (not taking into account the mean vector length of each individual; P<0.01). The mean vector for the experimental group was 348 and the mean vector length was 0.62 (not taking into account the mean vector length of each individual; P<0.01).

Sun compass orientation in pigeons 2345 A C E

= 1*** r = 0.98

= 326* r = 0.71

= 18 r = 0.40

U2 = NS

U2 = NS

U2 = NS

= 359*** r = 0.96

= 13** r = 0.80

= 225 r = 0.25

Control (trained to north) Experimental (trained to north)

Control (trained to south) Experimental (trained to south)

Fig. 3. The directional response of control and experimental pigeons for the rst trial on the last 5 days of training (A,B), for the rst trial of test 1 (C,D) and for the rst trial of test 2 (E,F). In A and B, each symbol represents the mean directional response and mean vector length for an individual pigeon. In CF, each symbol represents the directional response for an individual pigeon on the rst trial. The heavy solid arrow outside the circle indicates the training direction (the training direction is normalized to 360 for pigeons trained to the south). The arrows within the circles indicate the mean vector for each group. In C and D, the arrows within the circles indicate the mean vector for each group with the 95 % condence intervals (arc). The length of each line approximates the mean vector length with the radius of the circle equal to a mean vector of 1.0. , group mean vector; *P<0.05, **P<0.01, ***P<0.001 with a Rayleigh test. NS, not signicant; r, mean vector length. U2, between-group differences tested with the Watson U2-test.

Both the control and experimental groups were non-randomly oriented, and the distribution of mean vectors did not differ. These results are similar to the results of the analysis of the rst trial directional responses. The group results indicate that, when tested in the morning, control and experimental pigeons were oriented in the true compass direction in which they had been trained and were not simply adopting a xed angle relative to the sun. It should be noted that examination of Fig. 2A indicates that at least one control pigeon may have been relying on a xed angle response when tested in the morning. However, despite the response of this one control pigeon, the data critically demonstrate that the experimental pigeons were able to use the sun compass at a time of day when they had never seen the sun. The results of the ANOVA for the latency to the rst response between the last training day and test 1 are presented in Fig. 4. The ANOVA revealed a signicant interaction

between group and day (F1,14=5.62, P<0.05). Fishers post-hoc analysis revealed no signicant difference between the response latency for the control group and experimental group on the last day of training (P>0.05) and a signicant difference between the response latency for the control group and experimental group on the test day (P<0.05). The difference in response latency between the control and experimental pigeons during test 1 indicates that the two groups responded differently to the morning sun, a difference that presumably reects the lack of prior experience of the experimental birds with the morning sun. Test 2: orientation under overcast skies Test 2 was a control test to determine whether the orientation of the pigeons in the arena was based on some environmental cue other than the sun compass (e.g. the magnetic eld of the earth). The results from the rst trials of test 2 are presented

2346 C. A. BUDZYNSKI, F. C. DYER AND V. P. BINGMAN

150 Latency to first directional response (s) Control Experimental

125

100

75

50

25

0 Last training day Test day Fig. 4. The mean response latency (in s) to the rst directional response of trial 1 for control and experimental pigeons during the last training day and on test 1. Values are means + S.E.M., N=8.

in Fig. 3E,F. The mean vectors of the control group and experimental group for the rst trial data did not differ from uniform. Thus, the orientation of the pigeons recorded during test 2 was dependent on the availability of the sun. Although the analysis of the rst trial directional response was not signicant, the apparent orientation towards the training direction of four of the eight control pigeons was curious. However, for the ve trial test analysis, the mean vector for the control group was 225 and the mean vector length was 0.16 (not taking into account the mean vector length of each individual; P>0.05). Thus, the apparent orientation towards the training direction of these individual pigeons was not evident in the ve trial test analysis. The mean vector for the experimental group was 263 and the mean vector length was 0.15 (not taking into account the mean vector length of each individual; P>0.05). Both the control and experimental groups were randomly oriented, and the distribution of mean vectors did not differ. Discussion In this study, we examined whether young pigeons exposed only to the afternoon sun could generalize the information gained in the afternoon and use the morning sun for orientation. Insects and sh do not need complete exposure to the sun to use it as an orientation mechanism throughout the day (Braemer, 1960; Dyer and Dickinson, 1994; Lindauer, 1957, 1960; Wehner and Mller, 1993). However, there is conicting evidence about the amount of experience birds need to use the sun as an orientation cue. Schmidt-Koenig (1963a) tested sun compass orientation in adult pigeons under the arctic sun in summer using a cuecontrolled arena similar to our own. Two of his three pigeons

oriented in the training direction at a time of day when they had never experienced the sun (at midnight). Hoffmann (1959) obtained similar results with starlings displaced to the arctic and tested under the midnight sun. The results of SchmidtKoenig (1963a) and Hoffmann (1953, 1959, 1960) indicate that birds might be able to use the sun compass at a time of day when they have never seen the sun. However, their research involved mature birds that had received extensive exposure to the diurnal course of the sun in lower latitudes. In addition, the starlings in the study of Hoffmann (1959) may also have seen a portion of the midnight sun before testing. Although Hoffmann (1953, 1960) did examine sun compass orientation in young starlings, they were 12 days old when removed from the nest and may have had some passive exposure to the sun. In a more substantial study, Wiltschko and Wiltschko (1980) raised young homing pigeons with exposure only to the afternoon sun. They ew a control group throughout the day and an experimental group only in the afternoon. They then released both groups in the morning from an unfamiliar location. During the morning, both groups were oriented towards home. Both groups were then subjected to a 6 h slow clock-shift and released in the afternoon (the subjective morning for the pigeons). The control group displayed a 90 clockwise shift in orientation, whereas the experimental pigeons continued to orient towards home. In a subsequent study, Wiltschko et al. (1981) concluded that the experimental pigeons were relying on a magnetic compass to orient, and they suggested that pigeons needed complete exposure to the daytime arc of the sun to use it for orientation throughout the day. Pigeons use both a navigational map and a compass to navigate home. Wiltschko and Wiltschko (1980) conducted experiments that involved homing releases. With respect to understanding the sun compass, homing releases are confounded because the sun compass functions together with the navigational map. Therefore, the apparent failure of their experimental pigeons to use the sun as a compass during homing releases may not be due to the absence of a developed sun compass. Instead, the pigeons may have possessed a functioning sun compass but simply have given priority to their magnetic compass when using compass information in conjunction with their navigational map. In many other contexts, animals will ignore an orientation cue when some other source of directional information is more reliable, but such a result is not evidence that the cue that is ignored is not used under different circumstances (see, for example, Gagliardo et al., 1996). A paradigm that isolates sun compass learning is necessary when examining the development of sun compass orientation. In the present study, we trained and tested young pigeons in an outdoor arena that explicitly isolated sun compass learning from other navigational mechanisms. During training, there were no differences between the control and experimental groups in the mean number of sessions needed to reach the training criterion, and both groups learned to orient in the training direction. The results are consistent with previous

Sun compass orientation in pigeons 2347

studies: birds can use the sun as a compass for orientation when isolated in an experimental arena (Balda and Wiltschko, 1995; Bingman and Jones, 1994; Chappell and Guilford, 1995; Duff et al., 1998; Gagliardo et al., 1996; Hoffmann, 1960; Kramer, 1952a,b; Schmidt-Koenig, 1963a,b; Wiltschko and Balda, 1989). More substantially, the experimental pigeons were able to learn to use the sun compass with only partial experience of the course of the sun. Thus, when the sun compass has been experimentally isolated from other spatial mechanisms, pigeons with exposure only to the afternoon sun are able to generalize this information and use the morning sun to orient. Consequently, pigeons resemble insects and sh (Braemer, 1960; Dyer and Dickinson, 1994; Lindauer, 1957, 1960; Wehner and Mller, 1993), which also develop a sun compass that works at all times of day even after only partial experience with the arc of the sun. However, the results of the present study do not diminish the importance of the ndings of Wiltschko and Wiltschko (1980). During the homing releases, their pigeons may have possessed a fully functioning sun compass. However, the pigeons may have been impaired in integrating the sun compass with the navigational map because they were deprived of ight experience at a time of day when they had not seen the sun. In contrast, our experiment isolated the sun compass from the navigational map and other homing mechanisms. The contrast between our results and those of Wiltschko and Wiltschko (1980) suggests that the development of sun compass orientation in the context of homing may be more complex than previously appreciated. During test 2, both groups of pigeons displayed uniformly distributed directional choices under complete cloud cover. The results indicate that the orientation recorded during test 1 was dependent on the sun. Thus, in the experimental arena used in the present study, young pigeons with limited exposure to the arc of the sun do not rely on a magnetic compass or any other cue to orient when the sun is not visible. Honeybees with only late afternoon exposure to the sun do not compensate for the apparent movement of the sun with perfect accuracy at other times of day. Rather, the bees place the morning sun in the direction of a 180 reverse angle of the azimuth of the sun during training and maintain this estimate for the entire period before noon (approximately east in the study of Dyer and Dickinson, 1994). Do young pigeons with exposure only to the afternoon sun compensate accurately for the movement of the sun during the morning or do they compensate only approximately in the way bees do? Supercially, the data from the experimental pigeons indicate that their compensation is quite accurate in the morning. However, we tested all the pigeons at approximately the same time during the morning (approximately 09:00 h EST) when the sun was roughly in the east. Therefore, if they relied on a rough approximation (e.g. 180 reverse angle of the sun at sunset), as bees do, similar results would have been recorded. Further experiments are necessary to determine whether the sun compass orientation of the experimental pigeons during the morning was fully timecompensated or simply an approximation similar to that used by partially experienced bees. In summary, the data obtained from the experimental pigeons in this study indicate that birds resemble insects with respect to the development of sun compass orientation. As such, the data indicate that some properties of sun compass learning are shared across a broad array of taxonomic groups.

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