Influence of fuel load and weather on timing of nocturnal spring migratory departures in European robins, <Emphasis Type="Italic">Erithacus rubecula</Emphasis>

Time of departure and landing of nocturnal migrants are of great importance for understanding migratory strategy used by birds. It allows us to estimate flying time and hence the distance that migrants cover during a single night. In this paper, I studied the temporal schedule of nocturnal departures of European robins during spring migration. The study was done on the Courish Spit on the Baltic Sea in 1998–2003 by retrapping 51 ringed birds in high mist nets during nocturnal migratory departure. Take-offs of individual birds occurred between the first and tenth hour after sunset (median 176 min after sunset). Departure time was not related to fuel stores at arrival and departure, stopover duration and progress of the season. The results suggest that one reason for temporal variation in take-off time was differential response of European robins with high and low motivation to depart to such triggers as air pressure and its trend. If these parameters reach a certain minimum threshold shortly before sunset, robins with a high migratory motivation take off in the beginning of the night. When air pressure or its trend reaches a maximum, it may trigger to take off later during the night birds with lower initial motivation for departure, including those that have low refuelling efficiency. In regulation of timing of take-offs of robins, an important role is also played by their individual endogenous circadian rhythm of activity which is related to the environment in a complex way.     

The nature of the migration route shapes physiological traits and aerodynamic properties in a migratory songbird

Migration distance is supposed to represent an important selection pressure shaping physiological and morphological properties. Previous work has focussed on this effect, while the importance of ecological barriers in this context has been rarely considered. We studied two subspecies of a migratory songbird, the northern wheatear (Oenanthe oenanthe oenanthe and O. o. leucorhoa L.), on an island in the North Sea. The former subspecies reaches their Scandinavian breeding areas after a short sea crossing, whereas leucorhoa northern wheatears cross the North Atlantic towards Iceland, Greenland or Canada. Physiological traits (fuel deposition rate) and wings’ aerodynamic properties (wing pointedness independent of body size), both affecting migration speed, were hypothesized to be more pronounced in leucorhoa than in oenanthe northern wheatears. Within subspecies, the physiological and aerodynamic properties were hypothesized to explain arrival date at the stopover site with “fast migrants” arriving early. Physiological and aerodynamic properties in leucorhoa northern wheatears lead to a faster and less costly migration, favouring a sea crossing, but in trade-off lower flight manoeuvrability than in oenanthe birds. Wings’ aerodynamic properties affected the seasonal occurrence of leucorhoa females, whereas the physiological traits significantly influenced arrival date in oenanthe individuals. The less risky migration route in oenanthe birds with few short sea crossings may have favoured higher flight manoeuvrability for foraging (less pointed wings), in trade-off an energetically more costly flight. Hence, not the migration distance itself, but the presence/absence of a sea barrier presents an important selection pressure in migratory land birds favouring low flight costs.     

Exaggerated orientation scatter of nocturnal passerine migrants close to breeding grounds: comparisons between seasons and latitudes

Using tracking radars, we investigated the variability of flight directions of long-distance nocturnal passerine migrants across seasons (spring versus autumn migration) and sites at the southern (56° N) and northern (68° N) ends of the Scandinavian Peninsula (Lund versus Abisko). Whilst most migrants at Lund are on passage to and from breeding sites in Fennoscandia, the majority of the migrants at Abisko are close to their breeding sites, and migration at Abisko thus to a large degree reflects initial departure from breeding sites (autumn) or final approach to breeding destinations (spring). The radar data were used to test predictions about differences in orientation and wind drift effects between adult and juvenile birds (a large proportion of autumn migrants consists of juvenile birds on their first journey), between situations far away from or near the goals and between different phases of migration (initial departure, en route passage, final approach to goal). The concentrations (both total and within-night concentrations) of flight directions differed significantly between seasons as well as sites, with the highest concentration at Lund in spring (mean vector length of track directions, r = 0.79) and lowest at Abisko during spring (r = 0.35). Partial wind drift and partial compensation were recorded at Lund, with a similar effect size in spring and autumn, whilst possible wind drift effects at Abisko were obscured by the large directional scatter at this site. The results from Lund support the prediction that the high proportion of juveniles in autumn contributes to increase the directional scatter during this season, whilst there was no support for predictions of differential wind drift effects between seasons and situations with different goal distances. The most striking and surprising result was the exceedingly large scatter of flight directions at Abisko, particularly in spring. We suggest that such an exaggerated scatter may be associated with final approach orientation, where migrants reach their specific goals from all various directions by final navigation within a more wide-ranging goal region. The larger scatter of autumn flight directions at Abisko compared to Lund may be due to exploratory flights in variable directions being more common at initial departure from breeding sites than later during migratory passage. These surprising results highlight the importance of studying and analysing orientation during final approach to (and initial departure from) migratory goals for understanding the orientation systems of migratory birds.     

Assessing the aerodynamic effects of tail elongations in the house martin (Delichon urbica): implications for the initial selection pressures in hirundines

Of the three species of hirundine that breed sympatrically across the U.K., one, the barn swallow, has outer tail feathers elongated into streamers, whereas the other two species, the house martin and the sand martin, do not. The tail streamer of the barn swallow is regarded as a classic example of a sexually selected trait. Recent evidence, however, has suggested that streamers may have evolved largely through natural selection for enhanced flight performance and increased maneuverability. We tested the hypotheses that small streamers (1) increase performance in turning flight, but (2) decrease performance in flight variables related to velocity. We manipulated the lengths of house martin outer tail feathers and measured changes in their free-flight performance, using stereo-video to reconstruct the birds" three-dimensional flight paths. Five flight variables were found to best describe individual variation in flight performance. Of these five, the three variables determining maneuverability predicted that flight performance would be optimized by a 6- to 10-mm increase in the length of the outer tail feathers. In contrast, for mean velocity and mean acceleration, extension of the outer tail feathers appears to have a detrimental effect on flight performance. We suggest that the initial selection pressure for streamers in ancestral short-tailed "barn swallows" was via natural selection for increased maneuverability. In addition, we propose that the benefits of increased maneuverability have differed between hirundines in the past, such that the cost of increasing the length of the outer tail feather has, to date, outweighed the benefits of doing so in streamerless hirundines. Received: 14 February 2000 / Revised: 2 July 2000 / Accepted 18 July 2000     

Arrival timing and seasonal reproductive performance in a long-distance migratory landbird

The date when a landbird migrant arrives on its breeding grounds may have reproductive consequences. Generally, early arriving individuals begin breeding earlier and consequently experience greater seasonal reproductive performance. Here, we describe relationships between arrival timing and seasonal reproductive performance in the American redstart (Setophaga ruticilla), a long-distance passerine migrant, arriving at northerly breeding grounds in Michigans eastern Upper Peninsula. Evidence suggests that both males and females benefited from early arrival at the breeding grounds. Early males appeared to settle on higher quality territories and hatched nestlings sooner than later arrivals. Early females began their clutches early, produced heavier nestlings and possibly laid more eggs than later arrivals. Larger clutches and heavier offspring increase the likelihood of offspring recruiting into the breeding population. The findings of this study point to fitness consequences arising from when a bird arrives at its breeding grounds. These results also have implications for understanding how events occurring during spring migration influence reproductive performance as migratory delays likely influence arrival timing.Communicated by: W. Wiltschko     

Phenotype-dependent arrival time and its consequences in a migratory bird

Arrival times for migratory animals can be viewed as the result of an optimization process of costs and benefits of early arrival, and when the cost and benefit functions of early arrival depend on phenotypic quality, this will result in phenotype-dependent optimal arrival times. This hypothesis was tested for males of the migratory and sexually size-dimorphic barn swallow Hirundo rustica. The major cost of early arrival is poor environmental conditions which resulted in mortality of short-tailed early-arriving males in one year. The major benefits of early arrival are higher mating success, enhanced reproductive success, improved recruitment rates for offspring, and enhanced quality of the mate acquired. Annual variation in male arrival date is related to weather conditions at the breeding grounds, but also to some extent to weather conditions in the African winter quarters. Individual variation in arrival time can be explained by phenotype-dependent cost and benefit functions of early arrival. Male barn swallows with long tail ornaments arrived earlier than short-tailed males. The costs of early arrival should be particularly high under poor environmental conditions and lead to a stronger negative relationship between arrival date and phenotypic quality in years with poor environmental conditions. This prediction was confirmed by a stronger negative relationship between male tail length and date of arrival in years when arrival was relatively late because of poor weather. A female preference for early-arriving males may result in acquisition of good genes for optimal migratory behaviour, if migratory direction and extent have a genetic basis as shown in a number of different bird species.     

Is there a different response to winds during migration between soaring and flapping raptors? An example with the Montagu’s harrier and the lesser kestrel

During migrations, birds have to cope with varying meteorological conditions, which shape their migratory routes and affect their performance. Amongst these, wind is the main meteorological agent influencing behaviour of birds in their migration journeys. Here we analyze the effect of winds during migrations of adult individuals of two raptor species tracked with satellite telemetry, the Montagu’s harrier (Circus pygargus) and the lesser kestrel (Falco naumanni). While harriers use mostly soaring flight, kestrels principally use flapping flight and thus, wind can differently affect these birds. We found that both forward and perpendicular winds significantly affected the movements of the Montagu’s harrier, which were drifted from their intended direction but also took advantage of tailwinds. On the contrary, lesser kestrels moved more regardless of forward winds, despite they were highly drifted by crosswinds. Our results also support that the drifting effect of winds at the onset of the spring migration may explain the loop migration observed for both species, with birds compensating later the effect of crosswinds to arrive to their breeding areas. Results presented here illustrate how winds can differently affect migrating birds according to their flight modes.     

Timing and flight mode of departure in migrating European bee-eaters in relation to multi-scale meteorological processes

Understanding departure decisions of migratory birds and the environmental factors affecting them is important for predicting their distribution, abundance, and arrival times to breeding and wintering areas. In the past, methodological difficulties to obtain fine-scale bird departure and meteorological data have limited testing the multi-scale effects of meteorology on bird departure during migration. We investigated departure timing of European bee-eaters (Merops apiaster) staging in southern Israel, identified their departure flight mode (flapping or soaring) using radio telemetry, and measured local meteorological conditions to study if bird departure was affected by these. Departure timing was examined using a timescale analysis design. The conditions before, during, and after the time of departure were compared using timescales of 24 h, 6 h, 1 h, and 10 min and in relation to bird flight mode. At the between-days timescale, barometric pressure at departure time was significantly lower compared with 2–1 day earlier, whereas temperature at departure was significantly higher compared with 3–2 days earlier. Temperature at departure was also higher compared with 6 h and 3–2 h earlier. Tailwind assistance had no significant effect at any timescale. Soaring birds departed at significantly higher temperature compared with flapping birds. We suggest that bee-eater departure is tuned to the infrequent passage of warm atmospheric depressions at the between-days timescale and with an increasing temperature trend within these days enabling the birds to use energetically cheap soaring flight. We thus suggest that energetic considerations dictate the departure decisions of migrating European bee-eaters.     

Orientation in captive migratory and sedentary Australian silvereyes <Emphasis Type="Italic">Zosterops lateralis</Emphasis> (Zosteropidae)

The orientation of two closely related subspecies of Australian silvereye Zosterops lateralis was studied in captivity over 14 months. Migratory silvereyes Z. lateralis lateralis showed significant directional preferences during the spring and autumn migration periods and also displayed orientated behaviour during the breeding period. In contrast, the non-migratory subspecies Z. lateralis familiaris did not display any significant directional preferences at any time. This is the first time that the orientation behaviour of a migratory and non-migratory subspecies has been compared over the duration of an annual cycle, both during and outside the migratory periods. The results suggest that migratory silvereyes possess an endogenous program determining the timing and direction of autumn and spring migration and that this program is unique to the migratory subspecies. This is also the first comparison of this nature on southern hemisphere birds and demonstrates that the migratory behaviour of southern hemisphere migrants may be more similar to that of northern hemisphere migrants than previously thought.Submitted to Behavioral Ecology and Sociobiology: 23 Jan 2006.     

The relevance of environmental conditions for departure decision changes en route in migrating geese

The timing of life-cycle events crucially influences fitness, particularly in migratory birds, which visit chains of sites with varying seasonality. Here, we used a proportional hazards model to identify local environmental factors, which a long-distance migrant, the Pink-footed Goose (Anser brachyrhynchus), uses for departure decisions on multiple sites along its spring flyway from Denmark via Norway to Svalbard. Our results not only identified day length, local accumulated temperature, and their interaction as likely candidates, but also (more importantly) showed for the first time that their relevance changes en route. The results suggest that the birds switch on a "migratory program" in their wintering grounds, with day length providing general information on time of the year and integrated temperatures providing information on larger scale climate trends. Thereafter, on the stopover sites, local accumulated temperatures allow the geese to infer information on the advancement of spring, which is then used to adjust the speed of progressing northward.     

Influence of body condition and weather on departures of first-year European robins, <Emphasis Type="Italic">Erithacus rubecula</Emphasis>, from an autumn migratory stopover site

How migratory birds decide when to leave a stopover site is important to the understanding of bird migration strategies. Our study looks at how body condition and the weather affect the decision to depart on nocturnal migratory flight. During two autumn migration seasons (2002–2003), we radio tracked 51 first-year European robins, Erithacus rubecula, at a stopover site on the Courish Spit (Eastern Baltic) from the first day after landing until their migratory departure. The tagged robins stopped over for 1–14 days. There was no clear relationship between stopover duration and energetic condition on arrival. Weather conditions (wind, precipitation, and cloud cover) on departure differed measurably between years. In 2002, robins took off mainly under following winds and clear skies. In 2003, there were mainly light head winds and partially cloudy or overcast skies. This could be explained by the year-specific role of weather factors in making the decision to depart. In both years, robins making short (1–2 days) stopovers took off in more varied weather situations than those individuals with long stopovers. This suggests that robins from the former group were more inclined to continue with migration than longer-stay birds that, apart from re-fuelling, could be waiting for favourable weather. The lack of a relationship between stopover duration and body condition and some departures under unfavourable weather conditions suggest that endogenous spatiotemporal programmes may play an important role in controlling stopover duration in robins.     

Male tail streamer length predicts fertilization success in the North American barn swallow (Hirundo rustica erythrogaster)

In socially monogamous species, extra-pair paternity has the potential to increase the variance in male reproductive success, thereby affecting the opportunity for sexual selection on male extravagant ornamentation. In the European barn swallow (Hirundo rustica rustica), the tail streamer length is a sexually selected male ornament and an honest indicator of viability. The North American barn swallow (Hirundo rustica erythrogaster) also shows sexual dimorphism in tail streamer length, but whether this trait holds the same signalling function in this subspecies is a controversial issue, and the available literature is presently scarce. Here, we present data on paternity in the North American barn swallow, including a complete sampling of extra-pair sires in four colonies. We analysed how extra-pair paternity affected the variance in male fertilization success and examined whether male tail streamer (i.e. the outermost tail feather) length correlated with fertilization success (n=86 males). Extra-pair paternity constituted 31% of all offspring and significantly increased the variance in male fertilization success. The number of offspring sired by extra-pair males accounted for almost half of the total variance in male fertilization success. Males with naturally long tail streamers had a higher fertilization success than males with shorter tail streamers, and this pattern was mainly caused by a higher extra-pair success for males with long tail streamers. Males with long tail streamers also paired with early breeding females in prime body condition. These results are consistent with the idea that there is directional sexual selection on male tail streamer length, possibly mediated through male extra-pair mating success or the timing of breeding onset.     

Seasonal modulation of flight speed among nocturnal passerine migrants: differences between short- and long-distance migrants

Migrating birds are expected to fly at higher airspeeds when minimizing time rather than energy costs of their migratory journeys. Spring migration has often been suggested to be more time selected than autumn migration, because of the advantage of early arrival at breeding sites. We have earlier demonstrated that nocturnal passerine migrants fly at higher airspeeds during spring compared to autumn, supporting time-selected spring migration. In this study, we test the hypothesis that seasonal airspeeds are modulated differently between short- and long-distance migrants, because of a stronger element of time selection for autumn migration over long distances. In support of this hypothesis, we demonstrate that the seasonal difference in airspeed is significantly larger (spring airspeed exceeding autumn airspeed by a factor of 1.16 after correcting for the influence of altitude, wind and climb/descent on airspeed) among short-distance compared to long-distance (factor 1.12) migrants. This result is based on a large sample of tracking radar data from 3 years at Falsterbo, South Sweden. Short-distance migrants also tend to fly with more favourable winds during autumn, indicating relaxed time constraints (being able to afford to wait for favourable winds) compared to long-distance migrants. These results indicate surprisingly fine-tuned seasonal modulation of airspeed and responses to wind, associated with behavioural strategies adapted to different levels of time selection pressures during spring and autumn migration.     

Autumn migratory fuelling: a response to simulated magnetic displacements in juvenile wheatears, Oenanthe oenanthe

Recent experiments exposing migratory birds to altered magnetic fields simulating geographical displacements have shown that the geomagnetic field acts as an external cue affecting migratory fuelling behaviour. This is the first study investigating fuel deposition in relation to geomagnetic cues in long-distance migrants using the western passage of the Mediterranean region. Juvenile wheatears (Oenanthe oenanthe) were exposed to a magnetically simulated autumn migration from southern Sweden to West Africa. Birds displaced parallel to the west of their natural migration route, simulating an unnatural flight over the Atlantic Ocean, increased their fuel deposition compared to birds experiencing a simulated migration along the natural route. These birds, on the other hand, showed relatively low fuel loads in agreement with earlier data on wheatears trapped during stopover. The experimental displacement to the west, corresponding to novel sites in the Atlantic Ocean, led to a simulated longer distance to the wintering area, probably explaining the observed larger fuel loads. Our data verify previous results suggesting that migratory birds use geomagnetic cues for fuelling decisions and, for the first time, show that birds, on their first migration, can use geomagnetic cues to compensate for a displacement outside their normal migratory route, by adjusting fuel deposition.     

The strategy of fly-and-forage migration,illustrated for the osprey (<Emphasis Type="Italic">Pandion haliaetus</Emphasis>)

Migrating birds often alternate between flight steps, when distance is covered and energy consumed, and stopover periods, when energy reserves are restored. An alternative strategy is fly-and-forage migration, useful mainly for birds that hunt or locate their prey in flight, and thus, enables birds to combine foraging with covering migration distance. The favourability of this strategy in comparison with the traditional stopover strategy depends on costs of reduced effective travel speed and benefits of offsetting energy consumption during migration flights. Evaluating these cost-benefit effects, we predict that fly-and-forage migration is favourable under many conditions (increasing total migration speed), both as a pure strategy and in combination with stopover behaviour. We used the osprey (Pandion haliaetus) as test case for investigating the importance of this strategy during spring and autumn migration at a lake in southern Sweden. The majority, 78%, of passing ospreys behaved according to the fly-and-forage migration strategy by deviating from their migratory track to visit or forage at the lake, while 12% migrated past the lake without response, and 10% made stopovers at the lake. Foraging success of passing ospreys was almost as good as for birds on stopover. Timing of foraging demonstrated that the birds adopted a genuine fly-and-forage strategy rather than intensified foraging before and after the daily travelling period. We predict that fly-and-forage migration is widely used and important among many species besides the osprey, and the exploration of its occurrence and consequences will be a challenging task in the field of optimal migration.     

Least auklet ornaments: do they function as quality indicators?

Summary In the monogamous least auklet (Aethia pusilla, Alcidae) both males and females have three highly variable ornamental traits (facial plumes, a colourful bill and a knob-like bill ornament) and both sexes perform courtship displays. To assess whether mating preferences could be related to the expression of these ornaments, we performed model presentation experiments in which we varied the bill colour and the size of both the bill ornament and the facial plumes. Auklets reacted to models that had brighter red bills and accentuated facial plumes with more frequent sexual displays than to models with average bills and plumes. We conclude that these two ornamental traits are likely to be favoured by sexual selection through mating preferences. In general, however, ornaments were weak predictors of individual quality. Multiple regression indicated that all ornaments taken together explained a small (R ² = 0.07) but significant proportion of the variability in adult body condition but only in a poor year for reproduction when the birds were in relatively poor body condition and a small proportion of the population bred successfully. The degree of ornamentation was also not related to timing of breeding, chick feeding rate or reproductive success and there was no relationship between adult survival and ornaments. We conclude, therefore, that least auklet ornaments are, at best, weak indicators of quality that may be the result of sexual selection operating only in years when breeding conditions are poor. Offprint requests to: R. Montgomerie     

Social dominance and migratory restlessness in the dark-eyed junco (Junco hyemalis)

Summary I investigated the effect of restricted food and social dominance on nocturnal migratory activity (Zugunruhe) in dark-eyed juncos (Junco hyemalis) in late fall and winter. Highly restricted food tended to increase Zugunruhe in both dominant and subordinate members of pairs, however, subordinates showed significantly more migratory activity than dominants or solitary controls. Further, subordinate birds continued Zugunruhe after dominants and solitary controls had ceased this activity for the remainder of the winter. From mid-December through mid-January when birds had access to food ad lib, migratory activity decreased significantly. These results indicate that toward the end of fall migration, migratory behavior is subject to ecological and social conditions that influence the probability of overwinter survival. Presumably, such a system enables these birds to minimize the distance of migration; at the same time, they are able to track a relatively unpredictable, temperate, winter environment by prolonging migration if necessary.     

Factors influencing the movement biology of migrant songbirds confronted with an ecological barrier

Whether or not a migratory songbird embarks on a long-distance flight across an ecological barrier is likely a response to a number of endogenous and exogenous factors. During autumn 2008 and 2009, we used automated radio tracking to investigate how energetic condition, age, and weather influenced the departure timing and direction of Swainson’s thrushes (Catharus ustulatus) during migratory stopover along the northern coast of the Gulf of Mexico. Most birds left within 1 h after sunset on the evening following capture. Those birds that departed later on the first night or remained longer than 1 day were lean. Birds that carried fat loads sufficient to cross the Gulf of Mexico generally departed in a seasonally appropriate southerly direction, whereas lean birds nearly always flew inland in a northerly direction. We did not detect an effect of age or weather on departures. The decision by lean birds to reorient movement inland may reflect the suitability of the coastal stopover site for deposition of fuel stores and the motivation to seek food among more extensive forested habitat away from the barrier.     

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     

Is wind drift in migrating barnacle and brent geese, Branta leucopsis and Branta bernicla, adaptive or non-adaptive?

Whether migrating birds compensate for wind drift or not has been discussed frequently during the last decades. Observed behaviours seem to differ between species and situations. Even though complete compensation allows a bird to fly the shortest distance during its migration, this is not necessarily the optimal solution in all cases. There are certain situations when drifting with the wind, and thus getting higher groundspeed, will be more beneficial, i.e. adaptive drift. In this article, I analyse flight directions of spring migrating barnacle and brent geese, tracked by radar and optical range finder, and compare these with prevailing winds to see if these birds compensate or drift with the wind. I also use wind data from the days when the trackings were made to construct expected flight paths to see whether wind drift or compensation would be the most beneficial behaviour. The geese were found to drift partially with the wind. The drift effect was concluded to be true partial drift, i.e. not pseudodrift or drift forced by strong winds. The drift was not found to be of any obvious adaptive value, as the geese drifted irrespective of the strategy that would have been most beneficial to them. None of the situations in which drift is predicted to be adaptive was applicable to the birds and migration days in this study. A possible explanation for the observed pattern is that since these birds usually have access to favourable winds during their spring migration, selection pressures for adaptive drift or compensation behaviours might be weak.