Wind selectivity of migratory flight departures in birds

Optimal migration theory predicts that birds minimizing the overall time of migration should adjust stopover duration with respect to the rate of fuel accumulation. Recent theoretical developments also take into account the wind situation and predict that there is a time window (a set of days) during which birds should depart when assisted by winds but will not do so if there are head winds. There is also a final day when birds will depart irrespective of wind conditions. Hence, the wind model of optimal migration theory predicts that birds should be sensitive to winds and that there should be a correlation between departures and winds blowing towards the intended migration direction. We tested this assumption by tracking the departures of radio-tagged passerines during autumn migration in southern Sweden. Our birds were moderately to very fat when released and therefore energetically ready for departure. There was a significant correlation between direction of departure and wind direction. We also found that during days when birds departed there was a significantly larger tail wind component than during days when birds were present but did not depart. Our results show that passerines do take the current wind situation into account when departing on migratory flights. We also briefly discuss possible clues that birds use when estimating wind direction and strength. The inclusion of wind is an important amendment to optimal migration theory of birds and should be explored further. Received: 1 March 1999 / Received in revised form: 4 October 1999 / Accepted: 16 October 1999     

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.     

Flight speeds of birds observed with radar: Evidence for two phases of migratory flight

Summary Horizontal flight speeds relative to the air (air speeds) of about 3,500 birds flying at night over southeastern New York State were measured with a tracking radar, taking special care to obtain accurate local wind measurements. Results were in agreement with previous data (Larkin et al. 1979) indicating that migrating birds sometimes fly at slower speeds than seem consistent with energetically efficient flight.In both spring and fall, about 30% of the birds flew at air speeds of less than 5 m/s (Fig. 5). Two phases of flight were observed, often occurring together on the same night (Figs. 7 and 9). Faster birds generally flew at headings closer to the seasonally appropriate direction and sometimes tended to fly at higher altitudes. Slower birds' headings tended toward the expected direction only part of the time and were more scattered, both between and within nights of observation.The observed relationship between heading, flight speed, and wind direction confirms the following component correlation reported by previous investigators. But the strongest correlation was with relative wind direction rather than the speed of tail winds.Direct visual observations of a representative subsample of the targets confirmed other indications that they were small birds engaged in flapping flight.Nights when a large proportion of birds had air speeds below 5 m/s were often characterized by seasonally unfavorable winds. Nights with a single air speed mode below 8 m/s occurred when winds were unfavorable or when a high-pressure system was centered near the site of observation.A small but appreciable proportion of birds, here called retros, had such low air speeds that, even though heading more or less upwind, they were being carried backward relative to the ground.A number of other variables, including altitude, time of night, time of year, and rate of ascent/descent, sometimes correlated with speed of flight (Table 3).Generally accepted methods of estimating such parameters as birds' headings and wind drift may be seriously in error if the relationships found in the present investigation are characteristic of other geographical areas.     

Pigeon homing: A repetition of the deflector loft experiment

Summary Pigeons brought up in closed cages deflecting the winds to the left or to the right show, on release, a bias of the mean vanishing directions to the left or right of the mean direction of untreated control birds. The results are in full accord with the findings of Baldaccini et al. (1975).Supported by the Deutsche Forschungsgemeinschaft     

Homing pigeons do extract directional information from olfactory stimuli

Summary Two groups of pigeons were kept from fledging time in two cages fully exposed to winds. From time to time, the cage containing experimentals was additionally exposed to an artificial air current coming from a specific direction and carrying a scent of benzaldehyde. When both groups were exposed to benzaldehyde scent during transportation and at the release site, the control birds flew homeward, whereas the experimentals oriented in the direction roughly opposite that from which they were used to perceiving the benzaldehyde at the loft. When benzaldehyde was not applied, experimental pigeons were homeward oriented like controls.     

Correlating seabird movements with ocean winds: linking satellite telemetry with ocean scatterometry

Satellite telemetry studies of the movements of seabirds are now common and have revealed impressive flight capabilities and extensive distributions among individuals and species at sea. Linking seabird movements with environmental conditions over vast expanses of the world’s open ocean, however, remains difficult. Seabirds of the order Procellariiformes (e.g., petrels, albatrosses, and shearwaters) depend largely on wind and wave energy for efficient flight. We present a new method for quantifying the movements of far-ranging seabirds in relation to ocean winds measured by the SeaWinds scatterometer onboard the QuikSCAT satellite. We apply vector correlation (as defined by Crosby et al. in J Atm Ocean Tech 10:355–367, 1993) to evaluate how the trajectories (ground speed and direction) for five procellariiform seabirds outfitted with satellite transmitters are related to ocean winds. Individual seabirds (Sooty Shearwater, Pink-footed Shearwater, Hawaiian Petrel, Grey-faced Petrel, and Black-footed Albatross) all traveled predominantly with oblique, isotropic crossing to quartering tail-winds (i.e., 105–165° in relation to birds’ trajectory). For all five seabirds, entire track line trajectories were significantly correlated with co-located winds. Greatest correlations along 8-day path segments were related to wind patterns during birds’ directed, long-range migration (Sooty Shearwater) as well as movements associated with mega-scale meteorological phenomena, including Pacific Basin anticyclones (Hawaiian Petrel, Grey-faced Petrel) and eastward-propagating north Pacific cyclones (Black-footed Albatross). Wind strength and direction are important factors related to the overall movements that delineate the distribution of petrels at sea. We suggest that vector correlation can be used to quantify movements for any marine vertebrate when tracking and environmental data (winds or currents) are of sufficient quality and sample size. Vector correlation coefficients can then be used to assess population—or species-specific variability and used to test specific hypotheses related to how animal movements are associated with fluid environments.     

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.     

Flight speeds observed with radar,a correction: slow “birds” are insects

Summary Based upon new data and re-analysis of previous data, some results concerning migration of birds at night are now attributed to insects. In particular, slowly flying radar targets over land (Larkin and Thompson 1980) and, by analogy, over the ocean (Larkin et al. 1979) are shown to lack wing beat patterns of birds and the appearance of birds when viewed with a radar-controlled high-power telescope and spotlamp. The insect targets overlap with birds in the amount of radar echo returned, are abundant in warmer months, and fly even in March and November in temperate North America. Other previous results are largely substantiated; however, a previous report (Larkin 1980) that migrating birds can detect the wind direction over the ocean is not substantiated.     

Mass-based depth and velocity scales for gravity currents and related flows

Gravity driven flows on inclines can be caused by cold, saline or turbid inflows into water bodies. Another example are cold downslope winds, which are caused by cooling of the atmosphere at the lower boundary. In a well-known contribution, Ellison and Turner (ET) investigated such flows by making use of earlier work on free shear flows by Morton, Taylor and Turner (MTT). Their entrainment relation is compared here with a spread relation based on a diffusion model for jets by Prandtl. This diffusion approach is suitable for forced plumes on an incline, but only when the channel topography is uniform, and the flow remains supercritical. A second aspect considered here is that the structure of ET’s entrainment relation, and their shallow water equations, agrees with the one for open channel flows, but their depth and velocity scales are those for free shear flows, and derived from the velocity field. Conversely, the depth of an open channel flow is the vertical extent of the excess mass of the liquid phase, and the average velocity is the (known) discharge divided by the depth. As an alternative to ET’s parameterization, two sets of flow scales similar to those of open channel flows are outlined for gravity currents in unstratified environments. The common feature of the two sets is that the velocity scale is derived by dividing the buoyancy flux by the excess pressure at the bottom. The difference between them is the way the volume flux is accounted for, which—unlike in open channel flows—generally increases in the streamwise direction. The relations between the three sets of scales are established here for gravity currents by allowing for a constant co-flow in the upper layer. The actual ratios of the three width, velocity, and buoyancy scales are evaluated from available experimental data on gravity currents, and from field data on katabatic winds. A corresponding study for free shear flows is referred to. Finally, a comparison of mass-based scales with a number of other flow scales is carried out for available data on a two-layer flow over an obstacle. Mass-based flow scales can also be used for other types of flows, such as self-aerated flows on spillways, water jets in air, or bubble plumes.     

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.     

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.     

Gap Crossing Decisions by Forest Songbirds during the Post-Fledging Period

Gaps in forest cover, created by agriculture, forestry, and other anthropogenic activities, are assumed to impede the movements of many forest songbirds. Little is known, however, about the reluctance of different species of birds to cross habitat gaps. We studied this by inducing birds in the post-fledging period to cross gaps of varying widths and to choose between routes through woodland or across open areas by attracting them to a recording of mobbing calls by Chickadees (  Parus atricapillus). In 278 experiments conducted in boreal forest and agricultural landscapes near Québec city, 157 birds or flocks of birds of five species were attracted. Overall, birds were twice as likely to travel through 50 m of woodland than through 50 m in the open to reach the recording. When given a choice of traveling through woodland or across a gap, the majority of respondents preferred woodland routes, even when they were three times longer than shortcuts in the open. However, species differed greatly in their response to gaps. Our results show that woodland links significantly facilitate movements of birds across fragmented landscapes.     

Northeastward reverse migration of birds over Nova Scotia,Canada, in autumn

Summary During 4 seasons of study, small numbers of birds flew overland to the NNE-E, counter to the main SW migration, intermittently throughout the autumn (31 July – 16 November). Radars also detected overwater reverse migration (RM) from New England to Nova Scotia and from Nova Scotia toward Newfoundland. RM occurred at all hours of the day and night, especially when few birds were migrating SW. Most but not all cases of RM occurred with following SW winds. RM was more common with cloud and/or poor visibility than in fair weather, but was not restricted to cloudy occasions. Mean tracks were correlated with wind direction, but were not consistently downwind. Tracks tended to be closer to downwind in early than in late autumn, and on clear than on overcast nights. Dispersion in tracks was not discernibly related to weather variables, time of day or night, or magnetic disturbance. Eleven hypotheses concerning the reasons for reverse migration in autumn are evaluated; cases of RM recorded in this study are attributable to at least three of these hypotheses (late summer dispersal, hurricanes, dawn reorientation toward coast) and possibly to several others.     

Homing behavior of wood thrushes (Hylocichla mustelina)

Summary Using the wood thrush (Hylocichla mustelina) as an example of a typical nocturnal migrant, we employed radio telemetry to follow breeding birds during the course of homing movements following displacements. Seven thrushes were displaced over distances of 6.5 to 17.3 km in a variety of directions from their nesting territories. The thrushes moved in a series of short flights (mean = 2.1 km) performed primarily at dawn. Consequently, the birds took several days to home from even these relatively short displacements. Thrushes flew under clear and solid overcast skies and even in light rain. The pooled individual flights of the birds were significantly oriented in the homeward direction (Fig. 2). Their orientation relative to home did not improve significantly as they progressed toward the goal. Three of the birds were documented to have returned to their home territories. The detailed tracks of the birds preclude the possibility that they homed by random search.     

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     

Trans-Sahara migrants select flight altitudes to minimize energy costs rather than water loss

Meteorological conditions influence strongly the energy and water budget of birds. By adjusting their flights spatially and temporally with respect to these conditions, birds can reduce their energy expenditure and water loss considerably. By radar, we quantified songbird migration across the western Sahara in spring and autumn. There autumn migrants face the trade-off between (a) favorable winds combined with hot and dry air at low altitudes and (b) unfavorable winds combined with humid and cold air higher up. Thus, it can be tested whether birds may chose altitudes to minimize water loss instead of energy expenditure. We predicted optimal flight altitudes with respect to water loss and energy expenditure based on a physiological flight model when crossing the western Sahara and compared these model predictions spatially and temporally with measured songbird densities. The model aiming for minimal water consumption predicted a mean flight altitude of 3,400 m under autumn conditions. However, 64% of the nocturnal songbird migration flew at altitudes below 1,000 m above ground level profiting from tailwind. This preference for tailwind in autumn, despite the hot and dry air, emphasizes the importance of energy savings and diminishes the significance of possible water stress for the selection of flight altitude. Nevertheless, during daytime, high energy expenditure due to air turbulences and water loss due to warmer air and direct solar radiation prevent songbirds from prolonging their nocturnal flights regularly into the day. Birds crossing the Sahara save water by nocturnal flights and diurnal rests.     

Corticosterone levels in relation to migratory readiness in red-eyed vireos (<Emphasis Type="Italic">Vireo olivaceus</Emphasis>)

We examined the relationship between plasma levels of corticosterone and the migratory activity and directional preference of red-eyed vireos during fall migration at the northern coast of the Gulf of Mexico. Corticosterone is thought to play a role in physiological and behavioural processes before, during, and after long-distance migratory flights. An increase in corticosterone at the onset of migratory flights can be expected in birds that are energetically prepared to migrate in a seasonally appropriate southerly direction. Red-eyed vireos ( Vireo olivaceus) were tested in orientation cages under clear twilight skies. Just prior to the orientation experiments, blood was sampled to assay baseline corticosterone levels. Average corticosterone level for all birds was 22.8 ng/ml. Red-eyed vireos with higher than average baseline levels of corticosterone were significantly more active in orientation cages compared to birds with lower levels of corticosterone. Moreover, birds with higher than average levels oriented in a southwesterly direction, which is consistent with a trans-Gulf flight, whereas individuals with levels below average showed a NNW mean direction. Although there was no significant difference in baseline levels of corticosterone between fat and lean birds, individual mass loss between capture and test was negatively correlated with corticosterone levels. Results from this study clearly demonstrate that corticosterone influences departure decisions and the choice of direction during migration.     

The air speed of migrating birds and its relationship to the wind

Summary This paper deals with the flight speeds of migrating birds observed by radar over a Swiss Alpine pass. The distributions of air speeds for different classes of birds and the influence of the wind on air speeds were investigated. Our findings differ greatly from observations of bird migration over the North Atlantic Ocean and the North American continent. Our data reveal: (1) that the air speeds of the bulk of migrating birds were in the range of 8–18 m/s and that the amount of slow flyers (air speed below 5 m/s) was less than 5%; (2) that there was an obvious influence of the wind on air speeds and, moreover, the data showed a distinctly variable compensatory behavior among different bird classes; (3) that the wind component in the direction of the bird's heading was a better predictor for air speed than the wind direction. Although we do not speculate about possibilities and mechanisms of wind detection, a simple argument shows how birds could estimate wind directions if they do use the moving surface as a reference system.     

Regional-scale meteorological characteristics of the Vento Norte phenomenon observed in Southern Brazil

Environmental Fluid Mechanics - The sudden increase in air temperature associated with strong gusty winds of northerly direction is a phenomenon occasionally observed during the cold season in the...     

Effects of Roads on Movements by Understory Birds in Mixed-Species Flocks in Central Amazonian Brazil

Abstract: Roads through tropical forest create linear disturbances that have unknown consequences for forest birds. We studied how a narrow, rarely used road through otherwise undisturbed Amazonian forest affected the movements and area requirements of understory birds that form mixed-species flocks. Differences in road maintenance led to two distinct treatments along the same road. Trees along the "closed" road formed a partial canopy connecting the two sides of the road, although the roadway itself was kept open. The "open" road was regularly maintained, making a complete opening 10–30 m wide.We followed 15 flocks, 5 each in interior forest, along the open road, and along the closed road. These flocks were led by Thamnomanes antshrikes, and each flock had a discreet, permanent territory. Flock territory size (mean = 8.5 ha) did not differ among the three locations. The open road formed the territorial boundary for all five flocks, although birds moved within a few meters of the edge of the road. The closed road was less of a barrier: 2 of 5 flocks used both sides of the closed road. Playback experiments showed that flocks readily crossed the closed road to approach agonistic vocalizations. Along the open road, even though birds responded to playback by becoming agitated and moving to the extreme edge of the roadside vegetation, they were less likely to cross the road and did so only after a longer duration of playback. Our results suggest that flocks respond to a road as they would to a long linear gap. They use the vegetation along the edges of the road, but because they are unwilling to cross the open area, it becomes a flock territory boundary. Similarly, as in forest gaps, successional change along the closed road produced suitable habitat for flocks. Although this suggests that roads are a trivial problem, we caution that this result applies only to narrow roads that are not accompanied by deforestation or other disturbance.