The influence of weather on the flight altitude of nocturnal migrants in mid‐latitudes

By altering its flight altitude, a bird can change the atmospheric conditions it experiences during migration. Although many factors may influence a bird's choice of altitude, wind is generally accepted as being the most influential. However, the influence of wind is not clearly understood, particularly outside the trade‐wind zone, and other factors may play a role. We used operational weather radar to measure the flight altitudes of nocturnally migrating birds during spring and autumn in the Netherlands. We first assessed whether the nocturnal altitudinal distribution of proportional bird density could be explained by the vertical distribution of wind support using three different methods. We then used generalized additive models to assess which atmospheric variables, in addition to altitude, best explained variability in proportional bird density per altitudinal layer each night. Migrants generally remained at low altitudes, and flight altitude explained 52 and 73% of the observed variability in proportional bird density in spring and autumn, respectively. Overall, there were weak correlations between altitudinal distributions of wind support and proportional bird density. Improving tailwind support with height increased the probability of birds climbing to higher altitude, but when birds did fly higher than normal, they generally concentrated around the lowest altitude with acceptable wind conditions. The generalized additive model analysis also indicated an influence of temperature on flight altitudes, suggesting that birds avoided colder layers. These findings suggested that birds increased flight altitudes to seek out more supportive winds when wind conditions near the surface were prohibitive. Thus, birds did not select flight altitudes only to optimize wind support. Rather, they preferred to fly at low altitudes unless wind conditions there were unsupportive of migration. Overall, flight altitudes of birds in relation to environmental conditions appear to reflect a balance between different adaptive pressures.     

Towards a new understanding of migration timing: slower spring than autumn migration in geese reflects different decision rules for stopover use and departure

According to migration theory and several empirical studies, long‐distance migrants are more time‐limited during spring migration and should therefore migrate faster in spring than in autumn. Competition for the best breeding sites is supposed to be the main driver, but timing of migration is often also influenced by environmental factors such as food availability and wind conditions. Using GPS tags, we tracked 65 greater white‐fronted geese Anser albifrons migrating between western Europe and the Russian Arctic during spring and autumn migration over six different years. Contrary to theory, our birds took considerably longer for spring migration (83 days) than autumn migration (42 days). This difference in duration was mainly determined by time spent at stopovers. Timing and space use during migration suggest that the birds were using different strategies in the two seasons: In spring they spread out in a wide front to acquire extra energy stores in many successive stopover sites (to fuel capital breeding), which is in accordance with previous results that white‐fronted geese follow the green wave of spring growth. In autumn they filled up their stores close to the breeding grounds and waited for supportive wind conditions to quickly move to their wintering grounds. Selection for supportive winds was stronger in autumn, when general wind conditions were less favourable than in spring, leading to similar flight speeds in the two seasons. In combination with less stopover time in autumn this led to faster autumn than spring migration. White‐fronted geese thus differ from theory that spring migration is faster than autumn migration. We expect our findings of different decision rules between the two migratory seasons to apply more generally, in particular in large birds in which capital breeding is common, and in birds that meet other environmental conditions along their migration route in autumn than in spring.     

Flight speeds of swifts (Apus apus): seasonal differences smaller than expected

We have studied the nocturnal flight behaviour of the common swift (Apus apus L.), by the use of a tracking radar. Birds were tracked from Lund University in southern Sweden during spring migration, summer roosting flights and autumn migration. Flight speeds were compared with predictions from flight mechanical and optimal migration theories. During spring, flight speeds were predicted to be higher than during both summer and autumn due to time restriction. In such cases, birds fly at a flight speed that maximizes the overall speed of migration. For summer roosting flights, speeds were predicted to be lower than during both spring and autumn since the predicted flight speed is the minimum power speed that involves the lowest energy consumption per unit time. During autumn, we expected flight speeds to be higher than during summer but lower than during spring since the expected flight speed is the maximum range speed, which involves the lowest energy consumption per unit distance. Flight speeds during spring were indeed higher than during both summer and autumn, which indicates time-selected spring migration. Speeds during autumn migration were very similar to those recorded during summer roosting flights. The general result shows that swifts change their flight speed between different flight behaviours to a smaller extent than expected. Furthermore, the difference between flight speeds during migration and roosting among swifts was found to be less pronounced than previously recorded.     

Can wind help explain seasonal differences in avian migration speed?

A bird's ground speed is influenced by the wind conditions it encounters. Wind conditions, although variable, are not entirely random. Instead, wind exhibits persistent spatial and temporal dynamics described by the general circulation of the atmosphere. As such, in certain geographical areas wind's assistance (or hindrance) on migratory flight is also persistent, being dependent upon the bird's migratory direction in relation to prevailing wind conditions. We propose that, considering the western migration route of nocturnal migrants through Europe, winds should be more supportive in spring than in autumn. Thus, we expect higher ground speeds, contributing to higher overall migration speeds, in spring. To test whether winds were more supportive in spring than autumn, we quantified monthly wind conditions within western Europe relative to the seasonal direction of migration using 30 years (1978–2008) of wind data from the NCEP/NCAR Reanalysis dataset. We found that supporting winds were significantly more frequent for spring migration compared to autumn and up to twice as frequent at higher altitudes. We then analyzed three years (2006–2008) of nocturnal migratory ground speeds measured with radar in the Netherlands which confirmed higher ground speeds in spring than autumn. This seasonal difference in ground speed suggests a 16.9% increase in migration speed from autumn to spring. These results stress the importance of considering the specific wind conditions experienced by birds when interpreting migration speed. We provide a simple methodological approach enabling researchers to quantify regional wind conditions for any geographic area and time period of interest.     

Navigating north: how body mass and winds shape avian flight behaviours across a North American migratory flyway

The migratory patterns of birds have been the focus of ecologists for millennia. What behavioural traits underlie these remarkably consistent movements? Addressing this question is central to advancing our understanding of migratory flight strategies and requires the integration of information across levels of biological organisation, e.g. species to communities. Here, we combine species‐specific observations from the eBird citizen‐science database with observations aggregated from weather surveillance radars during spring migration in central North America. Our results confirm a core prediction of migration theory at an unprecedented national scale: body mass predicts variation in flight strategies across latitudes, with larger‐bodied species flying faster and compensating more for wind drift. We also find evidence that migrants travelling northward earlier in the spring increasingly compensate for wind drift at higher latitudes. This integration of information across biological scales provides new insight into patterns and determinants of broad‐scale flight strategies of migratory birds.     

Flight responses by a migratory soaring raptor to changing meteorological conditions

Soaring birds that undertake long-distance migration should develop strategies to minimize the energetic costs of endurance flight. This is relevant because condition upon completion of migration has direct consequences for fecundity, fitness and thus, demography. Therefore, strong evolutionary pressures are expected for energy minimization tactics linked to weather and topography. Importantly, the minute-by-minute mechanisms birds use to subsidize migration in variable weather are largely unknown, in large part because of the technological limitations in studying detailed long-distance bird flight. Here, we show golden eagle (Aquila chrysaetos) migratory response to changing meteorological conditions as monitored by high-resolution telemetry. In contrast to expectations, responses to meteorological variability were stereotyped across the 10 individuals studied. Eagles reacted to increased wind speed by using more orographic lift and less thermal lift. Concomitantly, as use of thermals decreased, variation in flight speed and altitude also decreased. These results demonstrate how soaring migrant birds can minimize energetic expenditures, they show the context for avian decisions and choices of specific instantaneous flight mechanisms and they have important implications for design of bird-friendly wind energy.     

Geographic comparison of seasonal migration events of the large white butterfly,Pieris brassicae

1. The multivoltine butterfly Pieris brassicae (Linnaeus) inhabits most climatic zones in Europe and Asia. Its regional populations are adapted to the respective climatic conditions and hibernate in those regions. Yet how these adaptations are stabilised in spite of the species' regular migration events is unclear. 2. In this study, significant differences were found in the preferred flight direction of P. brassicae depending on the season and the butterfly's geographic origin. The individual flight behaviours indicated that flight direction is a heritable character. 3. The bivoltine population from northern Germany undertook a typical return flight in the second generation of the year. This reverse flight was induced by the developmental mode of the caterpillars. A diapause in the pupal stage triggered a northward migration, and a non‐diapause a southward migration. 4. In contrast to bivoltine populations, the second generation of multivoltine populations maintained the direction of the hibernating generation. 5. A unique situation was determined for the population from the northern coast of Spain. The North Sea seemed to be a migration barrier that forced an adaptation, with the first generation flying southeastward, and the second generation northeastward. 6. The flight pattern of at least some of the investigated populations suggests that, after their yearly migration, the descendants of a local population are able to reach the home range of their ancestors again.     

Seasonal differences in energy requirements of Garden Warblers Sylvia borin migrating across the Sahara desert

The Sahara desert acts as an ecological barrier for billions of passerine birds on their way to and from their African wintering areas. The Garden Warbler Sylvia borin is one of the most common migrants involved. We used body mass of this species from Greece in autumn and spring to simulate the desert crossing and to assess how body mass relates to fuel requirement. The flight range estimates were adjusted to the seasonal extent of the desert, 2200 km in autumn and about 2800 km in spring. In autumn, with an average fuel load of about 100% of body mass without fuel, birds were not able to cross the desert in still air, but northerly winds prevail during September and with the average wind assistance only one in 14 was predicted to fail to make the crossing. Body mass data from spring, after the desert crossing, was used to estimate departure body mass from south of the desert. The average wind assistance in spring is close to zero and departure body mass of the average bird arriving at Antikythira, a small Greek island, under such conditions was estimated to be 34.6 g, which corresponded to a fuel load of 116%. Calculations based on 1% body mass loss per hour of flight showed slightly larger body mass loss than that calculated from flight range estimates. The results suggest that passerine birds about to cross the eastern part of the Sahara desert need to attain a larger fuel load in spring than in autumn.     

Flight directionality of Tribolium castaneum soon after take‐off under glasshouse and field conditions

Flight directionality of the rust‐red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), was investigated under glasshouse and field conditions using sticky traps placed around dense experimental infestations of T. castaneum derived from field‐collected samples. Although beetles of this species are known to fly quite readily, information on flight of beetles away from grain resources is limited. Under still glasshouse conditions, T. castaneum does not demonstrate strong horizontal or vertical trajectories in their initial flight behaviour. Flight was significantly directional in half of the replicates, but trapped beetles were only weakly concentrated around the mean direction of flight. In the field, by contrast, emigration of T. castaneum was strongly directional soon after flight initiation. The mean vector lengths were generally >0.5 which indicates that trapped beetles were strongly concentrated around the calculated mean flight direction. A circular‐circular regression of mean flight vs. mean downwind direction suggested that flight direction was generally correlated with downwind direction. The mean height at which T. castaneum individuals initially flew was 115.4 ± 7.0 cm, with 58.3% of beetles caught no more than 1 m above the ground. The height at which beetles were trapped did not correlate with wind speed at the time of sampling, but the data do indicate that wind speed significantly affected T. castaneum flight initiation, because no beetles (or very few; no more than three) were trapped in the field when the mean wind speed was above 3 m s^?1. This study thus demonstrates that wind speed and direction are both important aspects of flight behaviour of T. castaneum, and therefore of the spatio‐temporal dynamics of this species.     

南京地区棉蚜的飞行活动节律及其飞行能力

昆虫的飞行活动规律及飞行能力是研究其能否迁飞的基础。通过采用春季到秋季20 m高空黄盆诱蚜、高空所诱蚜和春季木槿树上有翅蚜的卵巢解剖,以及春夏秋三季田间有翅蚜的吊飞试验等方法,研究了南京地区棉蚜Aphis gossypii的飞行活动节律和飞行能力。结果表明,有翅棉蚜的日羽化高峰出现在19:00～20:00。2001年南京地区棉蚜的春、秋两季迁飞高峰分别在5月8日和10月25日。5月份高空诱集的棉蚜中,95.7%个体的卵巢小管数在7条以下,而木槿上羽化后1天的有翅蚜中有35.2%个体的卵巢小管数在7条以上;高空诱蚜和木槿上蚜的平均卵巢小管数存在极显著差异,分别为3.94±1.65和5.88±1.92。8月中下旬棉田棉蚜存在低空飞行行为,并且出现飞行高峰时有翅蚜的卵巢小管数平均在6条以下,超过6条则停止飞行。羽化后1～2天有翅棉蚜吊飞个体的飞行比率和平均飞行距离表现为春、秋季显著大于夏季,三季的最长飞行距离分别为3.89 km、6.15 km和1.44 km。     

High Altitude Bird Migration at Temperate Latitudes: A Synoptic Perspective on Wind Assistance

At temperate latitudes the synoptic patterns of bird migration are strongly structured by the presence of cyclones and anticyclones, both in the horizontal and altitudinal dimensions. In certain synoptic conditions, birds may efficiently cross regions with opposing surface wind by choosing a higher flight altitude with more favourable wind. We observed migratory passerines at mid-latitudes that selected high altitude wind optima on particular nights, leading to the formation of structured migration layers at varying altitude up to 3 km. Using long-term vertical profiling of bird migration by C-band Doppler radar in the Netherlands, we find that such migration layers occur nearly exclusively during spring migration in the presence of a high-pressure system. A conceptual analytic framework providing insight into the synoptic patterns of wind assistance for migrants that includes the altitudinal dimension has so far been lacking. We present a simple model for a baroclinic atmosphere that relates vertical profiles of wind assistance to the pressure and temperature patterns occurring at temperate latitudes. We show how the magnitude and direction of the large scale horizontal temperature gradient affects the relative gain in wind assistance that migrants obtain through ascending. Temperature gradients typical for northerly high-pressure systems in spring are shown to cause high altitude wind optima in the easterly sectors of anticyclones, thereby explaining the frequent observations of high altitude migration in these synoptic conditions. Given the recurring synoptic arrangements of pressure systems across temperate continents, the opportunities for exploiting high altitude wind will differ between flyways, for example between easterly and westerly oceanic coasts.     

Flight of the bluegreen aphid Acyrthosiphon kondoi Shinji (Homoptera: Aphididae) I. Flight periodicity and phenological relationships

Abstract

Seasonal flight patterns of the bluegreen aphid (BGA), Acyrthosiphon kondoi Shinji (Homoptera: Aphididae) were studied. BGA had flight peaks twice a year in spring and autumn, BGA flight appeared to be diurnal. The main period of activity extended from 5 am to 7.30 pm.

The accumulated spring and autumn flight peaks lasted 422 Degree Days (DD) and 351 DD (2.9 and 2.4 BGA generations) respectively. Accumulated DD between the median of spring and autumn flights, and the autumn and following spring flight, was 1667 and 1676 DD; this corresponds to 11.5 and 11.6 BGA generations respectively. The use of DD for predicting BGA flight is discussed.     

The exception to the rule: retreating ice front makes Bewick's swans Cygnus columbianus bewickii migrate slower in spring than in autumn

In the vast majority of migratory bird species studied so far, spring migration has been found to proceed faster than autumn migration. In spring, selection pressures for rapid migration are purportedly higher, and migratory conditions such as food supply, daylength, and/or wind support may be better than in autumn. In swans, however, spring migration appears to be slower than autumn migration. Based on a comparison of tundra swan Cygnus columbianus tracking data with long‐term temperature data from wheather stations, it has previously been suggested that this was due to a capital breeding strategy (gathering resources for breeding during spring migration) and/or to ice cover constraining spring but not autumn migration. Here we directly test the hypothesis that Bewick's swans Cygnus columbianus bewickii follow the ice front in spring, but not in autumn, by comparing three years of GPS tracking data from individual swans with concurrent ice cover data at five important migratory stop‐over sites. In general, ice constrained the swans in the middle part of spring migration, but not in the first (no ice cover was present in the first part) nor in the last part. In autumn, the swans migrated far ahead of ice formation, possibly in order to prevent being trapped by an early onset of winter. We conclude that spring migration in swans is slower than autumn migration because spring migration speed is constrained by ice cover. This restriction to spring migration speed may be more common in northerly migrating birds that rely on freshwater resources.     

粘虫成虫在气流场中飞行行为的观察研究

通过改进悬吊测飞技术、室内风洞和野外雷达相结合的观测方法,研究了粘虫在气流场中的飞行行为特征及其与气流的关系。直筒风洞自由飞行观测的结果表明,粘虫蛾对气流有明显行为反应,表现为头部迎风起飞和迎风飞行的特性;在3.0～5.5 m/s风速下,有92%～94% 的个体可一次逆风飞行通过2 m长的风洞;当风速≥6.0 m/s时,有71.9%的蛾子沿螺旋状的飞行轨迹逆风通过风洞。环形风洞悬吊飞行测试的结果表明,粘虫可逆风飞行的最大风速为7.2 m/s;在风速≤4 m/s条件下,90%以上个体头部迎风飞行或头部朝向与风向成一定的夹角,侧逆风飞行。 雷达观测发现粘虫在空中迁飞过程中具有成层现象,并有较强的秋季回迁定向行为,其头部总是朝向西南;迁飞的最终位移与风向及风速大小有关,迁飞位移速度是飞行速度与风速的矢量和。     

Adverse wind conditions during northward Sahara crossings increase the in‐flight mortality of Black‐tailed Godwits

Long‐distance migratory flights are predicted to be associated with higher mortality rates when individuals encounter adverse weather conditions. However, directly connecting environmental conditions experienced in‐flight with the survival of migrants has proven difficult. We studied how the in‐flight mortality of 53 satellite‐tagged Black‐tailed Godwits (Limosa limosa limosa) during 132 crossings of the Sahara Desert, a major geographical barrier along their migration route between The Netherlands and sub‐Saharan Africa, is correlated with the experienced wind conditions and departure date during both southward and northward migration. We show that godwits experienced higher wind assistance during southward crossings, which seems to reflect local prevailing trade winds. Critically, we found that fatal northward crossings (15 deaths during 61 crossings) were associated with adverse wind conditions. Wind conditions during migration can thus directly influence vital rates. Changing wind conditions associated with global change may thus profoundly influence the costs of long‐distance migration in the future.     

Migratory and targeted flight in seasonal forms of the black bean aphid, Aphis fabae

ABSTRACT. Adult winged forms of Aphis fabae Scop., the spring migrants (fundatrigeniae; alate virginoparae), the summer migrants (alate virginoparae) and the autumn migrants (gynoparae), were flown in an automated vertical wind tunnel, with a small green target presented every 60 s. On average, targeted flight developed in the spring and summer migrants after 19 and 15 min, respectively. However, in autumn migrants targeted flight occurred after 184 min of flight. No evidence of target approach was found up to 30 min prior to this time, although some movement towards the target was observed in the 30 min before the first response of autumn migrants, suggesting that a gradual change from migratory to targeted flight occurred. No correlation was found between aphid weight and the time to first response for spring or summer migrants, but a positive correlation was found for autumn migrants. Flight patterns of autumn migrants, in the absence of a target, are described and compared with flight patterns previously reported for summer migrants.     

Flight speeds and climb rates of Brent Geese: mass-dependent differences between spring and autumn migration

Aerodynamic theories of bird flight predict that horizontal flight speed will increase with increasing load whereas vertical flight speed will decrease. Horizontal flight speed for birds minimizing overall time on migration is predicted to be higher than flight speed for birds minimizing energy expenditure. In this study we compare flight speeds of Brent Geese Branta b. bernicla recorded by tracking radar and optical range finder during spring and autumn migration in southernmost Sweden, testing the above-mentioned predictions. Geese passing Sweden in spring are substantially heavier than in autumn and there might also be a stronger element of time-selection in spring than in autumn. Recorded airspeeds were significantly higher in spring (mean 19.0 m s⁻¹) than in autumn (mean 17.3 m s⁻¹), the average difference being slightly larger than predicted due to the mass difference alone. The effects on airspeed of wind, vertical speed, flock size and altitude were also analysed, but none of these factors could explain the seasonal difference in airspeed. Hence, the results support the hypothesis of mass-dependent flight speed adjustment. The difference between the two seasons was not large enough to corroborate the hypothesis of a stronger element of time-selection in spring, but this hypothesis cannot be rejected. Vertical flight speeds were lower in spring than in autumn, supporting a negative effect of load on birds' flight power margin.     

Flocking behaviour in the twilight ascents of Common Swifts Apus apus

Among the many unique flight behaviours of Common Swifts Apus apus, the most puzzling may be their ascents to high altitudes during both dusk and dawn. Twilight ascents have been hypothesized to be functionally related to information acquisition, including integration of celestial orientation cues, high‐altitude visual landmarks and sampling of weather conditions. However, their exact purpose remains unknown. We tracked Common Swifts with tracking radar at their breeding grounds in southern Sweden, and present evidence that during the dusk ascent and dawn descent they often occur in flocks, whereas during the dusk descent and dawn ascent phase they do not. This flocking behaviour suggests that swifts may benefit from conspecific interactions during twilight ascents and descents, possibly through more robust cue acquisition and information exchange in groups, or extending social behaviour also seen in screaming parties before dusk.     

Wind conditions facilitate the seasonal water‐crossing behaviour of Oriental Honey‐buzzards Pernis ptilorhynchus over the East China Sea

Migratory raptors rarely fly over stretches of water larger than 25 km, although different species undertake water crossings of varying lengths, depending mainly on their wing morphology. Oriental Honey‐buzzards fly c. 680 km over the East China Sea in autumn from breeding areas in Japan to wintering areas in Southeast Asia, but avoid this long water crossing in spring. We investigated the effects of weather on this exceptional migratory behaviour and its seasonality through a maximum entropy niche modelling approach. We used data collected through satellite tracking of 31 adult birds as presence points and a set of variables related to wind, precipitation and convective condition as environmental predictors. Results of modelling showed very different, almost non‐overlapping, areas suitable for migration over the East China Sea region in autumn and spring. Suitable migration routes in autumn mostly occurred over the sea, whereas suitable areas for spring migration mostly occurred over land, suggesting that circumnavigating the East China Sea is preferable in spring. At the regional scale, wind conditions facilitate water‐crossing behaviour of Oriental Honey‐buzzards in autumn, but not in spring. Specifically, suitable tailwinds over the sea enable water‐crossing in autumn, whereas in spring, wind support and convective conditions are best over land. Our modelling did not suggest any importance of convective conditions for autumn migration. However, we expect that at smaller temporal scales, convective conditions would be a considerable facilitator of the water‐crossing behaviour in this species.     

Wing‐beat characteristics of birds recorded with tracking radar and cine camera

This study presents wing‐beat frequency data measured mainly by radar, complemented by video and cinematic recordings, for 153 western Palaearctic and two African species. Data on a further 45 Palaearctic species from other sources are provided in an electronic appendix. For 41 species with passerine‐type flight, the duration of flapping and pausing phases is given. The graphical presentations of frequency ranges and wing‐beat patterns show within‐species variation and allow easy comparison between species, taxonomic groups and types of flight. Wing‐beat frequency is described by Pennycuick (J. Exp. Biol. 2001; 204: 3283–3294) as a function of body‐mass, wing‐span, wing‐area, gravity and air density; for birds with passerine‐type flight the power‐fraction has also to be considered. We tested Pennycuick’s general allometric model and estimated the coefficients based on our data. The general model explained a high proportion of variation in wing‐beat frequency and the coefficients differed only slightly from Pennycuick’s original values. Modelling continuous‐flapping flyers alone resulted in coefficients not different from those predicted (within 95% intervals). Doing so for passerine‐type birds resulted in a model with non‐significant contributions of body‐mass and wing‐span to the model. This was mainly due to the very high correlation between body‐mass, wing‐span and wing‐area, revealing similar relative scaling properties within this flight type. However, wing‐beat frequency increased less than expected with respect to power‐fraction, indicating that the drop in flight level during the non‐flapping phases, compensated by the factor (g/q)^0.5 in Pennycuick’s model, is smaller than presumed. This may be due to lift produced by the body during the bounding phase or by only partial folding of the wings.