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.     

Effects of weather on avian migration at proposed ridgeline wind energy sites

With the popularity of wind energy increasing globally, concerns surfaced in the 1980s as to the potential adverse effects of wind turbines on migrating birds. Understanding how weather conditions influence passage rates can help determine the potential for increased avian–turbine collisions. Using vertical and horizontal mounted marine radars, raptor stand watch observations, and portable handheld weather stations, we studied how temperature, cloud cover, barometric pressure, wind direction, and wind speed affected avian passage rates and height of migrants over 3 ridges (Wartenbe, North Dokie, and South Dokie) being developed for wind energy in northern British Columbia. Using an Akaike's Information Criterion (AIC), we determined that a reduced model combining wind speed, barometric pressure, and cloud cover was best at explaining and predicting higher passage rates (expressed as no. birds/hr) in the fall migration for both diurnal and nocturnal migrants. Wind speed proved the most important predictor of passage rates for spring nocturnal migrants and a combination of cloud cover, temperature, and wind direction for diurnal spring migrants. Wind speed also predicted decreases in flight altitude among nocturnal migrants but increased altitude in diurnal migrants. This information coupled with migration timing and topographical areas of higher migrant activity can be useful to wind energy proponents who wish to mitigate collision risk with migrating birds. © 2011 The Wildlife Society.     

Altitude choice by night migrants in a desert area predicted by meteorological factors

The height distribution of nocturnal migrants in southern Israel was determined by con-ically scanning the sky with the pencil-beam of an X-band radar at different elevation angles. Altitudinal profiles of meteorological parameters were derived from radio sondes launched at midnight and from pilot balloons launched every 4 h. A model to predict the height distribution of birds by means of meteorological variables was developed by assuming that the observed proportions of birds within a height zone, compared with the neighbouring height zones, reflect the degree of the birds' preference for that height zone. Only one among the variables included in the multiple regression analysis proved to have a significant influence on the height distribution of migrants: the difference of tailwind speed between height zones. Simulations with 1000 birds choosing altitudes by means of the night's altitudinal profile of tailwind speed closely traced the observed distributions. The fact that all the other meteorological factors which were previously suggested to have an influence on the flight range in trans-desert migration were not selected as relevant factors is discussed. The following basic information on nocturnal bird migration in the Negev is provided as a background for the statistical analysis: Directions of migration are within very narrow limits. During the first hour after take-off, 60% of autumn migrants and 75% of spring migrants are climbing, with vertical speeds of 0.1–2 m per s and 0.1–4 m per s, respectively. During the rest of the night, climbing and descending birds are in nearly equal proportions. Thus, there is a high potential of sampling atmospheric conditions at different altitudes. Height distributions in spring and autumn show the influence of the trade wind situation, autumn migrants making use of the northerly winds at low levels in spite of high temperatures, while spring migrants tend to reach the southwesterly winds at higher levels.     

Projected changes in wind assistance under climate change for nocturnally migrating bird populations

Current climate models and observations indicate that atmospheric circulation is being affected by global climate change. To assess how these changes may affect nocturnally migrating bird populations, we need to determine how current patterns of wind assistance at migration altitudes will be enhanced or reduced under future atmospheric conditions. Here, we use information compiled from 143 weather surveillance radars stations within the contiguous United States to estimate the daily altitude, density, and direction of nocturnal migration during the spring and autumn. We intersected this information with wind projections to estimate how wind assistance is expected to change during this century at current migration altitudes. The prevailing westerlies at midlatitudes are projected to increase in strength during spring migration and decrease in strength to a lesser degree during autumn migration. Southerly winds will increase in strength across the continent during both spring and autumn migration, with the strongest gains occurring in the center of the continent. Wind assistance is projected to increase across the central (0.44 m/s; 10.1%) and eastern portions of the continent (0.32 m/s; 9.6%) during spring migration, and wind assistance is projected to decrease within the central (0.32 m/s; 19.3%) and eastern portions of the continent (0.17 m/s; 6.6%) during autumn migration. Thus, across a broad portion of the continent where migration intensity is greatest, the efficiency of nocturnal migration is projected to increase in the spring and decrease in the autumn, potentially affecting time and energy expenditures for many migratory bird species. These findings highlight the importance of placing climate change projections within a relevant ecological context informed through empirical observations, and the need to consider the possibility that climate change may generate both positive and negative implications for natural systems.     

Nocturnal passerine migration without tailwind assistance

Nocturnal passerine migrants could substantially reduce the amount of energy spent per distance covered if they fly with tailwind assistance and thus achieve ground speeds that exceed their airspeeds (the birds’ speed in relation to the surrounding air). We analysed tracking radar data from two study sites in southern and northern Scandinavia and show that nocturnally migrating passerines, during both spring and autumn migration, regularly travelled without tailwind assistance. Average ground and airspeeds of the birds were strikingly similar for all seasonal and site‐specific samples, demonstrating that winds had little overall influence on the birds’ resulting travel speeds. Distributions of wind effects, measured as (1) the difference between ground and airspeed and (2) the tail/headwind component along the birds’ direction of travel, showed peaks close to a zero wind effect, indicating that the migratory flights often occurred irrespective of wind direction. An assessment of prevailing wind speeds at the birds’ mean altitude indicated a preference for lower wind speeds, with flights often taking place in moderate winds of 3–10 m/s. The limited frequency of wind‐assisted flights among the nocturnal passerine migrants studied is surprising and in clear contrast to the strong selectivity of tailwinds exhibited by some other bird groups. Relatively high costs of waiting for favourable winds, rather low probabilities of occurrence of tailwind conditions and a need to use a large proportion of nights for flying are probably among the factors that explain the lack of a distinct preference for wind‐assisted flights among nocturnal passerine migrants.     

Winds at departure shape seasonal patterns of nocturnal bird migration over the North Sea

On their migratory journeys, terrestrial birds can come across large inhospitable areas with limited opportunities to rest and refuel. Flight over these areas poses a risk especially when wind conditions en route are adverse, in which case inhospitable areas can act as an ecological barrier for terrestrial migrants. Thus, within the east-Atlantic flyway, the North Sea can function as an ecological barrier. The main aim of this study was to shed light on seasonal patterns of bird migration in the southern North Sea and determine whether departure decisions on nights of intense migration were related to increased wind assistance. We measured migration characteristics with a radar that was located 18 km off the NW Dutch coast and used simulation models to infer potential departure locations of birds on nights with intense nocturnal bird migration. We calculated headings, track directions, airspeeds, groundspeeds on weak and intense migration nights in both seasons and compared speeds between seasons. Moreover, we tested if departure decisions on intense migration nights were associated with supportive winds. Our results reveal that on the intense migration nights in spring, the mean heading was towards E, and birds departed predominantly from the UK. On intense migration nights in autumn, the majority of birds departed from Denmark, Germany and north of the Netherlands with the mean heading towards SW. Prevailing winds from WSW at departure were supportive of a direct crossing of the North Sea in spring. However, in autumn winds were generally not supportive, which is why many birds exploited positive wind assistance which occurred on intense migration nights. This implies that the seasonal wind regimes over the North Sea alter its migratory dynamics which is reflected in headings, timing and intensity of migration.     

Breeding latitude predicts timing but not rate of spring migration in a widespread migratory bird in South America

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Flight altitude of trans-Sahara migrants in autumn: a comparison of radar observations with predictions from meteorological conditions and water and energy balance models

Radar observations on the altitude of bird migration and altitudinal profiles of meteorological conditions over the Sahara desert are presented for the autumn migratory period. Migratory birds fly at an average altitude of 1016 m (a.s.l.) during the day and 571 m during the night. Weather data served to calculate flight range using two models: an energy model (EM) and an energy-and-water model (EWM). The EM assumes that fuel supply limits flight range whereas the EWM assumes that both fuel and water may limit flight range. Flight ranges estimated with the EM were generally longer than those with the EWM. This indicates that trans-Sahara migrants might have more problems balancing their water than their energy budget. However, if we assume fuel stores to consist of 70% instead of 100% fat (the remainder consisting of 9% protein and 21% water), predicted flight ranges of the EM and EWM largely overlap. Increased oxygen extraction, reduced flight costs, reduced exhaled air temperature, reduced cutaneous water loss and increased tolerance to water loss are potential physiological adaptations that would improve the water budget in migrants. Both the EM and EWM predict optimal flight altitudes in agreement with radar observations in autumn. Optimal flight altitudes are differently predicted by the EM and EWM for nocturnal spring migration. During spring, the EWM predicts moderately higher and the EM substantially higher flight altitudes than during autumn. EWM predictions are therefore in better agreement with radar observations on flight altitude of migrants over the Negev desert in spring than EM predictions.     

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.     

Optimal use of wind by migrating birds: combined drift and overcompensation

Migrating birds may save flying time by allowing themselves to be partially drifted by strong winds at high altitude and correcting for the displacement at low altitude under relatively weaker winds. This behaviour will be favourable with strong upper winds and with wind direction approx. 30 ° to 90 ° in relation to the goal direction (following side winds). Radar observations of drift in high altitude bird migration and visual records of low altitude overcompensation are compatible with the optimal flight behaviour of migrants at high and low altitude, respectively, as predicted from this hypothesis.     

Structure of communities of ground-living spiders along altitudinal gradients

Differences in the altitudinal composition of ground-living spider communities were studied by pitfall trapping in western Norway. These data plus compilations of information from the literature were used to investigate features in the size composition, temporal appearance and spatial distribution of species. Spiders at the highest altitude had the widest distribution both in terms of geographical and altitudinal range. At the lowest altitude species were more often stenochronous than at higher altitudes. The size composition, reflecting food choice of the spiders, tended to be bimodal with increasing altitude. It is argued that small spiders are segregated mainly along the habitat dimension (searchers), while the larger ones are segregated with respect to diel activity patterns and food choice (pursuers). We suggest that mountain spider faunas in temperate regions at high latitudes mainly consist of widely distributed and easily dispersed species.     

白马雪山黑白仰鼻猴(Rhinopithecus bieti)垂直迁移

2000-2001年间,在白马雪山国家级自然保护区南任村附近,通过在四个山坡每隔海拔100米的样带(宽5.0米)内搜集黑白仰鼻猴的粪便来研究猴群不同季节的垂直迁移模式。同时,通过在每个地区山脊上和山沟内搜集2.5米宽垂直样带内的粪便,来研究猴群不同季节对山脊和山沟的选择性。结果表明:猴群在海拔3500-4300米的区域活动。虽然猴群全年集中利用3900-4200米的林带;但是具有季节性差异:夏季最高(4200米),依次是秋季(4100米)和冬季(4000米),夏季最低(3900米)。另外,冬季山沟中的粪便多于山脊说明猴群在沟中停留时间长,这与沟中少风且温度高有关。不同海拔带上的粪便密度和松萝量间正相关,这意味着食物资源的垂直分布是影响猴群垂直迁移的重要因素。春季,猴群会下到低海拔采食嫩芽/叶,而冬季则下到低海拔处躲避首次大雪,这很可能导致猴群集中利用第二个海拔带。     

BIRD MIGRATION OVER THE MALTESE ISLANDS

The present paper is based on data obtained during several years' observations and three recent surveys. Little has previously been published on migration through the Maltese Islands. The geographical, vegetational and climatic factors of the islands are discussed in so far as they affect the migrants. Visible migration is seen with anticyclonic weather and westerly winds. Birds are found grounded after night migration in cyclonic weather with southwesterly or easterly winds, much larger numbers and variety being seen with the latter. When there is a deterioration in the weather during the night, a large influx of birds is seen on the following morning, and in addition large flocks of migrating Turtle Doves are seen. Several trans-Saharan migrants may pass in smaller numbers during autumn than spring, but the difference may be more apparent than real because in early autumn the birds may depart after only a very short stay, and a few conspicuous species are absent or scarce. By contrast several species which winter north of the Sahara pass only or in much larger numbers during autumn, and these more than make up for those which are absent or rare. There is no evidence from bird ringing that in spring Malta regularly gets birds from Tunisia, at any rate from that part covered by the ringing stations (Cap Bon, Enfidaville, Gabes). The migrants which pass through Malta probably originate from an area in North Africa around Tripoli and some way westwards of it. During autumn the bulk of recoveries is from eastern European countries with a smaller percentage from northern and central Europe. Several species or groups of species are dealt with individually. In the discussion stress is laid on the very close relationship between migration and weather, especially the wind component. The comparatively small numbers of birds seen at Malta probably form part of a larger movement travelling on a broad front. It is argued that the large “falls” of migrants in bad weather result from drift acting on a mass passing mainly to one side or the other of the islands. Since much larger densities are seen with easterly than with westerly winds, it follows, if the hypothesis of drift be correct, that the numbers of birds travelling to the east of Malta are larger than those to the west of it. Moreover, since day migrants are seen with westerly winds and the bulk of night migrants with easterly ones, it is inferred that day migrants normally pass to the west, and the bulk of night migrants to the east, of the Maltese Islands. A parallel is drawn between the autumn migration and the performance of racing pigeons which are flown from the north and NE at this season.     

Assessing the origin of Neotropical mountain dung beetle assemblages (Scarabaeidae: Scarabaeinae): the comparative influence of vertical and horizontal colonization

Aims The fauna of mountains and their surrounding regions are likely to be influenced principally by two biological processes: horizontal colonization along similar altitudinal levels by elements originating from lineages inhabiting higher latitudes; and vertical colonization by lineages from the same latitude, but at lower altitudes. We examine whether the expected patterns derived from the latter process can be observed in mountain dung beetle assemblages. Specifically, we study the variation in species composition and richness with altitude in five regions spanning elevation gradients, analysing whether the altitudinal rates of change in the number of species and genera differ, and whether beta‐diversity scores for adjacent sites in each altitudinal gradient are different for species and genera. Location Eastern Cordillera of the Colombian Andes. Methods Field work was carried out in 1997–99 at 27 sites in five regions with elevation gradients, with 10–32 pitfall traps placed in each site. For each altitudinal level the numbers of species and genera were analysed with respect to altitude, and the slope of the linear regression between these variables was calculated. The slope of the curve of the altitude against the cumulative number of species and genera was also calculated for each altitudinal gradient to describe the compositional change between adjacent sites (beta diversity). Species and generic slopes were compared using analysis of covariance. The turnover of species along each altitudinal gradient was measured using presence/absence data and Cody's beta‐diversity index between adjacent pairs of sites. A cluster analysis was used to detect faunistically homogeneous groups of localities. Results Species richness always decreased with altitude, although the slopes did not differ significantly from zero. The number of genera also decreased with increasing altitude, but generally at a significantly slower rate than for species. Variation in the species beta‐diversity scores between altitudinal levels did not follow a homogeneous pattern in the different regions. Two main altitudinal groups of sites with a boundary c. 1500–1750 m a.s.l. can be detected with respect to faunistic similarity. Low‐ and mid‐altitude sites are inhabited by all of the genera (19) and 80% of all species collected. Eight genera and 61 species (c. 60% of the total) are unable to inhabit high‐altitude sites, and only 20 species appear to be exclusive to these high‐altitude environments (> 2000 m a.s.l.). Main conclusions The dominant processes explaining dung beetle composition in the high north‐eastern Andean mountains are probably those of vertical colonization. The limited role of horizontal colonization processes, or colonization from northern or southern lineages, could be a consequence of the isolation and recent geological origin of these mountains.     

Altitudinal migration by birds: a review of the literature and a comprehensive list of species

Altitudinal migration is the seasonal altitudinal movement of birds from breeding areas to non‐breeding or wintering areas at different elevations. Although this type of migration is widely reported, questions remain concerning the number of species that perform altitudinal migration, possible variation among different taxa and geographic locations in the extent of altitudinal migration, and the foraging guilds of altitudinal migrants. We conducted an extensive bibliographic survey and compiled a list of altitudinal migrant birds worldwide. We characterized species in terms of their foraging guilds because the spatial distribution of food resources along altitudinal gradients is often evoked as a driver of bird altitudinal migration. We identified 1238 species of altitudinal migrants, ~10% of the ~10,000 extant species of birds. We found a strong geographic bias in publications focusing on avian altitudinal migration toward the United States and Costa Rica, and a paucity of studies in megadiverse regions such as the Afrotropical and Indomalayan realms, and areas in the Neotropics other than Costa Rica. We also found that most species of altitudinal migrants were invertivores rather than frugivores or nectarivores. This general pattern held true for all zoogeographic realms except the Neotropics, where nectarivores and frugivores predominated among altitudinal migrants. The prevalence of invertivore birds among altitudinal migrants is not unexpected because this is the most common foraging guild among birds worldwide. Overall, we found no prevalence of any specific foraging guild among altitudinal migrants across zoogeographic regions. The results of studies to date suggest that altitudinal migration by birds may be driven by a number of factors, including access to increased food resources for breeding or molting, weather conditions, and mating and nesting opportunities. However, to better understand the mechanisms underlying altitudinal migration, broadening the geographic scope of studies is paramount, with additional study of altitudinal migration especially needed in the megadiverse tropical regions of sub‐Saharan Africa, Southeast Asia, and South America.     

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.     

Process in the evolution of bird migration and pattern in avian ecogeography

Current ideas about the evolution of bird migration equate its origin with the first appearance of fully migratory populations, and attribute its evolution to a selective advantage generated by increased breeding success, gained through temporary emigration from resident populations to breed in under-exploited seasonal areas. I propose an alternative hypothesis in which migration first appears as a temporary directional shift away from the breeding site outside the reproductive period, in response to seasonal variation in the direction and/or severity of environmental gradients. Fully migratory populations then appear through either extinction of sedentary phenotypes, or colonisation of vacant seasonal areas by migrants. Where colonisation occurs, resident ancestral populations can be driven to extinction by competition from migrants which invade their range outside the breeding season, resulting in fully migratory species. An analogous process drives the evolution of migration between high latitudes and the tropics, since extension of breeding range into higher latitudes may drive low latitude populations to extinction, resulting in an overall shift of breeding range. This process can explain reverse latitudinal gradients in avian diversity in the temperate zone, since the breeding ranges of migratory species concentrate in latitudes where they enjoy the highest breeding success. Near absence of forest-dwelling species among Palaearctic-African migrants is attributable to the lack of forest in northern Africa for much of the Tertiary, which has precluded selection both for southward extension of migration by west Palaearctic forest species, and northward breeding colonisation by African forest species.     

Latitudinal distribution, migration, and testosterone levels in birds

Tropical bird species usually have lower testosterone (T) levels during breeding than temperate species. However, the potential mechanisms behind the positive interspecific correlation between T and latitude remain unexplored. In a comparative study of more than 100 bird species, we examined whether social constraints during male-male competition arising from migration and breeding synchrony are responsible for the latitude effects. Species that breed at higher latitudes are more likely to migrate and experience more intense intrasexual competition upon spring arrival than nonmigrant species from lower latitudes. Additionally, species from higher latitudes cope with shorter breeding seasons and thus with more synchronous breeding, which selects for high T titers via increased male-male conflicts. Accordingly, peak T levels were associated with migration and the duration of the egg laying period that reflects breeding synchrony. Because migration and breeding synchrony were related to latitudinal distribution, they appear to be important components of the latitude effects on T. A multivariate model controlling for covariation of predictor variables revealed that latitude remained the strongest predictor of peak T. Therefore, selection due to migration and breeding synchrony may partially cause the latitude effect, but other geographically varying factors may also play a role in mediating peak T levels at different latitudes.     

Detecting Altitudinal Migration Events in Neotropical Birds Using Stable Isotopes

We evaluated the utility of stable‐hydrogen isotope ratios in tropical bird tissues for detecting altitudinal migration events. Our results identified two of five species as altitudinal migrants in Nicaragua. This approach may circumvent the current limitations of mark–recapture techniques and enhance our ability to study this poorly characterized behavior.     

Bird movements at rotor heights measured continuously with vertical radar at a Dutch offshore wind farm

Assessing the impacts of avian collisions with wind turbines requires reliable estimates of avian flight intensities and altitudes, to enable accurate estimation of collision rates, avoidance rates and related effects on populations. At sea, obtaining such estimates visually is limited not only by weather conditions but, more importantly, because a high proportion of birds fly at night and at heights above the range of visual observation. We used vertical radar with automated bird‐tracking software to overcome these limitations and obtain data on the magnitude, timing and altitude of local bird movements and seasonal migration measured continuously at a Dutch offshore wind farm. An estimated 1.6 million radar echoes representing individual birds or flocks were recorded crossing the wind farm annually at altitudes between 25 and 115 m (the rotor‐swept zone). The majority of these fluxes consisted of gull species during the day and migrating passerines at night. We demonstrate daily, monthly and seasonal patterns in fluxes at rotor heights and the influence of wind direction on flight intensity. These data are among the first to show the magnitude and variation of low‐altitude flight activity across the North Sea, and are valuable for assessing the consequences of developments such as offshore wind farms for birds.