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1.  Bird orientation: compensation for wind drift in migrating raptors is age dependent. 
Despite the potentially strong effect of wind on bird orientation, our understanding of how wind drift affects migrating birds is still very limited. Using data from satellite-based radio telemetry, we analysed the effect of changing winds on the variation of the track direction of individual birds. We studied adults and juveniles of two raptor species, osprey Pandion haliaetus and honey buzzard Pernis apivorus, on autumn migration between North Europe and Africa, and demonstrate an important difference between the age categories of both species in the extent of wind drift. For juveniles, side- and following-wind components affected the rates of movement perpendicular to and along the mean direction, respectively, to a similar degree, suggesting full wind drift. By contrast, for adults the rate of crosswind displacement was significantly smaller than the effect of wind on forward movement, showing much reduced wind drift (29%). This indicates that adults have acquired a more sophisticated orientation system, permitting detection of and compensation for wind drift, than juveniles. These drift effects are likely to reduce the ability of juveniles to locate species-specific wintering areas in case of rapid climatic wind change.
PMCID: PMC1698035  PMID: 12952622
2.  Juvenile Songbirds Compensate for Displacement to Oceanic Islands during Autumn Migration 
PLoS ONE  2011;6(3):e17903.
To what degree juvenile migrant birds are able to correct for orientation errors or wind drift is still largely unknown. We studied the orientation of passerines on the Faroe Islands far off the normal migration routes of European migrants. The ability to compensate for displacement was tested in naturally occurring vagrants presumably displaced by wind and in birds experimentally displaced 1100 km from Denmark to the Faroes. The orientation was studied in orientation cages as well as in the free-flying birds after release by tracking departures using small radio transmitters. Both the naturally displaced and the experimentally displaced birds oriented in more easterly directions on the Faroes than was observed in Denmark prior to displacement. This pattern was even more pronounced in departure directions, perhaps because of wind influence. The clear directional compensation found even in experimentally displaced birds indicates that first-year birds can also possess the ability to correct for displacement in some circumstances, possibly involving either some primitive form of true navigation, or ‘sign posts’, but the cues used for this are highly speculative. We also found some indications of differences between species in the reaction to displacement. Such differences might be involved in the diversity of results reported in displacement studies so far.
PMCID: PMC3064565  PMID: 21464975
3.  Flying with the wind: scale dependency of speed and direction measurements in modelling wind support in avian flight 
Movement Ecology  2013;1(1):4.
Understanding how environmental conditions, especially wind, influence birds' flight speeds is a prerequisite for understanding many important aspects of bird flight, including optimal migration strategies, navigation, and compensation for wind drift. Recent developments in tracking technology and the increased availability of data on large-scale weather patterns have made it possible to use path annotation to link the location of animals to environmental conditions such as wind speed and direction. However, there are various measures available for describing not only wind conditions but also the bird's flight direction and ground speed, and it is unclear which is best for determining the amount of wind support (the length of the wind vector in a bird’s flight direction) and the influence of cross-winds (the length of the wind vector perpendicular to a bird’s direction) throughout a bird's journey.
We compared relationships between cross-wind, wind support and bird movements, using path annotation derived from two different global weather reanalysis datasets and three different measures of direction and speed calculation for 288 individuals of nine bird species. Wind was a strong predictor of bird ground speed, explaining 10-66% of the variance, depending on species. Models using data from different weather sources gave qualitatively similar results; however, determining flight direction and speed from successive locations, even at short (15 min intervals), was inferior to using instantaneous GPS-based measures of speed and direction. Use of successive location data significantly underestimated the birds' ground and airspeed, and also resulted in mistaken associations between cross-winds, wind support, and their interactive effects, in relation to the birds' onward flight.
Wind has strong effects on bird flight, and combining GPS technology with path annotation of weather variables allows us to quantify these effects for understanding flight behaviour. The potentially strong influence of scaling effects must be considered and implemented in developing sampling regimes and data analysis.
Electronic supplementary material
The online version of this article (doi:10.1186/2051-3933-1-4) contains supplementary material, which is available to authorized users.
PMCID: PMC4337751  PMID: 25709818
NOAA; ECMWF; GPS; Aves; Doppler-shift; Scaling; Measurement error; Flight direction; Flight speed
4.  Convergent patterns of long-distance nocturnal migration in noctuid moths and passerine birds 
Vast numbers of insects and passerines achieve long-distance migrations between summer and winter locations by undertaking high-altitude nocturnal flights. Insects such as noctuid moths fly relatively slowly in relation to the surrounding air, with airspeeds approximately one-third of that of passerines. Thus, it has been widely assumed that windborne insect migrants will have comparatively little control over their migration speed and direction compared with migrant birds. We used radar to carry out the first comparative analyses of the flight behaviour and migratory strategies of insects and birds under nearly equivalent natural conditions. Contrary to expectations, noctuid moths attained almost identical ground speeds and travel directions compared with passerines, despite their very different flight powers and sensory capacities. Moths achieved fast travel speeds in seasonally appropriate migration directions by exploiting favourably directed winds and selecting flight altitudes that coincided with the fastest air streams. By contrast, passerines were less selective of wind conditions, relying on self-powered flight in their seasonally preferred direction, often with little or no tailwind assistance. Our results demonstrate that noctuid moths and passerines show contrasting risk-prone and risk-averse migratory strategies in relation to wind. Comparative studies of the flight behaviours of distantly related taxa are critically important for understanding the evolution of animal migration strategies.
PMCID: PMC3158935  PMID: 21389024
seasonal migration; songbirds; Lepidoptera; Autographa gamma; flight speed; orientation
5.  Confronting the winds: orientation and flight behaviour of roosting swifts, Apus apus. 
Swifts, Apus apus, spend the night aloft and this offers an opportunity to test the degree of adaptability of bird orientation and flight to different ecological situations. We predicted the swifts' behaviour by assuming that they are adapted to minimize energy expenditure during the nocturnal flight and during a compensatory homing flight if they become displaced by wind. We tested the predictions by recording the swifts' altitudes, speeds and directions under different wind conditions with tracking radar; we found an agreement between predictions and observations for orientation behaviour, but not for altitude and speed regulation. The swifts orientated consistently into the head wind, with angular concentration increasing with increasing wind speed. However, contrary to our predictions, they did not select altitudes with slow or moderate winds, nor did they increase their airspeed distinctly when flying into strong head winds. A possible explanation is that their head-wind orientation is sufficient to keep nocturnal displacement from their home area within tolerable limits, leaving flight altitude to be determined by other factors (correlated with temperature), and airspeed to show only a marginal increase in strong winds. The swifts were often moving "backwards", heading straight into the wind but being overpowered by wind speeds exceeding their airspeed. The regular occurrence of such flights is probably uniquely associated with the swifts' remarkable habit of roosting on the wing.
PMCID: PMC1088711  PMID: 11375093
6.  Commuting fruit bats beneficially modulate their flight in relation to wind 
When animals move, their tracks may be strongly influenced by the motion of air or water, and this may affect the speed, energetics and prospects of the journey. Flying organisms, such as bats, may thus benefit from modifying their flight in response to the wind vector. Yet, practical difficulties have so far limited the understanding of this response for free-ranging bats. We tracked nine straw-coloured fruit bats (Eidolon helvum) that flew 42.5 ± 17.5 km (mean ± s.d.) to and from their roost near Accra, Ghana. Following detailed atmospheric simulations, we found that bats compensated for wind drift, as predicted under constant winds, and decreased their airspeed in response to tailwind assistance such that their groundspeed remained nearly constant. In addition, bats increased their airspeed with increasing crosswind speed. Overall, bats modulated their airspeed in relation to wind speed at different wind directions in a manner predicted by a two-dimensional optimal movement model. We conclude that sophisticated behavioural mechanisms to minimize the cost of transport under various wind conditions have evolved in bats. The bats’ response to the wind is similar to that reported for migratory birds and insects, suggesting convergent evolution of flight behaviours in volant organisms.
PMCID: PMC3973273  PMID: 24648227
atmospheric modelling; biotelemetry; crosswind compensation; Eidolon helvum; flight behaviour; tailwind assistance
7.  High Altitude Bird Migration at Temperate Latitudes: A Synoptic Perspective on Wind Assistance 
PLoS ONE  2013;8(1):e52300.
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.
PMCID: PMC3536796  PMID: 23300969
8.  Geographical and temporal flexibility in the response to crosswinds by migrating raptors 
Wind and ocean currents may potentially have important effects on travelling animals, as an animal which does not respond to lateral flow will be drifted from its intended direction of movement. By analysing daily movements of migrating ospreys Pandion haliaetus and marsh harriers Circus aeruginosus, as recorded by satellite telemetry, in relation to global wind data, we showed that these raptors allow on average 47 per cent drift. Furthermore, our analyses revealed significant geographical and temporal variation in the response to crosswinds. During some parts of the migration, the birds drifted and in other parts they compensated or even overcompensated. In some regions, the response of marsh harriers depended on the wind direction. They drifted when the wind came from one side and (over)compensated when the wind came from the opposite side, and this flexible response was different in different geographical regions. These results suggest that migrating raptors modulate their response to crosswinds at different places and times during their travels and show that individual birds use a much more varied repertoire of behavioural responses to wind than hitherto assumed. Our results may also explain why contrasting and variable results have been obtained in previous studies of the effect of wind on bird migration.
PMCID: PMC3061148  PMID: 20980299
bird orientation; migration; satellite tracking; wind drift
9.  Evidence for Sex-Segregated Ocean Distributions of First-Winter Wandering Albatrosses at Crozet Islands 
PLoS ONE  2014;9(2):e86779.
The highly mobile wandering albatrosses (Diomedea exulans) are adapted to navigate the extreme environment of the Southern Ocean and return to isolated islands to breed. Each year they cover several hundreds of thousands of kilometers during travels across the sea. Little is known about the dispersal flights and migration of young albatrosses. We tracked, by satellite telemetry, the departure dispersal of 13 juvenile wandering albatrosses from the Crozet Islands and compared them with tracks of 7 unrelated adults during the interbreeding season. We used the satellite tracks to identify different behavioural steps of the inherited migration program used by juvenile wandering albatrosses during their first solo-migration. Our results show that the juvenile wandering albatrosses from Crozet Islands moved to sex-specific foraging zones of the ocean using at departures selectively the wind. The results suggest that the inherited migration program used by the juvenile wandering albatrosses encode several distinct steps, based on inherited preferred departure routes, differences in migration distance between sexes, and selective use of winds. During long transportation flights the albatrosses were influenced by winds and both adult and juveniles followed approximate loxodrome (rhumbline) routes coinciding with the foraging zone and the specific latitudes of their destination areas. During the long segments of transportation flights across open seas the juveniles selected routes at more northerly latitudes than adults.
PMCID: PMC3937278  PMID: 24586254
10.  Weather, host and vector — their interplay in the spread of insect-borne animal virus diseases 
The Journal of Hygiene  1980;85(1):65-102.
The spread of insect-borne animal virus diseases is influenced by a number of factors. Hosts migrate, move or are conveyed over long distances: vectors are carried on the wind for varying distances in search of hosts and breeding sites; weather and climate affect hosts and vectors through temperature, moisture and wind. As parasites of host and vector, viruses are carried by animals, birds and insects, and their spread can be correlated with the migration of hosts and the carriage of vectors on winds associated with the movements of the Intertropical Convergence Zone (ITCZ) and warm winds to the north and south of the limits of the ITCZ. The virus is often transmitted from a local cycle to a migratory cycle and back again.
Examples of insect-borne virus diseases and their spread are analysed. Japanese, Murray Valley, Western equine, Eastern equine and St Louis encephalitis represent viruses transmitted by mosquito—bird or pig cycles.
The areas experiencing infection with these viruses can be divided into a number of zones: A, B, C, D, E and F. In zone A there is a continuous cycle of virus in host and vector throughout the year; in zone B, there is an upsurge in the cycle during the wet season, but the cycle continues during the dry season; there is movement of infected vectors between and within zones A and B on the ITCZ and the virus is introduced to zone C by infected vectors on warm winds; persistence may occur in zone C if conditions are right. In zone D, virus is introduced each year by infected vectors on warm winds and the arrival of the virus coincides with the presence of susceptible nestling birds and susceptible piglets. The disappearance of virus occurs at the time when migrating mosquitoes and birds are returning to warmer climates. The virus is introduced to zone E only on occasions every 5-10 years when conditions are suitable. Infected hosts introduced to zone F do not lead to circulation of virus, since the climate is unsuitable for vectors. Zones A, B and C correspond to endemic and zones D and E to epidemic conditions.
Similar zones can be recognized for African horse sickness, bluetongue, Ibaraki disease and bovine ephemeral fever — examples of diseases transmitted in a midge-mammal cycle. In zones A and B viruses are transported by infected midges carried on the wind in association with the movement of ITCZ and undergo cycles in young animals. In these zones and in zone C there is a continual movement of midges on the warm wind between one area and another, colonizing new sites or reinforcing populations of midges already present. Virus is introduced at times into fringe areas (zones D and E) and, as there is little resistance in the host, gives rise to clinical signs of disease. In some areas there is persistence during adverse conditions; in others, the virus is carried back to the endemic zones by infected midges or vectors.
Examples of viruses maintained in a mosquito/biting fly—mammal cycle are Venezuelan equine encephalitis and vesicular stomatitis. These viruses enter a migratory cycle from a local cycle and the vectors in the migratory cycle are carried over long distances on the wind. Further examples of virus spread by movement of vectors include West Nile, Rift Valley fever, yellow fever, epizootic haemorrhagic disease of deer and Akabane viruses.
In devising means of control it is essential to decide the relationship of host, vector and virus and the nature of the zone in which the area to be controlled lies. Because of the continual risk of reintroduction of infected vectors, it is preferable to protect the host by dipping, spraying or by vaccination rather than attempting to eliminate the local population of insects.
PMCID: PMC2134001  PMID: 6131919
11.  Juvenile Osprey Navigation during Trans-Oceanic Migration 
PLoS ONE  2014;9(12):e114557.
To compensate for drift, an animal migrating through air or sea must be able to navigate. Although some species of bird, fish, insect, mammal, and reptile are capable of drift compensation, our understanding of the spatial reference frame, and associated coordinate space, in which these navigational behaviors occur remains limited. Using high resolution satellite-monitored GPS track data, we show that juvenile ospreys (Pandion haliaetus) are capable of non-stop constant course movements over open ocean spanning distances in excess of 1500 km despite the perturbing effects of winds and the lack of obvious landmarks. These results are best explained by extreme navigational precision in an exogenous spatio-temporal reference frame, such as positional orientation relative to Earth's magnetic field and pacing relative to an exogenous mechanism of keeping time. Given the age (<1 year-old) of these birds and knowledge of their hatching site locations, we were able to transform Enhanced Magnetic Model coordinate locations such that the origin of the magnetic coordinate space corresponded with each bird's nest. Our analyses show that trans-oceanic juvenile osprey movements are consistent with bicoordinate positional orientation in transformed magnetic coordinate or geographic space. Through integration of movement and meteorological data, we propose a new theoretical framework, chord and clock navigation, capable of explaining the precise spatial orientation and temporal pacing performed by juvenile ospreys during their long-distance migrations over open ocean.
PMCID: PMC4262435  PMID: 25493430
12.  Windscapes shape seabird instantaneous energy costs but adult behavior buffers impact on offspring 
Movement Ecology  2014;2:17.
Windscapes affect energy costs for flying animals, but animals can adjust their behavior to accommodate wind-induced energy costs. Theory predicts that flying animals should decrease air speed to compensate for increased tailwind speed and increase air speed to compensate for increased crosswind speed. In addition, animals are expected to vary their foraging effort in time and space to maximize energy efficiency across variable windscapes.
We examined the influence of wind on seabird (thick-billed murre Uria lomvia and black-legged kittiwake Rissa tridactyla) foraging behavior. Airspeed and mechanical flight costs (dynamic body acceleration and wing beat frequency) increased with headwind speed during commuting flights. As predicted, birds adjusted their airspeed to compensate for crosswinds and to reduce the effect of a headwind, but they could not completely compensate for the latter. As we were able to account for the effect of sampling frequency and wind speed, we accurately estimated commuting flight speed with no wind as 16.6 ms?1 (murres) and 10.6 ms?1 (kittiwakes). High winds decreased delivery rates of schooling fish (murres), energy (murres) and food (kittiwakes) but did not impact daily energy expenditure or chick growth rates. During high winds, murres switched from feeding their offspring with schooling fish, which required substantial above-water searching, to amphipods, which required less above-water searching.
Adults buffered the adverse effect of high winds on chick growth rates by switching to other food sources during windy days or increasing food delivery rates when weather improved.
PMCID: PMC4445632  PMID: 26019870
13.  Using Nocturnal Flight Calls to Assess the Fall Migration of Warblers and Sparrows along a Coastal Ecological Barrier 
PLoS ONE  2014;9(3):e92218.
Atmospheric conditions fundamentally influence the timing, intensity, energetics, and geography of avian migration. While radar is typically used to infer the influence of weather on the magnitude and spatiotemporal patterns of nocturnal bird migration, monitoring the flight calls produced by many bird species during nocturnal migration represents an alternative methodology and provides information regarding the species composition of nocturnal migration. We used nocturnal flight call (NFC) recordings of at least 22 migratory songbirds (14 warbler and 8 sparrow species) during fall migration from eight sites along the mainland and island coasts of Rhode Island to evaluate five hypotheses regarding NFC detections. Patterns of warbler and sparrow NFC detections largely supported our expectations in that (1) NFC detections associated positively and strongly with wind conditions that influence the intensity of coastal bird migration and negatively with regional precipitation; (2) NFCs increased during conditions with reduced visibility (e.g., high cloud cover); (3) NFCs decreased with higher wind speeds, presumably due mostly to increased ambient noise; and (4) coastal mainland sites recorded five to nine times more NFCs, on average, than coastal nearshore or offshore island sites. However, we found little evidence that (5) nightly or intra-night patterns of NFCs reflected the well-documented latitudinal patterns of migrant abundance on an offshore island. Despite some potential complications in inferring migration intensity and species composition from NFC data, the acoustic monitoring of NFCs provides a viable and complementary methodology for exploring the spatiotemporal patterns of songbird migration as well as evaluating the atmospheric conditions that shape these patterns.
PMCID: PMC3958503  PMID: 24643060
14.  A single wind-mediated mechanism explains high-altitude ‘non-goal oriented’ headings and layering of nocturnally migrating insects 
Studies made with both entomological and meteorological radars over the last 40 years have frequently reported the occurrence of insect layers, and that the individuals forming these layers often show a considerable degree of uniformity in their headings—behaviour known as ‘common orientation’. The environmental cues used by nocturnal migrants to select and maintain common headings, while flying in low illumination levels at great heights above the ground, and the adaptive benefits of this behaviour have long remained a mystery. Here we show how a wind-mediated mechanism accounts for the common orientation patterns of ‘medium-sized’ nocturnal insects. Our theory posits a mechanism by which migrants are able to align themselves with the direction of the flow using a turbulence cue, thus adding their air speed to the wind speed and significantly increasing their migration distance. Our mechanism also predicts that insects flying in the Northern Hemisphere will typically be offset to the right of the mean wind line when the atmosphere is stably stratified, with the Ekman spiral in full effect. We report on the first evidence for such offsets, and show that they have significant implications for the accurate prediction of the flight trajectories of migrating nocturnal insects.
PMCID: PMC2842736  PMID: 19889697
orientation behaviour; windborne insect migration; nocturnal boundary layer; atmospheric dispersion models
15.  Flight Speeds among Bird Species: Allometric and Phylogenetic Effects 
PLoS Biology  2007;5(8):e197.
Flight speed is expected to increase with mass and wing loading among flying animals and aircraft for fundamental aerodynamic reasons. Assuming geometrical and dynamical similarity, cruising flight speed is predicted to vary as (body mass)1/6 and (wing loading)1/2 among bird species. To test these scaling rules and the general importance of mass and wing loading for bird flight speeds, we used tracking radar to measure flapping flight speeds of individuals or flocks of migrating birds visually identified to species as well as their altitude and winds at the altitudes where the birds were flying. Equivalent airspeeds (airspeeds corrected to sea level air density, Ue) of 138 species, ranging 0.01–10 kg in mass, were analysed in relation to biometry and phylogeny. Scaling exponents in relation to mass and wing loading were significantly smaller than predicted (about 0.12 and 0.32, respectively, with similar results for analyses based on species and independent phylogenetic contrasts). These low scaling exponents may be the result of evolutionary restrictions on bird flight-speed range, counteracting too slow flight speeds among species with low wing loading and too fast speeds among species with high wing loading. This compression of speed range is partly attained through geometric differences, with aspect ratio showing a positive relationship with body mass and wing loading, but additional factors are required to fully explain the small scaling exponent of Ue in relation to wing loading. Furthermore, mass and wing loading accounted for only a limited proportion of the variation in Ue. Phylogeny was a powerful factor, in combination with wing loading, to account for the variation in Ue. These results demonstrate that functional flight adaptations and constraints associated with different evolutionary lineages have an important influence on cruising flapping flight speed that goes beyond the general aerodynamic scaling effects of mass and wing loading.
Author Summary
Analysing the variation in flight speed among bird species is important in understanding flight. We tested if the cruising speed of different migrating bird species in flapping flight scales with body mass and wing loading according to predictions from aerodynamic theory and to what extent phylogeny provides an additional explanation for variation in speed. Flight speeds were measured by tracking radar for bird species ranging in size from 0.01 kg (small passerines) to 10 kg (swans). Equivalent airspeeds of 138 species ranged between 8 and 23 m/s and did not scale as steeply in relation to mass and wing loading as predicted. This suggests that there are evolutionary restrictions to the range of flight speeds that birds obtain, which counteract too slow and too fast speeds among bird species with low and high wing loading, respectively. In addition to the effects of body size and wing morphology on flight speed, we also show that phylogeny accounted for an important part of the remaining speed variation between species. Differences in flight apparatus and behaviour among species of different evolutionary origin, and with different ecology and flight styles, are likely to influence cruising flight performance in important ways.
Measurement of flight speeds of 138 species of bird reveals that mass and wing loading do not scale according to aerodynamic theory but vary significantly depending on phylogeny.
PMCID: PMC1914071  PMID: 17645390
16.  The influence of wind direction on the capture of the wood warbler (Phylloscopus sibilatrix), an uncommon migratory species in the western Mediterranean 
The wood warbler (Phylloscopus sibilatrix) is a migratory species in the western Mediterranean wintering in the Gulf of Guinea region, West Africa. In autumn and spring, this species, along with the populations breeding in Ireland and Britain, uses the Italian peninsula as its main axis of migration. From the data of captured birds at several ringing stations in the western Mediterranean (Balearic Islands and coastal Iberian Peninsula), we analyzed capture rates of the species during spring migration from 1993 to 2007. Based on the selection of days with a significant number of captures and those without captures, we analyzed the effect of wind direction over the western Mediterranean to determine a relationship between winds and the number of captures. From a total of 663 wood warblers captured between 1993 and 2007, a total of 31 days have been selected as significant days with a high number of captures, and 31 days have been selected as no-capture days. On days of maximum captures, winds coming from an easterly direction, i.e. Algeria and Tunisia, were dominant, indicating days with a clear eastern component. Contrary to expected results, captures were also made on days when the wind direction was predominantly from a northerly direction. Analysis of the origin of the winds in north eastern Spain (western Mediterranean) revealed that the majority of northerly winds originated from Africa and not from Europe as is usual for this region. Days or periods selected as no-capture days were characterized by winds coming from a northerly (European origin) or westerly direction.
Electronic supplementary material
The online version of this article (doi:10.1007/s00484-011-0443-4) contains supplementary material, which is available to authorized users.
PMCID: PMC3212683  PMID: 21604153
Wind direction; Synoptic situation; Mediterranean; Bird migration; Wood warbler
17.  Analysis of the Contribution of Wind Drift Factor to Oil Slick Movement under Strong Tidal Condition: Hebei Spirit Oil Spill Case 
PLoS ONE  2014;9(1):e87393.
The purpose of this study is to investigate the effects of the wind drift factor under strong tidal conditions in the western coastal area of Korea on the movement of oil slicks caused by the Hebei Spirit oil spill accident in 2007. The movement of oil slicks was computed using a simple simulation model based on the empirical formula as a function of surface current, wind speed, and the wind drift factor. For the simulation, the Environmental Fluid Dynamics Code (EFDC) model and Automatic Weather System (AWS) were used to generate tidal and wind fields respectively. Simulation results were then compared with 5 sets of spaceborne optical and synthetic aperture radar (SAR) data. From the present study, it was found that highest matching rate between the simulation results and satellite imagery was obtained with different values of the wind drift factor, and to first order, this factor was linearly proportional to the wind speed. Based on the results, a new modified empirical formula was proposed for forecasting the movement of oil slicks on the coastal area.
PMCID: PMC3907538  PMID: 24498094
18.  Stopover optimization in a long-distance migrant: the role of fuel load and nocturnal take-off time in Alaskan northern wheatears (Oenanthe oenanthe) 
Frontiers in Zoology  2013;10:26.
In long-distance migrants, a considerably higher proportion of time and energy is allocated to stopovers rather than to flights. Stopover duration and departure decisions affect consequently subsequent flight stages and overall speed of migration. In Arctic nocturnal songbird migrants the trade-off between a relatively long migration distance and short nights available for travelling may impose a significant time pressure on migrants. Therefore, we hypothesize that Alaskan northern wheatears (Oenanthe oenanthe) use a time-minimizing migration strategy to reach their African wintering area 15,000 km away.
We estimated the factors influencing the birds’ daily departure probability from an Arctic stopover before crossing the Bering Strait by using a Cormack-Jolly-Seber model. To identify in which direction and when migration was resumed departing birds were radio-tracked. Here we show that Alaskan northern wheatears did not behave as strict time minimizers, because their departure fuel load was unrelated to fuel deposition rate. All birds departed with more fuel load than necessary for the sea crossing. Departure probability increased with stopover duration, evening fuel load and decreasing temperature. Birds took-off towards southwest and hence, followed in general the constant magnetic and geographic course but not the alternative great circle route. Nocturnal departure times were concentrated immediately after sunset.
Although birds did not behave like time-minimizers in respect of the optimal migration strategies their surplus of fuel load clearly contradicted an energy saving strategy in terms of the minimization of overall energy cost of transport. The observed low variation in nocturnal take-off time in relation to local night length compared to similar studies in the temperate zone revealed that migrants have an innate ability to respond to changes in the external cue of night length. Likely, birds maximized their potential nightly flight range by taking off early in the night which in turn maximizes their overall migration speed. Hence, nocturnal departure time may be a crucial parameter shaping the speed of migration indicating the significance of its integration in future migration models.
PMCID: PMC3665591  PMID: 23663358
Arctic; Capture-recapture; Cormack-Jolly-Seber model; Departure probability; Departure time; Migration speed; Northern wheatear; Optimization; Songbird; Stopover ecology
19.  Ocean Surface Winds Drive Dynamics of Transoceanic Aerial Movements 
PLoS ONE  2008;3(8):e2928.
Global wind patterns influence dispersal and migration processes of aerial organisms, propagules and particles, which ultimately could determine the dynamics of colonizations, invasions or spread of pathogens. However, studying how wind-mediated movements actually happen has been hampered so far by the lack of high resolution global wind data as well as the impossibility to track aerial movements. Using concurrent data on winds and actual pathways of a tracked seabird, here we show that oceanic winds define spatiotemporal pathways and barriers for large-scale aerial movements. We obtained wind data from NASA SeaWinds scatterometer to calculate wind cost (impedance) models reflecting the resistance to the aerial movement near the ocean surface. We also tracked the movements of a model organism, the Cory's shearwater (Calonectris diomedea), a pelagic bird known to perform long distance migrations. Cost models revealed that distant areas can be connected through “wind highways” that do not match the shortest great circle routes. Bird routes closely followed the low-cost “wind-highways” linking breeding and wintering areas. In addition, we found that a potential barrier, the near surface westerlies in the Atlantic sector of the Intertropical Convergence Zone (ITCZ), temporally hindered meridional trans-equatorial movements. Once the westerlies vanished, birds crossed the ITCZ to their winter quarters. This study provides a novel approach to investigate wind-mediated movements in oceanic environments and shows that large-scale migration and dispersal processes over the oceans can be largely driven by spatiotemporal wind patterns.
PMCID: PMC2491555  PMID: 18698354
20.  Migration Routes and Strategies in a Highly Aerial Migrant, the Common Swift Apus apus, Revealed by Light-Level Geolocators 
PLoS ONE  2012;7(7):e41195.
The tracking of small avian migrants has only recently become possible by the use of small light-level geolocators, allowing the reconstruction of whole migration routes, as well as timing and speed of migration and identification of wintering areas. Such information is crucial for evaluating theories about migration strategies and pinpointing critical areas for migrants of potential conservation value. Here we report data about migration in the common swift, a highly aerial and long-distance migrating species for which only limited information based on ringing recoveries about migration routes and wintering areas is available. Six individuals were successfully tracked throughout a complete migration cycle from Sweden to Africa and back. The autumn migration followed a similar route in all individuals, with an initial southward movement through Europe followed by a more southwest-bound course through Western Sahara to Sub-Saharan stopovers, before a south-eastward approach to the final wintering areas in the Congo basin. After approximately six months at wintering sites, which shifted in three of the individuals, spring migration commenced in late April towards a restricted stopover area in West Africa in all but one individual that migrated directly towards north from the wintering area. The first part of spring migration involved a crossing of the Gulf of Guinea in those individuals that visited West Africa. Spring migration was generally wind assisted within Africa, while through Europe variable or head winds were encountered. The average detour at about 50% could be explained by the existence of key feeding sites and wind patterns. The common swift adopts a mixed fly-and-forage strategy, facilitated by its favourable aerodynamic design allowing for efficient use of fuel. This strategy allowed swifts to reach average migration speeds well above 300 km/day in spring, which is higher than possible for similar sized passerines. This study demonstrates that new technology may drastically change our views about migration routes and strategies in small birds, as well as showing the unexpected use of very limited geographical areas during migration that may have important consequences for conservation strategies for migrants.
PMCID: PMC3399846  PMID: 22815968
21.  Testing an Emerging Paradigm in Migration Ecology Shows Surprising Differences in Efficiency between Flight Modes 
PLoS ONE  2012;7(4):e35548.
To maximize fitness, flying animals should maximize flight speed while minimizing energetic expenditure. Soaring speeds of large-bodied birds are determined by flight routes and tradeoffs between minimizing time and energetic costs. Large raptors migrating in eastern North America predominantly glide between thermals that provide lift or soar along slopes or ridgelines using orographic lift (slope soaring). It is usually assumed that slope soaring is faster than thermal gliding because forward progress is constant compared to interrupted progress when birds pause to regain altitude in thermals. We tested this slope-soaring hypothesis using high-frequency GPS-GSM telemetry devices to track golden eagles during northbound migration. In contrast to expectations, flight speed was slower when slope soaring and eagles also were diverted from their migratory path, incurring possible energetic costs and reducing speed of progress towards a migratory endpoint. When gliding between thermals, eagles stayed on track and fast gliding speeds compensated for lack of progress during thermal soaring. When thermals were not available, eagles minimized migration time, not energy, by choosing energetically expensive slope soaring instead of waiting for thermals to develop. Sites suited to slope soaring include ridges preferred for wind-energy generation, thus avian risk of collision with wind turbines is associated with evolutionary trade-offs required to maximize fitness of time-minimizing migratory raptors.
PMCID: PMC3338847  PMID: 22558166
22.  Orientation Cues for High-Flying Nocturnal Insect Migrants: Do Turbulence-Induced Temperature and Velocity Fluctuations Indicate the Mean Wind Flow? 
PLoS ONE  2010;5(12):e15758.
Migratory insects flying at high altitude at night often show a degree of common alignment, sometimes with quite small angular dispersions around the mean. The observed orientation directions are often close to the downwind direction and this would seemingly be adaptive in that large insects could add their self-propelled speed to the wind speed, thus maximising their displacement in a given time. There are increasing indications that high-altitude orientation may be maintained by some intrinsic property of the wind rather than by visual perception of relative ground movement. Therefore, we first examined whether migrating insects could deduce the mean wind direction from the turbulent fluctuations in temperature. Within the atmospheric boundary-layer, temperature records show characteristic ramp-cliff structures, and insects flying downwind would move through these ramps whilst those flying crosswind would not. However, analysis of vertical-looking radar data on the common orientations of nocturnally migrating insects in the UK produced no evidence that the migrants actually use temperature ramps as orientation cues. This suggests that insects rely on turbulent velocity and acceleration cues, and refocuses attention on how these can be detected, especially as small-scale turbulence is usually held to be directionally invariant (isotropic). In the second part of the paper we present a theoretical analysis and simulations showing that velocity fluctuations and accelerations felt by an insect are predicted to be anisotropic even when the small-scale turbulence (measured at a fixed point or along the trajectory of a fluid-particle) is isotropic. Our results thus provide further evidence that insects do indeed use turbulent velocity and acceleration cues as indicators of the mean wind direction.
PMCID: PMC3012097  PMID: 21209956
23.  Bird orientation at high latitudes: flight routes between Siberia and North America across the Arctic Ocean 
Bird migration and orientation at high latitudes are of special interest because of the difficulties associated with different compass systems in polar areas and because of the considerable differences between flight routes conforming to loxodromes (rhumblines) or orthodromes (great circle routes). Regular and widespread east-north-east migration of birds from the northern tundra of Siberia towards North America across the Arctic Ocean (without landmark influences) were recorded by ship-based tracking radar studies in July and August. Field observations indicated that waders, including species such as Phalaropusfulicarius and Calidris melanotos, dominated, but also terns and skuas may have been involved. Analysis of flight directions in relation to the wind showed that these movements are not caused by wind drift. Assuming possible orientation principles based on celestial or geomagnetic cues, different flight trajectories across the Arctic Ocean were calculated: geographical loxodromes, sun compass routes, magnetic loxodromes and magnetoclinic routes. The probabilities of these four alternatives are evaluated on the basis of both the availability of required orientation cues and the predicted flight paths. This evaluation supports orientation along sun compass routes. Because of the longitudinal time displacement sun compass routes show gradually changing compass courses in close agreement with orthodromes. It is suggested that an important migration link between Siberia and North American stopover sites 1000-2500km apart across the Arctic Ocean has evolved based on sun compass orientation along orthodrome-like routes.
PMCID: PMC1690484  PMID: 10693821
24.  Multi-locus phylogeography of the dusky dolphin (Lagenorhynchus obscurus): passive dispersal via the west-wind drift or response to prey species and climate change? 
The dusky dolphin (Lagenorhynchus obscurus) is distributed along temperate, coastal regions of New Zealand, South Africa, Argentina, and Peru where it feeds on schooling anchovy, sardines, and other small fishes and squid tightly associated with temperate ocean sea surface temperatures. Previous studies have suggested that the dusky dolphin dispersed in the Southern Hemisphere eastward from Peru via a linear, temperate dispersal corridor provided by the circumpolar west-wind drift. With new mitochondrial and nuclear DNA sequence data, we propose an alternative phylogeographic history for the dusky dolphin that was structured by paleoceanographic conditions that repeatedly altered the distribution of its temperate prey species during the Plio-Pleistocene.
In contrast to the west-wind drift hypothesis, phylogenetic analyses support a Pacific/Indian Ocean origin, with a relatively early and continued isolation of Peru from other regions. Dispersal of the dusky dolphin into the Atlantic is correlated with the history of anchovy populations, including multiple migrations from New Zealand to South Africa. Additionally, the cooling of the Eastern Equatorial Pacific led to the divergence of anchovy populations, which in turn explains the north-south equatorial transgression of L. obliquidens and the subsequent divergence of L. obscurus in the Southern Hemisphere.
Overall, our study fails to support the west-wind drift hypothesis. Instead, our data indicate that changes in primary productivity and related abundance of prey played a key role in shaping the phylogeography of the dusky dolphin, with periods of ocean change coincident with important events in the history of this temperate dolphin species. Moderate, short-term changes in sea surface temperatures and current systems have a powerful effect on anchovy populations; thus, it is not infeasible that repeated fluctuations in anchovy populations continue to play an important role in the history of coastal dolphin populations.
PMCID: PMC1973136  PMID: 17683557
25.  Increased Flight Altitudes among Migrating Golden Eagles Suggest Turbine Avoidance at a Rocky Mountain Wind Installation 
PLoS ONE  2014;9(3):e93030.
Potential wind-energy development in the eastern Rocky Mountain foothills of British Columbia, Canada, raises concerns due to its overlap with a golden eagle (Aquila chrysaetos) migration corridor. The Dokie 1 Wind Energy Project is the first development in this area and stands as a model for other projects in the area because of regional consistency in topographic orientation and weather patterns. We visually tracked golden eagles over three fall migration seasons (2009–2011), one pre- and two post-construction, to document eagle flight behaviour in relation to a ridge-top wind energy development. We estimated three-dimensional positions of eagles in space as they migrated through our study site. Flight tracks were then incorporated into GIS to ascertain flight altitudes for eagles that flew over the ridge-top area (or turbine string). Individual flight paths were designated to a category of collision-risk based on flight altitude (e.g. flights within rotor-swept height; ≤150 m above ground) and wind speed (winds sufficient for the spinning of turbines; >6.8 km/h at ground level). Eagles were less likely to fly over the ridge-top area within rotor-swept height (risk zone) as wind speed increased, but were more likely to make such crosses under headwinds and tailwinds compared to western crosswinds. Most importantly, we observed a smaller proportion of flights within the risk zone at wind speeds sufficient for the spinning of turbines (higher-risk flights) during post-construction compared to pre-construction, suggesting that eagles showed detection and avoidance of turbines during migration.
PMCID: PMC3966850  PMID: 24671199

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