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1.  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.
doi:10.1371/journal.pone.0017903
PMCID: PMC3064565  PMID: 21464975
2.  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.
doi:10.1098/rsbl.2003.0014
PMCID: PMC1698035  PMID: 12952622
3.  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.
doi:10.1098/rspb.2010.2106
PMCID: PMC3061148  PMID: 20980299
bird orientation; migration; satellite tracking; wind drift
4.  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.
doi:10.1007/s00484-011-0443-4
PMCID: PMC3212683  PMID: 21604153
Wind direction; Synoptic situation; Mediterranean; Bird migration; Wood warbler
5.  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.
doi:10.1371/journal.pone.0052300
PMCID: PMC3536796  PMID: 23300969
6.  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.
doi:10.1098/rspb.2001.1622
PMCID: PMC1088711  PMID: 11375093
7.  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.
doi:10.1371/journal.pone.0092218
PMCID: PMC3958503  PMID: 24643060
8.  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.
doi:10.1098/rspb.2009.1221
PMCID: PMC2842736  PMID: 19889697
orientation behaviour; windborne insect migration; nocturnal boundary layer; atmospheric dispersion models
9.  Stochastic atmospheric assistance and the use of emergency staging sites by migrants 
Numerous animals move vast distances through media with stochastic dynamic properties. Avian migrants must cope with variable wind speeds and directions en route, which potentially jeopardize fine-tuned migration routes and itineraries. We show how unpredictable winds affect flight times and the use of an intermediate staging site by red knots (Calidris canutus canutus) migrating from west Africa to the central north Siberian breeding areas via the German Wadden Sea. A dynamic migration model incorporating wind conditions during flight shows that flight durations between Mauritania and the Wadden Sea vary between 2 and 8 days. The number of birds counted at the only known intermediate staging site on the French Atlantic coast was strongly positively correlated with simulated flight times. In addition, particularly light-weight birds occurred at this location. These independent results support the idea that stochastic wind conditions are the main driver of the use of this intermediate stopover site as an emergency staging area. Because of the ubiquity of stochastically varying media, we expect such emergency habitats to exist in many other migratory systems, both airborne and oceanic. Our model provides a tool to quantify the effect of winds and currents en route.
doi:10.1098/rspb.2009.2112
PMCID: PMC2871836  PMID: 20071381
Calidris canutus canutus; conservation; migration; modelling; stopover; wind assistance
10.  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.
doi:10.1371/journal.pone.0093030
PMCID: PMC3966850  PMID: 24671199
11.  Assessment of the Present and Future Offshore Wind Power Potential: A Case Study in a Target Territory of the Baltic Sea Near the Latvian Coast 
The Scientific World Journal  2013;2013:126428.
Offshore wind energy development promises to be a significant domestic renewable energy source in Latvia. The reliable prediction of present and future wind resources at offshore sites is crucial for planning and selecting the location for wind farms. The overall goal of this paper is the assessment of offshore wind power potential in a target territory of the Baltic Sea near the Latvian coast as well as the identification of a trend in the future wind energy potential for the study territory. The regional climate model CLM and High Resolution Limited Area Model (Hirlam) simulations were used to obtain the wind climatology data for the study area. The results indicated that offshore wind energy is promising for expanding the national electricity generation and will continue to be a stable resource for electricity generation in the region over the 21st century.
doi:10.1155/2013/126428
PMCID: PMC3745973  PMID: 23983619
12.  Characterisation of a transcriptome to find sequence differences between two differentially migrating subspecies of the willow warbler Phylloscopus trochilus 
BMC Genomics  2013;14:330.
Background
Animal migration requires adaptations in morphological, physiological and behavioural traits. Several of these traits have been shown to possess a strong heritable component in birds, but little is known about their genetic architecture. Here we used 454 sequencing of brain-derived transcriptomes from two differentially migrating subspecies of the willow warbler Phylloscopus trochilus to detect genes potentially underlying traits associated with migration.
Results
The transcriptome sequencing resulted in 1.8 million reads following filtering steps. Most of the reads (84%) were successfully mapped to the genome of the zebra finch Taeniopygia gutatta. The mapped reads were situated within at least 12,101 predicted zebra finch genes, with the greatest sequencing depth in exons. Reads that were mapped to intergenic regions were generally located close to predicted genes and possibly located in uncharacterized untranslated regions (UTRs). Out of 85,000 single nucleotide polymorphisms (SNPs) with a minimum sequencing depth of eight reads from each of two subspecies-specific pools, only 55 showed high differentiation, confirming previous studies showing that most of the genetic variation is shared between the subspecies. Validation of a subset of the most highly differentiated SNPs using Sanger sequencing demonstrated that several of them also were differentiated between an independent set of individuals of each subspecies. These SNPs were clustered in two chromosome regions that are likely to be influenced by divergent selection between the subspecies and that could potentially be associated with adaptations to their different migratory strategies.
Conclusions
Our study represents the first large-scale sequencing analysis aiming at detecting genes underlying migratory phenotypes in birds and provides new candidates for genes potentially involved in migration.
doi:10.1186/1471-2164-14-330
PMCID: PMC3660185  PMID: 23672489
454 Transcriptome sequencing; Genetics of migration; Phylloscopus
13.  Assessing the impact of marine wind farms on birds through movement modelling 
Advances in technology and engineering, along with European Union renewable energy targets, have stimulated a rapid growth of the wind power sector. Wind farms contribute to carbon emission reductions, but there is a need to ensure that these structures do not adversely impact the populations that interact with them, particularly birds. We developed movement models based on observed avoidance responses of common eider Somateria mollissima to wind farms to predict, and identify potential measures to reduce, impacts. Flight trajectory data that were  collected post-construction of the Danish Nysted offshore wind farm were used to parameterize competing models of bird movements around turbines. The model most closely fitting the observed data incorporated individual variation in the minimum distance at which birds responded to the turbines. We show how such models can contribute to the spatial planning of wind farms by assessing their extent, turbine spacing and configurations on the probability of birds passing between the turbines. Avian movement models can make new contributions to environmental assessments of wind farm developments, and provide insights into how to reduce impacts that can be identified at the planning stage.
doi:10.1098/rsif.2012.0121
PMCID: PMC3405758  PMID: 22552921
environmental impact assessment; barriers to movement; collision; common eider; Somateria mollissima; Nysted offshore wind farm
14.  Resistance of flight feathers to mechanical fatigue covaries with moult strategy in two warbler species 
Biology Letters  2004;1(1):27-30.
Flight feather moult in small passerines is realized in several ways. Some species moult once after breeding or once on their wintering grounds; others even moult twice. The adaptive significance of this diversity is still largely unknown. We compared the resistance to mechanical fatigue of flight feathers from the chiffchaff Phylloscopus collybita, a migratory species moulting once on its breeding grounds, with feathers from the willow warbler Phylloscopus trochilus, a migratory species moulting in both its breeding and wintering grounds. We found that flight feathers of willow warblers, which have a shaft with a comparatively large diameter, become fatigued much faster than feathers of chiffchaffs under an artificial cyclic bending regime. We propose that willow warblers may strengthen their flight feathers by increasing the diameter of the shaft, which may lead to a more rapid accumulation of damage in willow warblers than in chiffchaffs.
doi:10.1098/rsbl.2004.0244
PMCID: PMC1629048  PMID: 17148120
moult; migration; mechanical fatigue; biomechanics
15.  Nucleotide sequence and genomic organization of bird minisatellites. 
Nucleic Acids Research  1989;17(6):2203-2214.
Two minisatellite loci from a Eurasian songbird, the willow warbler (Phylloscopus trochilus) were isolated, sequenced and used as probes to detect more than 20 related hypervariable loci. In addition, a sequence flanking one of the minisatellite loci was isolated, and used to study a VNTR locus. The bird minisatellites have a repeat unit of either 12 (AGGGAAGGGCTC) or 17 bp (GGGGACAGGGGACACCC), repeated in tandem 40-100 times per locus, and shows partial similarity to the sequence motifs of human minisatellites. These sequences are among the most variable minisatellites known, with the incidence per gamete of new length alleles estimated from family studies of warblers to about 5.6% per locus. The bird minisatellite alleles show mendelian inheritance and segregation analysis indicates that they are derived from families of sequences with members on several autosomal linkage groups. Some of the warbler core sequences cross-hybridize to hypervariable loci in other species of birds, mammals and fishes.
Images
PMCID: PMC317590  PMID: 2704618
16.  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
17.  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.
doi:10.1371/journal.pone.0002928
PMCID: PMC2491555  PMID: 18698354
18.  Soaring Migratory Birds Avoid Wind Farm in the Isthmus of Tehuantepec, Southern Mexico 
PLoS ONE  2014;9(3):e92462.
The number of wind farms operating in the Isthmus of Tehuantepec, southern Mexico, has rapidly increased in recent years; yet, this region serves as a major migration route for various soaring birds, including Turkey Vultures (Cathartes aura) and Swainson's Hawks (Buteo swainsoni). We analyzed the flight trajectories of soaring migrant birds passing the La Venta II wind farm during the two migratory seasons of 2011, to determine whether an avoidance pattern existed or not. We recorded three polar coordinates for the flight path of migrating soaring birds that were detected using marine radar, plotted the flight trajectories and estimated the number of trajectories that intersected the polygon defined by the wind turbines of La Venta II. Finally, we estimated the actual number of intersections per kilometer and compared this value with the null distributions obtained by running 10,000 simulations of our datasets. The observed number of intersections per kilometer fell within or beyond the lower end of the null distributions in the five models proposed for the fall season and in three of the four models proposed for the spring season. Flight trajectories had a non-random distribution around La Venta II, suggesting a strong avoidance pattern during fall and a possible avoidance pattern during spring. We suggest that a nearby ridgeline plays an important role in this pattern, an issue that may be incorporated into strategies to minimize the potential negative impacts of future wind farms on soaring birds. Studies evaluating these issues in the Isthmus of Tehuantepec have not been previously published; hence this work contributes important baseline information about the movement patterns of soaring birds and its relationship to wind farms in the region.
doi:10.1371/journal.pone.0092462
PMCID: PMC3967695  PMID: 24647442
19.  Avian collision risk at an offshore wind farm 
Biology Letters  2005;1(3):296-298.
We have been the first to investigate whether long-lived geese and ducks can detect and avoid a large offshore wind farm by tracking their diurnal migration patterns with radar. We found that the percentage of flocks entering the wind farm area decreased significantly (by a factor 4.5) from pre-construction to initial operation. At night, migrating flocks were more prone to enter the wind farm but counteracted the higher risk of collision in the dark by increasing their distance from individual turbines and flying in the corridors between turbines. Overall, less than 1% of the ducks and geese migrated close enough to the turbines to be at any risk of collision.
doi:10.1098/rsbl.2005.0336
PMCID: PMC1617151  PMID: 17148191
migration; radar; wind turbines; avoidance; collision; waterbirds
20.  Stable isotopes examined across a migratory divide in Scandinavian willow warblers (Phylloscopus trochilus trochilus and Phylloscopus trochilus acredula) reflect their African winter quarters. 
The C and N isotopes of feathers from two subspecies of willow warblers Phylloscopus trochilus trochilus and Phylloscopus trochilus acredula) are isotopically distinct. Our analysis of 138 adult males from 14 sites distributed across Sweden shows that the mean delta15N and delta13C values of subspecies acredula (from latitudes above 63 degrees N) were significantly higher than the mean delta15N and delta13C values of subspecies trochilus (from latitudes below 61 degrees N). The analysed willow warbler feathers had been moulted in the winter quarters and the observed isotopic signatures should thus reflect the isotopic pattern of food assimilated in Africa. The isotopic data observed in Sweden match the cline in morphology, both showing abrupt changes around 62 degrees N. This result agrees with data from ringing recoveries indicating that the two subspecies occupy geographically and isotopically distinct wintering grounds in Africa. Our isotopic data suggest that analysis of stable isotopes of C and N is a promising method to track wintering quarters of European birds that migrate to Africa.
PMCID: PMC1690498  PMID: 10670951
21.  Wind Speed during Migration Influences the Survival, Timing of Breeding, and Productivity of a Neotropical Migrant, Setophaga petechia 
PLoS ONE  2014;9(5):e97152.
Over the course of the annual cycle, migratory bird populations can be impacted by environmental conditions in regions separated by thousands of kilometers. We examine how climatic conditions during discrete periods of the annual cycle influence the demography of a nearctic-neotropical migrant population of yellow warblers (Setophaga petechia), that breed in western Canada and overwinter in Mexico. We demonstrate that wind conditions during spring migration are the best predictor of apparent annual adult survival, male arrival date, female clutch initiation date and, via these timing effects, annual productivity. We find little evidence that conditions during the wintering period influence breeding phenology and apparent annual survival. Our study emphasizes the importance of climatic conditions experienced by migrants during the migratory period and indicates that geography may play a role in which period most strongly impacts migrant populations.
doi:10.1371/journal.pone.0097152
PMCID: PMC4020938  PMID: 24828427
22.  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.
Introduction
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.
Results
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.
Conclusion
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.
doi:10.1186/1742-9994-10-26
PMCID: PMC3665591  PMID: 23663358
Arctic; Capture-recapture; Cormack-Jolly-Seber model; Departure probability; Departure time; Migration speed; Northern wheatear; Optimization; Songbird; Stopover ecology
23.  Flight responses by a migratory soaring raptor to changing meteorological conditions 
Biology Letters  2012;8(5):710-713.
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.
doi:10.1098/rsbl.2012.0359
PMCID: PMC3440984  PMID: 22593085
movement ecology; flight behaviour; migration; flight response; high-resolution GPS–GSM telemetry
24.  Influence of weather conditions on the flight of migrating black storks 
This study tested the potential influence of meteorological parameters (temperature, humidity, wind direction, thermal convection) on different migration characteristics (namely flight speed, altitude and direction and daily distance) in 16 black storks (Ciconia nigra). The birds were tracked by satellite during their entire autumnal and spring migration, from 1998 to 2006. Our data reveal that during their 27-day-long migration between Europe and Africa (mean distance of 4100 km), the periods of maximum flight activity corresponded to periods of maximum thermal energy, underlining the importance of atmospheric thermal convection in the migratory flight of the black stork. In some cases, tailwind was recorded at the same altitude and position as the birds, and was associated with a significant rise in flight speed, but wind often produced a side azimuth along the birds' migratory route. Whatever the season, the distance travelled daily was on average shorter in Europe than in Africa, with values of 200 and 270 km d−1, respectively. The fastest instantaneous flight speeds of up to 112 km h−1 were also observed above Africa. This observation confirms the hypothesis of thermal-dependant flight behaviour, and also reveals differences in flight costs between Europe and Africa. Furthermore, differences in food availability, a crucial factor for black storks during their flight between Europe and Africa, may also contribute to the above-mentioned shift in daily flight speeds.
doi:10.1098/rspb.2010.0422
PMCID: PMC2981984  PMID: 20427337
endangered species; black stork; satellite tracking; migration; meteorological conditions
25.  Experimental Evidence for the Effect of Small Wind Turbine Proximity and Operation on Bird and Bat Activity 
PLoS ONE  2012;7(7):e41177.
The development of renewable energy technologies such as wind turbines forms a vital part of strategies to reduce greenhouse gas emissions worldwide. Although large wind farms generate the majority of wind energy, the small wind turbine (SWT, units generating <50 kW) sector is growing rapidly. In spite of evidence of effects of large wind farms on birds and bats, effects of SWTs on wildlife have not been studied and are likely to be different due to their potential siting in a wider range of habitats. We present the first study to quantify the effects of SWTs on birds and bats. Using a field experiment, we show that bird activity is similar in two distance bands surrounding a sample of SWTs (between 6–18 m hub height) and is not affected by SWT operation at the fine scale studied. At shorter distances from operating turbines (0–5 m), bat activity (measured as the probability of a bat “pass” per hour) decreases from 84% (71–91%) to 28% (11–54%) as wind speed increases from 0 to 14 m/s. This effect is weaker at greater distances (20–25 m) from operating turbines (activity decreases from 80% (65–89%) to 59% (32–81%)), and absent when they are braked. We conclude that bats avoid operating SWTs but that this effect diminishes within 20 m. Such displacement effects may have important consequences especially in landscapes where suitable habitat is limiting. Planning guidance for SWTs is currently lacking. Based on our results we recommend that they are sited at least 20 m away from potentially valuable bat habitat.
doi:10.1371/journal.pone.0041177
PMCID: PMC3408485  PMID: 22859969

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