Loop migration in adult marsh harriers Circus aeruginosus,as revealed by satellite telemetry

Loop migration among birds is characterized by the spring route lying consistently west or east of the autumn route. The existence of loops has been explained by general wind conditions or seasonal differences in habitat distribution. Loop migration has predominantly been studied at the population level, for example by analysing ring recoveries. Here we study loop migration of individual marsh harriers Circus aeruginosus tracked by satellite telemetry. We show that despite a generally narrow migration corridor the harriers travelled in a distinct clockwise loop through Africa and southern Europe, following more westerly routes in spring than in autumn. We used the Normalized Difference Vegetation Index (NDVI) to identify potential feeding habitat in Africa. Suitable habitat seemed always more abundant along the western route, both in spring and autumn, and no important stopover site was found along the eastern route. Observed routes did thus not coincide with seasonal variation in habitat availability. However, favourable habitat might be more important during spring migration, when the crossing of the Sahara seems more challenging, and thus habitat availability might play an indirect role in the harriers’ route choice. Grid‐based wind data were used to reconstruct general wind patterns, and in qualitative agreement with the observed loop marsh harriers predominantly encountered westerly winds in Europe and easterly winds in Africa, both in autumn and in spring. By correlating tail‐ and crosswinds with forward and perpendicular movement rates, respectively, we show that marsh harriers are partially drifted by wind. Thus, we tentatively conclude that wind rather than habitat seems to have an overriding effect on the shape of the migration routes of marsh harriers. General wind conditions seem to play an important role also in the evolution of narrow migratory loops as demonstrated for individual marsh harriers.     

Effect of wind,thermal convection,and variation in flight strategies on the daily rhythm and flight paths of migrating raptors at Georgia's Black Sea coast

Every autumn, large numbers of raptors migrate through geographical convergence zones to avoid crossing large bodies of water. At coastal convergence zones, raptors may aggregate along coastlines because of convective or wind conditions. However, the effect of wind and thermal convection on migrating raptors may vary depending on local landscapes and weather, and on the flight strategies of different raptors. From 20 August to 14 October 2008 and 2009, we studied the effect of cloud development and crosswinds on the flight paths of raptors migrating through the eastern Black Sea convergence zone, where coastal lowlands at the foothills of the Pontic Mountains form a geographical bottleneck 5‐km‐wide near Batumi, the capital of the Independent Republic of Ajaria in southwestern Georgia. To identify key correlates of local aggregation, we examined diurnal variation in migration intensity and coastal aggregation of 11 species of raptors categorized based on size and flight strategies. As reported at other convergence zones, migration intensity of large obligate‐soaring species peaked during the core period of thermal activity at mid‐day. When clouds developed over interior mountains and limited thermal convection, these large obligate‐soaring species aggregated near the coast. However, medium‐sized soaring migrants that occasionally use flapping flight did not aggregate at the coast when clouds over the mountains weakened thermal convection. Numbers of alternate soaring‐flapping harriers (Circus spp.) peaked during early morning, with these raptors depending more on flapping flight during a time of day with poor thermal convection. Small sparrowhawks (Accipiter spp.) aggregated at the coast during periods when winds blew offshore, suggesting aggregation caused by wind drift. Thus, weather conditions, including cloud cover and wind speed and direction, can influence the daily rhythm and flight paths of migrating raptors and, therefore, should be accounted for before inferring population trends from migration counts.     

Regional and seasonal flight speeds of soaring migrants and the role of weather conditions at hourly and daily scales

Given that soaring birds travel faster with supportive winds or in good thermal soaring conditions, we expect weather conditions en route of migration to explain commonly observed regional and seasonal patterns in the performance of soaring migrants. We used GPS‐loggers to track 13 honey buzzards and four Montagu's harriers for two to six migrations each. We determined how tailwinds, crosswinds, boundary layer height (a proxy for thermal convection) and precipitation affected hourly speeds, daily distances and daily mean speeds with linear regression models. Honey buzzards mostly travel by soaring while Montagu's harriers supplement soaring with flapping. Therefore, we expect that performance of harriers will be less affected by weather than for buzzards. Weather conditions explained between 30 and 50% of variation in migration performance of both species. Tailwind had the largest effect on hourly speeds, daily mean speeds and daily travel distances. Honey buzzards travelled significantly faster and farther, and Montagu's harriers non‐significantly faster, under better convective conditions. Honey buzzards travelled at slower speeds and shorter distances in crosswinds, whereas harriers maintained high speeds in crosswinds. Weather conditions varied between regions and seasons, and this variation accounted for nearly all regional and seasonal variation in flight performance. Hourly performance was higher than predicted at times when we suspect birds had switched to intermittent or continuous flapping flight, for example during sea‐crossings. The daily travel distance of Montagu's harriers was determined to a significant extent by their daily travel time, which differed between regions, possibly also due to weather conditions. We conclude with the implications of our work for studies on migration phenology and we suggest an important role for high‐resolution telemetry in understanding migratory behavior across entire migratory journeys.     

Why do migrating birds fly along coastlines?

If migrating birds compensate completely for wind drift over land but not over the sea, as indicated by recent radar studies, land-birds facing the sea should, under certain conditions, follow the coastline for some distance rather than depart immediately over the sea. Coastal migration is to be expected if onshore drift exceeds the angle between the birds' preferred track direction and the coastline, as will be the case when this angle is small and strong opposed sidewinds or crosswinds prevail. With offshore drift coasting should occur if the flying time to the goal along a route via the coastline is shorter than that along a direct over-sea route, as will be the case for wide sea-crossings, with strong opposed sidewinds or crosswinds and small angles between the migrants' goal direction and the coastline. Field data support this hypothesis.     

Migrating Montagu's harriers frequently interrupt daily flights in both Europe and Africa

Time budgets are a powerful but hitherto seldom used way to study how migrants organise their bi‐annual travels. We studied daily time budgets of travelling Montagu's harriers Circus pygargus, based on GPS tracking data, in which we were particularly interested in how time budgets differ between regions and seasons, and are affected by wind. We found that Montagu's harriers used a relatively broad daily time window for travelling by starting daily travels just after sunrise and ending daily travels just before sunset. Occasionally, flights were extended into the night. Montagu's harriers frequently interrupted their daily flights for on average 1.5 h d^–1. These interruptions occurred in all regions and seasons. The tracking data during interruptions suggested two different behaviours: in 41% of all interruptions the birds were moving (presumed foraging,) and in 32% they were stationary (presumed resting; the remaining interruptions could not be classified). The interruptions for foraging indicate that Montagu's harriers have a fly‐and‐forage migration strategy (i.e. combine travelling and foraging on the same day), but the interruptions for resting illustrate that their travels comprise of more than fly‐and‐forage behaviour alone. The large number of interruptions for foraging in the Sahara Desert indicates that this region is less hostile for a migrating raptor than presumed previously. Importantly, harriers spent more time on interruptions for resting on days with stronger headwinds, suggesting that interruptions for resting serve a function of waiting for more favourable weather conditions. Daily variation in time budgets was largely explained by wind; harriers flew more hours per day, and interrupted their flights fewer hours per day, on days they experienced stronger tailwinds. In contrast, time budgets were similar between regions and seasons, suggesting that wind rather than landscape and season shape travel routines of Montagu's harriers.     

Nocturnal autumn bird migration at Falsterbo, South Sweden

We investigated the patterns of nocturnal bird migration in autumn 1998 at a coastal site on the Falsterbo peninsula in south-western Sweden, by means of a passive infrared device. In total 17 411 flight paths, including track direction and altitude, of migrating birds were recorded for 68 nights from August to October. Mean migratory traffic rate per night varied between 6 and 6618 birds km⁻¹ h⁻¹, with an average of 1319 birds km⁻¹ h⁻¹. Migration at Falsterbo showed a similar seasonal pattern to that reported for central Europe, with pronounced peaks of migration and intermittent periods with relatively low migratory intensities. Weather factors explained two thirds of the variance in the intensity of bird migration. During nights with intense migration, associated with weak winds, the mean track direction was close to that in central western Europe (225°). Birds usually maintained a constant heading independent of wind directions and, in consequence, were drifted by the wind. The mean orientation clearly differed from that of the nearest coastline, suggesting that the birds did not use the topography below to compensate for wind drift.     

Testing an emerging paradigm in migration ecology shows surprising differences in efficiency between flight modes

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.     

Spatio-temporal distribution of migrating raptors: a comparison of ringing and satellite tracking

We describe the migration performance of three long-distance migrating raptors, osprey Pandion haliaetus , honey buzzard Pernis apivorus and marsh harrier Circus aeruginosus , and one short-distance migrating raptor, common buzzard Buteo buteo based on Swedish ringing recoveries and satellite telemetry, respectively. Tracking by satellite can provide detailed information about the exact timing of migration, migration speed, migration directions, stopover sites, and detours, thereby overcoming many of the potential biases found in ring recoveries. Comparison of the results from these two methods revealed agreement in the geographical distribution of the studied Swedish raptor populations during autumn migration and the winter period. Satellite tracking, nevertheless, provided much more detailed information in Africa and revealed significantly faster migration progress than indicated by ring recoveries. The implications of our findings for interpretation of migratory connectivity and the understanding of migration are discussed.     

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.     

RAPTOR MIGRATION ACROSS THE STRAITS OF GIBRALTAR

The visible migration of birds of prey at Gibraltar is analysed from records kept throughout the spring passages of 1967–70 and the autumn passages of 1967–69. In early spring most visible passage is noted in the afternoons, whereas radar observations by Houghton (1970) indicate passage in the mornings. Later in the year an additional burst of visible passage sometimes occurs in the early morning, but it is concluded that most morning movements take place above visible range. Visible migration is recorded on most days of westerly wind during the migration seasons at Gibraltar. Passage is rarely seen when the winds are easterly. It is argued that under the latter conditions a strong upcurrent of air (standing wave) is formed over Gibraltar, and that this carries nearly all migrants above visible range.
Observations of visible passage elsewhere in the Straits suggest that, in spring, raptors of all species cross on a broad front from Tangier to Ceuta, except Honey Buzzards, which probably cross chiefly near Ceuta. In autumn, all species from northern Europe cross chiefly between Tarifa and to the east of Gibraltar, while birds from western Iberia probably cross mainly near Tarifa.
The periods of passage of the common migrant species are summarised.
On the basis of visual observations and published radar results, it is argued that raptors can compensate for lateral drift by the wind and so fly on chosen courses; but that in very strong cross-winds, e.g. the easterly Levanters, they may have to let themselves be drifted off-course.     

Modeling the Distribution of Migratory Bird Stopovers to Inform Landscape-Scale Siting of Wind Development

Conservation of migratory birds requires understanding the distribution of and potential threats to their migratory habitats. However, although migratory birds are protected under international treaties, few maps have been available to represent migration at a landscape scale useful to target conservation efforts or inform the siting of wind energy developments that may affect migratory birds. To fill this gap, we developed models that predict where four groups of birds concentrate or stopover during their migration through the state of Wyoming, USA: raptors, wetland, riparian and sparse grassland birds. The models were based on existing literature and expert knowledge concerning bird migration behavior and ecology and validated using expert ratings and known occurrences. There was significant agreement between migratory occurrence data and migration models for all groups except raptors, and all models ranked well with experts. We measured the overlap between the migration concentration models and a predictive model of wind energy development to assess the potential exposure of migratory birds to wind development and illustrate the utility of migratory concentration models for landscape-scale planning. Wind development potential is high across 15% of Wyoming, and 73% of this high potential area intersects important migration concentration areas. From 5.2% to 18.8% of each group’s important migration areas was represented within this high wind potential area, with the highest exposures for sparse grassland birds and the lowest for riparian birds. Our approach could be replicated elsewhere to fill critical data gaps and better inform conservation priorities and landscape-scale planning for migratory birds.     

Do Arctic waders use adaptive wind drift?

We analysed five data sets of flight directions of migrating arctic waders in relation to winds, recorded by tracking radar and optical range finder, in order to find out if these birds compensate for wind drift, or allow themselves to be drifted by winds. Our purpose was to investigate whether arctic waders use adaptive wind drift strategies or not. The data sets were collected in Siberia (two sets) and Canada during post-breeding (autumn) migration, and in Mauritania and South Sweden during pre-breeding (spring) migration. Both significant drift and compensation effects were found in three of the data sets, Canada, Mauritania and South Sweden. Almost no compensation was found in birds departing in easterly directions from the Siberian tundra (complete drift), while no drift effect was found in birds departing in westerly directions (complete compensation). There were indications that at least some populations of waders may use an adaptive drift strategy consisting of drift at high altitude and/or in high wind speed combined with compensation at low altitude and/or in lower wind speeds, but support for this idea was rather weak and not consistent. Our results were instead more in accordance with the adaptive drift theory that predicts initial drift during the migratory journey, followed by compensation during later stages as the birds are approaching their destinations. Such a strategy implies that arctic waders, at least adult birds, have the capacity of true navigation. A comparison with earlier studies of migrating arctic waders from different parts of the world show that all results so far may be interpreted in accordance with this general adaptive drift strategy. An element of non-adaptive drift can, however, not be completely ruled out.     

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.     

Social learning of hunting skills in juvenile marsh harriers <Emphasis Type="Italic">Circus aeruginosus</Emphasis>

The impact of social factors on the improvement of hunting skills of juvenile marsh harriers during their first autumnal migration were studied in SE Poland. While foraging with adult birds, juveniles performed more dives on prey both in terms of number of trials and rates. Hunting sessions of juveniles were more efficient in the presence of adults than in the absence of adults. Juveniles hunting with adults and other juveniles could select adequate habitat patches in which access to prey is easier. The role of vertical and horizontal transmission of information in the development of hunting skills in juvenile marsh harrier were confirmed because faster development of hunting ability was achieved in the social hunting after the end of their postfledging dependency period.     

Long-Term Phenological Shifts in Raptor Migration and Climate

Climate change is having a discernible effect on many biological and ecological processes. Among observed changes, modifications in bird phenology have been widely documented. However, most studies have interpreted phenological shifts as gradual biological adjustments in response to the alteration of the thermal regime. Here we analysed a long-term dataset (1980-2010) of short-distance migratory raptors in five European regions. We revealed that the responses of these birds to climate-induced changes in autumn temperatures are abrupt and synchronous at a continental scale. We found that when the temperatures increased, birds delayed their mean passage date of autumn migration. Such delay, in addition to an earlier spring migration, suggests that a significant warming may induce an extension of the breeding-area residence time of migratory raptors, which may eventually lead to residency.     

The problem of estimating wind drift in migrating birds

Whether migrating birds compensate for wind drift or not is a fundamental question in bird migration research. The procedures to demonstrate and quantitatively estimate wind drift or compensation are fraught with difficulties and pitfalls. In this paper, we evaluate four methods that have been used in several studies over the past decades. We evaluate the methods by analysing a model migratory movement with a realistic scatter in flight directions, for the ideal cases of full drift and complete compensation. Results obtained with the different methods are then compared with the "true behaviour" of the model movement, illustrating that spurious patterns of drift and compensation arise in some cases. We also illustrate and evaluate the different methods of estimating drift for a real case, based on tracking radar measurements of bird migration in relation to winds. Calculating the linear regression of mean geographic track (resulting flight direction) and heading directions (directions of the birds' body axis) of a migratory movement under different wind conditions in relation to the angle alpha (the angle between mean track and heading) always provides robust and reliable results. Comparing mean flight directions between occasions with winds from the left and right of the mean flight direction of the whole migratory movement also always provides expected and correct measures of drift. In contrast, regressions of individual flight directions in relation to alpha (the angle between track and heading for the specific individuals or flocks) are liable to produce biased and spurious results, overestimating compensation/overcompensation if following winds dominate in the analysis and overestimating drift/overdrift if opposed winds are dominating. Comparing mean directions for cases with winds from the left and right in relation to individual flight directions also gives biased and spurious results unless there is full variation in wind directions or an equal distribution of crosswinds from left and right. The results of the methodological evaluation and the analysis of the real case indicate that some earlier analyses of wind drift may have to be re-evaluated.     

Environmental influences on the orientation of free-flying nocturnal bird migrants

Autumn nocturnal bird migration was investigated using radar and visual observations. Multivariate analyses assessed the influence of environmental variables on orientation. Two categories of birds were distinguished: (1) shorebirds and waterfowl migrating in flocks; and (2) passerine birds flying singly. These two classes of migrants employ different orientation mechanisms. Landbirds selectively flew with the wind, regardless of its direction or speed or whether the skies were clear or overcast. Shorebirds and waterfowl flew in directions independent of wind in light or moderate winds. The accuracy of passerine orientation was not correlated with any of the variables examined. The passerine orientation observed cannot be accounted for by stellar orientation, but is explicable via orientation on the basis of wind direction.     

Regional wind patterns likely shape a seasonal migration detour

Migrating animals should optimise time and energy use when migrating, travelling directly to their destination. Detours from the most direct route may arise however because of barriers and weather conditions. Identifying how such situations arise from variable weather conditions is crucial to understand population response in the light of increased anthropogenic climate change. Here we used light-level geolocators to follow Cyprus wheatears for their full annual cycle in two separate years migrating between Cyprus, over the Mediterranean and the Sahara to winter in north–east sub-Saharan Africa. We predicted that any route detours would be related to wind conditions experienced during migration. We found that spring migration for all birds included an eastern detour, whilst autumn migrations were direct across the Sahara. The direct autumn migration was likely a consequence of consistent tail-winds, whilst the eastern detour in spring is likely to be more efficient given the wind conditions which are against a direct route. Such variable migration routes shaped by coincidence with prevailing winds are probably common suggesting that some birds may be able to adapt to future changes in wind conditions.     

A RADAR STUDY OF THE AUTUMN MIGRATION OF WOOD PIGEONS COLUMBA PALUMBUS IN SOUTHERN SCANDINAVIA

The migration of Wood Pigeons in southern Scandinavia was studied from 21 September to 10 October 1971 and from 16 September to 15 November 1972 using radar stations supplemented with observations from an aircraft and a network of ground observers. By far the largest quantities of Wood Pigeons migrated after cold front passages with northwesterly to northeasterly tailwinds. Most birds departed on a few days, apparently as a consequence of strong preference for tailwind situations. With northwesterly winds a proportionately high migratory activity was recorded in the Kattegatt area. With northeasterly winds activity was higher in the Baltic area. This allowed the Wood Pigeons to make maximal use of the tailwind component, and their ground speed usually exceeded 80 km/h. The calculated mean air speed was 60 km/h. Their dependence on tailwind was particularly strong when the birds were engaging in long sea-crossings, such as across the Kattegatt. Different coastlines affected the geographical pattern of migration in different ways. Frequently Wood Pigeon flocks flew almost parallel to the coast but some distance off shore, until they finally departed. The deflective force of coastlines was greatest when the birds' ground speed was low, that is, under headwind conditions or in calm weather. Mean track directions measured over two areas in northern Skane, called Inland W and Inland E, situated about 60 km apart, differed by 11, those over the western area being directed more to the south than those over the eastern. No significant correlation with wind directions was found in these areas. Combining data from the whole land area, however, track directions were found to vary from day to day in significant correlation to the wind direction. Mean track directions over the Baltic agreed with those over Skane, but both differed significantly from those over the Kattegatt. Both over the Baltic and over the Kattegatt directions were significantly correlated with wind directions, and showed greater variation than track directions over land. Daily track differences over the Baltic resulted both from differences taking place over the land, and from real wind deflection (drift). Both over the land and over the sea heading directions were correlated with wind directions, suggesting compensatory efforts on the part of the birds. On three days extensive fog covered much of the study area. Wood Pigeons continued to migrate, but certain aberrations in their behaviour were noted. Over land migration was relatively heavier in the west with northwesterly winds and in the east with northeasterly winds. The correlation demonstrated between wind direction and the mean track direction was based upon the fact that populations with different inherent primary directions made up different proportions of the migrating cohorts under different wind conditions (pseudo-drift). The incomplete compensation for wind deflection over the sea is ascribed to the lack of visual orientation cues. The more accurate orientation possible over land suggests one reason for the birds' reluctance to flights across the open sea. When mean track directions of Wood Pigeons in different parts of southern Scandinavia were related to the migratory goals of these birds, it was found that they have to change their primary direction in the course of their journey from breeding to wintering areas.     

Advanced departure dates in long-distance migratory raptors

Evidences for phenological changes in response to climate change are now numerous. One of the most documented changes has been the advance of spring arrival dates in migratory birds. However, the effects of climate change on subsequent events of the annual cycle remain poorly studied and understood. Moreover, the rare studies on autumn migration have mainly concerned passerines. Here, we investigated whether raptor species have changed their autumn migratory phenology during the past 30 years at one of the most important convergent points of western European migration routes in France, the Organbidexka pass, in the Western Pyrenees. Eight out of the 14 studied raptor species showed significant phenological shifts during 1981–2008. Long-distance migrants displayed stronger phenological responses than short-distance migrants, and advanced their mean passage dates significantly. As only some short-distance migrants were found to delay their autumn migration and as their trends in breeding and migrating numbers were not significantly negative, we were not able to show any possible settling process of raptor populations. Negative trends in numbers of migrating raptors were found to be related to weaker phenological responses. Further studies using data from other migration sites are necessary to investigate eventual changes in migration routes and possible settling process.