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Abstract The Alps are a natural barrier for avian broad‐front migration in Central Europe. While most birds that approach the Alps are deflected and circumvent the mountains, some choose to make the crossing. Here, they are funnelled and channelled in valleys, leading to high bird densities. Many Alpine valleys are suitable locations for wind farms, potentially creating a conflict between wind energy production and bird conservation. Collisions can be reduced by temporarily shutting down wind turbines. This however requires timely coordination, either by locally monitoring migration intensity or by extrapolating and forecasting migratory fluxes from other sites. However, little is known about the timing and intensity of bird migration in valleys of the central Alps, especially during spring migration. This study presents a 2‐year quantification of avian migration across the Alps. We collected terrestrial radar data at three sites: two located in Alpine valleys and one in the lowland, close to the northern foothills of the Alps. We found high migration traffic rates (MTR) during both migration seasons in the Alpine valleys, with outstanding numbers of migrants during the spring season. The strong alignment of the flight directions with the main orientation of alpine valleys highlights the importance of valleys and the connected passes in channelling migratory fluxes through the Alps. However, extrapolating migration intensities and forecasting peak migration events for inner Alpine sites is difficult, likely due to how migratory patterns and activity are influenced by the complexity of the local topography and the associated dynamic wind and weather conditions. Instead, we call for year‐round on‐site monitoring of migration intensities and strategies tailored to the local context to reduce the risk of bird strikes at wind turbines in the Alps.

Abstract Background Migrating birds fly non-stop for hours or even for days. They rely mainly on fat as fuel complemented by a certain amount of protein. Studies on homing pigeons and birds flying in a wind-tunnel suggest that the shares of fat and protein on total energy expenditure vary with flight duration and body fat stores. Also, flight behaviour, such as descending flight, is expected to affect metabolism. However, studies on free flying migrant birds under natural conditions are lacking. Methods On a Swiss Alpine pass, we caught three species of nocturnal migrant passerines out of their natural migratory flight. Since most night migrants start soon after dusk, we used time since dusk as a measure of flight duration. We used plasma concentrations of metabolites of the fat, protein, and carbohydrate metabolism as indicators of relative fuel use. We used flight altitudes of birds tracked with radar and with atmospheric pressure loggers to characterize flight behaviour. Results The indicators of fat catabolism (triglycerides, very low-density lipoproteins, glycerol) were positively correlated with body energy stores, supporting earlier findings that birds with high fat stores have a higher fat catabolism. As expected, plasma levels of triglycerides, very low-density lipoproteins, glycerol and ß-hydroxy-butyrate increased at the beginning of the night, indicating that nocturnal migrants increased their fat metabolism directly after take-off. Surprisingly, fat catabolism as well as glucose levels decreased in the second half of the night. Data from radar observations showed that the number of birds aloft, their mean height above ground and vertical flight speed decreased after midnight. Together with the findings from atmospheric pressure-loggers put on three species, this shows that nocturnal migrants migrating over continental Europe descend slowly during about 1.5 h before final landfall at night, which results in 11–30% energy savings according to current flight models. Conclusions We suggest that this slow descent reduces energy demands to an extent which is noticeable in the plasma concentration of lipid, protein, and carbohydrate metabolites. The slow descent may facilitate the search for a suitable resting habitat and serve to refill glycogen stores needed for foraging and predator escape when landed.

Abstract Insects are of increasing conservation concern as a severe decline of both biomass and biodiversity have been reported. At the same time, data on where and when they occur in the airspace is still sparse, and we currently do not know whether their density is linked to the type of landscape above which they occur. Here, we combined data of high-flying insect abundance from six locations across Switzerland representing rural, urban and mountainous landscapes, which was recorded using vertical-looking radar devices. We analysed the abundance of high-flying insects in relation to meteorological factors, daytime, and type of landscape. Air pressure was positively related to insect abundance, wind speed showed an optimum, and temperature and wind direction did not show a clear relationship. Mountainous landscapes showed a higher insect abundance than the other two landscape types. Insect abundance increased in the morning, decreased in the afternoon, had a peak after sunset, and then declined again, though the extent of this general pattern slightly differed between landscape types. We conclude that the abundance of high-flying insects is not only related to abiotic parameters, but also to the type of landscapes and its characteristics, which, on a long-term, should be taken into account for when designing conservation measures for insects.

Abstract Background To understand the ecology of long-distance migrant bird species, it is necessary to study their full annual cycle, including migratory routes and stopovers. This is especially important for species in high-elevation habitats that are particularly vulnerable to environmental change. Here, we investigated both local and global movements during all parts of the annual cycle in a small trans-Saharan migratory bird breeding at high elevation. Methods Recently, multi-sensor geolocators have opened new research opportunities in small-sized migratory organisms. We tagged Northern Wheatears Oenanthe oenanthe from the central-European Alpine population with loggers recording atmospheric pressure and light intensity. We modelled migration routes and identified stopover and non-breeding sites by correlating the atmospheric pressure measured on the birds with global atmospheric pressure data. Furthermore, we compared barrier-crossing flights with other migratory flights and studied the movement behaviour throughout the annual cycle. Results All eight tracked individuals crossed the Mediterranean Sea, using islands for short stops, and made longer stopovers in the Atlas highlands. Single non-breeding sites were used during the entire boreal winter and were all located in the same region of the Sahel. Spring migration was recorded for four individuals with similar or slightly different routes compared to autumn. Migratory flights were typically nocturnal and characterized by fluctuating altitudes, frequently reaching 2000 to 4000 m a.s.l, with a maximum of up to 5150 m. Barrier-crossing flights, i.e., over the sea and the Sahara, were longer, higher, and faster compared to flights above favourable stopover habitat. In addition, we detected two types of altitudinal movements at the breeding site. Unexpected regular diel uphill movements were undertaken from the breeding territories towards nearby roosting sites at cliffs, while regional scale movements took place in response to local meteorological conditions during the pre-breeding period. Conclusion Our data inform on both local and global scale movements, providing new insights into migratory behaviour and local movements in small songbirds. This calls for a wider use of multi-sensor loggers in songbird migration research, especially for investigating both local and global movements in the same individuals.

Abstract Over the past decades, tracking technologies have become more ubiquitous and helped uncover crucial spatiotemporal relationships in nature. In order to apply these technologies to small animals and reduce any potential adverse impact of devices, geopositioning methodologies compatible with lightweight devices are highly sought after. Measured by lightweight geolocators, atmospheric pressure provides an untapped opportunity for global geopositioning, as its natural temporal variation is unique to each location. In this study, we estimate the position of birds by comparing pressure data recorded by the geolocator with reference data from a global weather reanalysis database. The method produces a likelihood map of the position based on (1) a mask of the locations for which the ground‐level elevation matches the pressure measured by the geolocator and (2) a mismatch between the temporal time series measured by the geolocator and the reanalysis dataset. This new method is introduced step by step and applied to 16 tracks of nine long‐ and short‐distance migrant species. Using known positions of double‐tagged individuals (light and pressure data), we demonstrate that our method is almost three times more accurate than light‐based positioning with an average error of 44 km in our trials. In contrast to the traditional light‐based approach, pressure geolocation can provide useful information for short stationary periods (less than a day) and is not affected by the equinox problem nor by any shading effects due to weather or animal behaviour. To facilitate the application of the method, we developed an R package GeoPressureR, together with a user guide GeopressureManual and starting code GeoPressureTemplate. The use of pressure sensors to position animals has the potential to become widespread thanks to the combination of both affordable lightweight devices (100 km), forest‐dwelling species, nocturnal animals and altitudinal migrants.

Abstract Tracking technologies have widely expanded our understanding of bird migration routes, destinations and underlying strategies. However, determining the entire trajectory of small birds equipped with lightweight geolocators remains a challenge. We develop a highly optimized hidden Markov model (HMM) for reconstructing bird trajectories. The observation model is defined by pressure and, optionally, light measurements, while the movement model incorporates wind data to constrain consecutive positions based on realistic airspeeds. To reduce the computational costs associated with a large state space, we prune the HMM states and transitions based on flight and observation constraints to efficiently model the entire trajectory. The approach presented is based on a mathematically exact procedure and is fast to compute. We demonstrate how to compute (1) the most likely trajectory, (2) the marginal probability map of each stationary period, (3) simulated trajectories and (4) the wind conditions (wind support/drift) encountered by the bird during each migratory flight. We construct a version of an HMM optimized for reconstructing a bird's migration trajectory based on lightweight geolocator data. To render this approach easily accessible to researchers, we designed a dedicated R package GeoPressureR (https://raphaelnussbaumer.com/GeoPressureR/).

Abstract During their annual migration, avian migrants alternate stopover periods, for refuelling, with migratory flight bouts. We hypothesise that European Nightjars (Caprimulgus europaeus) adapt their daily migration tactics in association with biomes. We tracked the autumn migration of 24 European Nightjars, from breeding populations in Mongolia, Belgium and UK, using GPS-loggers and multi-sensor data loggers. We quantified crepuscular and nocturnal migration and foraging probabilities, as well as daily travel speed and flight altitude during active migration in response to biomes. Nightjars adopt a rush tactic, reflected in high daily travel speed, flight altitude and high migration probabilities at dusk and at night, when travelling through ecological barriers. Migration is slower in semi-open, hospitable biomes. This is reflected in high foraging probabilities at dusk, lower daily travel speed and lower migration probabilities at dusk. Our study shows how nightjars switch migration tactics during autumn migration, and suggest nightjars alternate between feeding and short migratory flight bouts within the same night when travelling through suitable habitats. How this may affect individuals’ fuel stores and whether different biomes provide refuelling opportunities en route remains to be investigated, to understand how future land-use change may affect migration patterns and survival probabilities.

The northern wheatear Oenanthe oenanthe has an almost circumpolar breeding distribution in the Northern Hemisphere, but all populations migrate to sub‐Saharan Africa in winter. Currently, tracking data suggest two main access routes to the northern continents via the Middle East and the Iberian Peninsula. These routes would require detours for birds breeding in the European Alps. Our aim was to map the migration routes and determine annual schedules for birds breeding in Switzerland and Austria, using light level geolocators. We compared their migration patterns with birds from a lowland breeding population in Germany. Birds from the Alps cross the Mediterranean Sea directly heading straight to their non‐breeding sites. In contrast, birds from Germany travelled further west via the Iberian Peninsula. While the German population initiated autumn migration relatively early, arrival on the wintering sites was nearly synchronous across the three populations. During spring migration, German birds arrived earlier at their breeding grounds than birds from the Alps. A comparison with the literature indicated that the breeding populations in the Alps use their own route and are among the latest to arrive in spring, showing resemblance to the phenology of Arctic breeding populations. Our results indicate that the annual cycle of Alps‐breeding wheatears is influenced primarily by breeding ground conditions, and not solely by migration distance.

Abstract Wind has a significant yet complex effect on bird migration speed. With prevailing south wind, overall migration is generally faster in spring than in autumn. However, studies on the difference in airspeed between seasons have shown contrasting results so far, in part due to their limited geographical or temporal coverage. Using the first full‐year weather radar data set of nocturnal bird migration across western Europe together with wind speed from reanalysis data, we investigate variation of airspeed across season. We additionally expand our analysis of ground speed, airspeed, wind speed, and wind profit variation across time (seasonal and daily) and space (geographical and altitudinal). Our result confirms that wind plays a major role in explaining both temporal and spatial variabilities in ground speed. The resulting airspeed remains relatively constant at all scales (daily, seasonal, geographically and altitudinally). We found that spring airspeed is overall 5% faster in Spring than autumn, but we argue that this number is not significant compared to the biases and limitation of weather radar data. The results of the analysis can be used to further investigate birds' migratory strategies across space and time, as well as their energy use.

Abstract Biological rhythms of nearly all animals on earth are synchronized with natural light and are aligned to day‐and‐night transitions. Here, we test the hypothesis that the lunar cycle affects the nocturnal flight activity of European Nightjars (Caprimulgus europaeus). We describe daily activity patterns of individuals from three different countries across a wide geographic area, during two discrete periods in the annual cycle. Although the sample size for two of our study sites is small, the results are clear in that on average individual flight activity was strongly correlated with both local variation in day length and with the lunar cycle. We highlight the species’ sensitivity to changes in ambient light and its flexibility to respond to such changes in different parts of the world.

Abstract Each year, billions of birds migrate across the continents by day and night through airspaces increasingly altered by human activity, resulting in the deaths of millions of birds every year through collisions with man‐made structures. To reduce these negative impacts on wildlife, forecasts of high migration intensities are needed to apply mitigation actions. While existing weather radar networks offer a unique possibility to monitor and forecast bird migration at large spatial scales, forecasts at the fine spatial scale within a complex terrain, such as the mountainous Swiss landscape, require a small‐scale network of ornithological radars. Before attempting to build such a network, it is crucial to first investigate the consistency of the migratory flow across space and time. In this study, we simultaneously operated three ornithological radar systems across the Swiss lowlands to assess the spatio‐temporal consistency of diurnal and nocturnal bird movements during the spring and autumn migration season. The relative temporal course of migration intensities was generally consistent between sites during peak migration, in particular for nocturnal movements in autumn, but absolute intensities differed greatly between sites. Outside peak migration, bird movement patterns were much less consistent and, unexpectedly, some presumably non‐migratory bird activity achieved intensities close to peak migration intensities, but without spatial correlations. Only nocturnal migration intensity in autumn could be predicted with consistently high accuracy, but including parameters of atmospheric conditions in the model improved predictability of diurnal movements considerably. Predictions for spring were less reliable, probably because we missed an important part of the migration season. Our results show that reliable forecasts of bird movements within a complex terrain call for a network of year‐round bird monitoring systems, whereas accurate information of atmospheric conditions can help to limit the number of measurement points.

Thanks to their light weight and low cost relative to GPS trackers, light-level geolocators are uniquely positioned to uncover bird migration patterns across less well-financed and understudied regions of the world. A main drawback of geolocators is the need to recapture equipped birds to retrieve the data. Maximizing the recapture rate is therefore critical to the success of any geolocator study. In this paper, we present a methodology drawing on historical ringing data in order to inform the deployment of geolocators, both in terms of how many birds can be equipped, and when/which birds to equip in order to maximize retrieval. We illustrate this methodology with a geolocator study of Red-capped Robin-chats (Cossypha natalensis) on the coast of Kenya and find that it accurately estimates how many geolocators to source. It also provides insights into which classes of birds (based on age, capture history, and timing within the season) are most likely to be recaptured. Finally, the analysis of recapture rates allows minimization of geolocator use and thus potential negative impacts to a population.

Abstract Background Over the past decade, the miniaturisation of animal borne tags such as geolocators and GPS-transmitters has revolutionized our knowledge of the whereabouts of migratory species. Novel light-weight multi-sensor loggers (1.4 g), which harbour sensors for measuring ambient light intensity, atmospheric pressure, temperature and acceleration, were fixed to two long-distance migrant bird species - eurasian hoopoe (Upupa epops) and great reed warbler (Acrocephalus arundinaceus). Using acceleration and atmospheric pressure data recorded every 5 and 30 min, respectively, we aimed at reconstructing individual diurnal and seasonal patterns of flight activity and flight altitude and thereby, at describing basic, yet hitherto unknown characteristics of migratory flight behaviour. Furthermore, we wanted to characterise the variability in these migration characteristics between individuals, species and migration periods. Results The flight duration from breeding to sub-Saharan African non-breeding sites and back was more variable within than between the species. Great reed warblers were airborne for a total of 252 flight hours and thus, only slightly longer than eurasian hoopoes with 232 h. With a few exceptions, both species migrated predominantly nocturnally - departure around dusk and landing before dawn. Mean flight altitudes were higher during pre- than during post-breeding migration (median 1100 to 1600 m a.s.l.) and flight above 3000 m occurred regularly with a few great reed warblers exceeding 6000 m a.s.l. (max. 6458 m a.s.l.). Individuals changed flight altitudes repeatedly during a flight bout, indicating a continuous search for (more) favourable flight conditions. Conclusions We found high variation between individuals in the flight behaviour parameters measured – a variation that surprisingly even exceeded the variation between the species. More importantly, our results have shown that multi-sensor loggers have the potential to provide detailed insights into many fundamental aspects of individual behaviour in small aerial migrants. Combining the data recorded on the multiple sensors with, e.g., remote sensing data like weather and habitat quality on the spatial and temporal scale will be a great step forward to explore individual decisions during migration and their consequences.

Airspace is a key but not well-understood habitat for many animal species. Enormous amounts of insects and birds use the airspace to forage, disperse, and migrate. Despite numerous studies on migration, the year-round flight activities of both birds and insects are still poorly studied. We used a 2 year dataset from a vertical-looking radar in Central Europe and developed an iterative hypothesis-testing algorithm to investigate the general temporal pattern of migratory and local movements. We estimated at least 3 million bird and 20 million insect passages over a 1 km transect annually. Most surprisingly, peak non-directional bird movement intensities during summer were of the same magnitude as seasonal directional movement peaks. Birds showed clear peaks in seasonally directional movements during day and night, coinciding well with the main migration period documented in this region. Directional insect movements occurred throughout the year, paralleling non-directional movements. In spring and summer, insect movements were non-directional; in autumn, their movements concentrated toward the southwest, similar to birds. Notably, the nocturnal movements of insects did not appear until April, while directional movements mainly occurred in autumn. This simple monitoring reveals how little we still know about the movement of biomass through airspace.

Recent and archived data from weather radar networks are extensively used for the quantification of continent-wide bird migration patterns. While the process of discriminating birds from weather signals is well established, insect contamination is still a problem. We present a simple method combining two Doppler radar products within a Gaussian mixture model to estimate the proportions of birds and insects within a single measurement volume, as well as the density and speed of birds and insects. This method can be applied to any existing archives of vertical bird profiles, such as the European Network for the Radar surveillance of Animal Movement repository, with no need to recalculate the huge amount of original polar volume data, which often are not available.

Knowledge on whereabouts within the annual cycle of migratory species is prerequisite for many aspects in ecology and biological conservation. Spatial assignments of stable isotopes archived in tissues allows for later inference on sites where the specific tissue had been grown. It has been rarely tested whether spatial assignments match directly tracked non-breeding residences, especially for migratory songbirds. We here compare assignments of stable isotopes from feathers of Palaearctic Barn swallows Hirundo rustica with their African non-breeding residence sites tracked by geolocation.Assignments based on δ2H, δ13C and δ15N isotope compositions delineate three main non-breeding regions: a main cluster in central Africa, a second in West Africa, and the third cluster in Northern Africa. Using δ13C, δ15N only, non-breeding sites ranged from clusters in West/Southwest Africa to South East Africa with a centre in Central Africa. The non-breeding areas (50% and 75% Kernel density estimates, KDE) of the birds tracked by geolocation stretched from West Africa via central Africa to southern Africa. We found little overlap of 0.3% (assuming a 1:1 odds ratio) to 1.4% (3:1 odds ratio) in the three element assignments and KDEs for only 2 and 13 individuals out of 32 birds. Assignment maps for two elements (δ13C, δ15N) and KDEs showed higher consistencies with an overlap of 3.6 and 8.5% for 12 and 18 birds. We argue that the low matching between stable isotope assignments and non-breeding sites in our study arise from insufficient baseline data for Africa (concerning both isoscapes and specific discrimination functions). However, other factors like aerial foraging habit of the species, and a potential mismatch of non-breeding site location and the spatial origin of aerial plankton might further hamper accurate assignments. Finally we call for concerted analyses of tissues i.e. feathers and claws of birds which are grown at known sites across the continent and from species with various ecological requirements (diverse habitats, foraging behaviours, and diet compositions) to establish isoscapes for general applicability.

Quantifying nocturnal bird migration at high resolution is essential for (1) understanding the phenology of migration and its drivers, (2) identifying critical spatio-temporal protection zones for migratory birds, and (3) assessing the risk of collision with artificial structures. We propose a tailored geostatistical model to interpolate migration intensity monitored by a network of weather radars. The model is applied to data collected in autumn 2016 from 69 European weather radars. To validate the model, we performed a cross-validation and also compared our interpolation results with independent measurements of two bird radars. Our model estimated bird densities at high resolution (0.2° latitude−longitude, 15 min) and assessed the associated uncertainty. Within the area covered by the radar network, we estimated that around 120 million birds were simultaneously in flight (10−90 quantiles: 107−134). Local estimations can be easily visualized and retrieved from a dedicated interactive website. This proof-of-concept study demonstrates that a network of weather radar is able to quantify bird migration at high resolution and accuracy. The model presented has the ability to monitor population of migratory birds at scales ranging from regional to continental in space and daily to yearly in time. Near-real-time estimation should soon be possible with an update of the infrastructure and processing software.

Despite benefits of using light-sensitive geolocators to track animal movements and describe patterns of migratory connectivity, concerns have been raised about negative effects of these devices, particularly in small species of aerial insectivore. Geolocators may act as handicaps that increase energetic expenditure, which could explain reported effects of geolocators on survival. We tested this ‘Energetic Expenditure Hypothesis’ in 12 populations of tree swallows (Tachycineta bicolor) and barn swallows (Hirundo rustica) from North America and Europe, using measurements of corticosterone from feathers (CORTf) grown after deployment of geolocators as a measure of physiology relevant to energetics. Contrary to predictions, neither among- (both species) nor within-individual (tree swallows only) levels of CORTf differed with respect to instrumentation. Thus, to the extent that CORTf reflects energetic expenditure, geolocators apparently were not a strong handicap for birds that returned post-deployment. While this physiological evidence suggests that information about migration obtained from returning geolocator-equipped swallows is unbiased with regard to levels of stress, we cannot discount the possibility that corticosterone played a role in reported effects of geolocators on survival in birds, and suggest that future studies relate corticosterone to antecedent factors, such as reproductive history, and to downstream fitness costs.

Populations of most North American aerial insectivores have undergone steep population declines over the past 40 years but the relative importance of factors operating on breeding, wintering, or stopover sites remains unknown. We used archival light-level geolocators to track the phenology, movements and winter locations of barn swallows (Hirdundo rustica; n = 27) from populations across North America to determine their migratory connectivity. We identified an east-west continental migratory divide for barn swallows with birds from western regions (Washington State, USA (n = 8) and Saskatchewan, Canada (n = 5)) traveling shorter distances to wintering areas ranging from Oregon to northern Colombia than eastern populations (Ontario (n = 3) and New Brunswick (n = 10), Canada) which wintered in South America south of the Amazon basin. A single swallow from a stable population in Alabama shared a similar migration route to eastern barn swallows but wintered farther north in northeast Brazil indicating a potential leap frog pattern migratory among eastern birds. Six of 9 (67%) birds from the two eastern populations and Alabama underwent a loop migration west of fall migration routes including around the Gulf of Mexico travelling a mean of 2,224 km and 722 km longer on spring migration, respectively. Longer migration distances, including the requirement to cross the Caribbean Sea and Gulf of Mexico and subsequent shorter sedentary wintering periods, may exacerbate declines for populations breeding in northeastern North America.

Since 1899 ringing (or banding) remained the most important source of information about migration routes, stopover sites and wintering grounds for birds that are too small to carry satellite-based tracking systems. Despite the large quantity of migrating birds ringed in their breeding areas in Europe, the number of ring recoveries from sub-Saharan Africa is very low and therefore the whereabouts of most small bird species outside the breeding season remain a mystery. With new miniaturized light-level geolocators it is now possible to look beyond the limits of ring recovery data. Here we show for the first time year round tracks of a near passerine trans-Saharan migrant, the European Hoopoe (Upupa epops epops). Three birds wintered in the Sahel zone of Western Africa where they remained stationary for most of the time. One bird chose a south-easterly route following the Italian peninsula. Birds from the same breeding population used different migration routes and wintering sites, suggesting a low level of migratory connectivity between breeding and wintering areas. Our tracking of a near passerine bird, the European Hoopoe, with light-level geolocators opens a new chapter in the research of Palaearctic-African bird migration as this new tool revolutionizes our ability to discover migration routes, stopover sites and wintering grounds of small birds.