Solar heating may explain extreme diel flight altitude changes in migrating birds.

Great reed warblers, Acrocephalus arundinaceus,1 and great snipes, Gallinago media,2 exhibit a diel cycle in flight altitudes-flying much higher during the day than the night-when performing migratory flights covering both night and day. One hypothesis proposed to explain this behavior is that the birds face additional heating by solar radiation during daytime and hence must climb to very high, and thus also very cold, altitudes to avoid overheating during daytime flights.1,2 Yet, solar heat gain in birds has been shown to drastically decrease with wind speed,3,4 and the quantitative heating effect by solar radiation on a bird flying with an airspeed of 10 m/s or more is unknown. We analyzed temperature data from multisensor data loggers (MDLs)5,6 placed without direct exposure to solar radiation on great reed warblers (the logger covered by feathers on the back) and great snipes (the logger on the leg, covered from the sun by the tail). We found that logger temperatures were significantly higher (5.9°C-8.8°C in great reed warblers and 4.8°C-5.4°C in great snipes) during the day than during the night in birds flying at the same altitudes (and thus also the same expected ambient air temperatures). These results strongly indicate that the heat balance of the flying birds is indeed affected by solar radiation, which is in accordance with the hypothesis that solar radiation is a key factor causing the remarkable diel cycles in flight altitude observed in these two long-distance migrant bird species.1,2.

Activity patterns throughout the annual cycle in a long-distance migratory songbird, the red-backed shrike Lanius collurio.

BACKGROUND: Long-distance migratory birds undergo complex annual cycles during which they must adjust their behaviour according to the needs and conditions encountered throughout the year. Yet, variation in activity throughout the entire annual cycle has rarely been studied in wild migratory birds. METHODS: We used multisensor data loggers to evaluate the patterns of activity throughout the complete annual cycle of a long-distance migratory bird, the red-backed shrike Lanius collurio. Accelerometer data was used to identify life-history stages and to estimate levels of activity during various phases of the annual cycle. In this study, we analysed the variation in daytime activity along the annual cycle and between migratory and non-migratory days. RESULTS: The birds' daytime activity varied throughout the annual cycle while night-time activity was almost exclusively restricted to migratory flights. The highest daytime activity levels were observed during the breeding season, while it remained low during autumn migration and the winter period. Daytime activity differed between sexes during the breeding period, when the males showed the highest level in activity. During migratory periods, both sexes exhibited a higher daytime activity in spring compared to autumn migration, being particularly high in the final migratory leg towards the breeding ground. The birds showed a lower daytime activity on migratory days (days when a migratory flight took place during the succeeding night) than on non-migratory days during both migratory seasons. CONCLUSIONS: Activity measured during daytime results from a combination of several behaviours, and a high daytime activity during spring migration and the breeding period is possibly reflecting particularly energy-demanding periods in the annual cycle of migratory birds. The use of multisensor data loggers to track annual activity provides us with a full annual perspective on variation in activity in long-distance migratory species, an essential approach for understanding possible critical life-history stages and migration ecology.

Flight altitude dynamics of migrating European nightjars across regions and seasons.

Avian migrants may fly at a range of altitudes, but usually concentrate near strata where a combination of flight conditions is favourable. The aerial environment can have a large impact on the performance of the migrant and is usually highly dynamic, making it beneficial for a bird to regularly check the flight conditions at alternative altitudes. We recorded the migrations between northern Europe and sub-Saharan Africa of European nightjars Caprimulgus europaeus to explore their altitudinal space use during spring and autumn flights and to test whether their climbs and descents were performed according to predictions from flight mechanical theory. Spring migration across all regions was associated with more exploratory vertical flights involving major climbs, a higher degree of vertical displacement within flights, and less time spent in level flight, although flight altitude per se was only higher during the Sahara crossing. The nightjars commonly operated at ascent rates below the theoretical maximum, and periods of descent were commonly undertaken by active flight, and rarely by gliding flight, which has been assumed to be a cheaper locomotion mode during descents. The surprisingly frequent shifts in flight altitude further suggest that nightjars can perform vertical displacements at a relatively low cost, which is expected if the birds can allocate potential energy gained during climbs to thrust forward movement during descents. The results should inspire future studies on the potential costs associated with frequent altitude changes and their trade-offs against anticipated flight condition improvements for aerial migrants.

Extreme altitude changes between night and day during marathon flights of great snipes.

Several factors affect the flight altitude of migratory birds, such as topography, ambient temperature, wind conditions, air humidity, predation avoidance, landmark orientation, and avoiding over-heating from direct sunlight.1-6 Recent tracking of migratory birds over long distances has shown that migrants change flight altitude more commonly and dramatically than previously thought.4-8 The reasons behind these altitude changes are not well understood. In their seasonal migrations between Sweden and sub-Saharan Africa, great snipes Gallinago media make non-stop flights of 4,000-7,000 km, lasting 60-90 h.9,10 Activity and air pressure data from multisensor dataloggers showed that great snipes repeatedly changed altitudes around dawn and dusk, between average cruising heights about 2,000 m (above sea level) at night and around 4,000 m during daytime. Frequency and autocorrelation analyses corroborated a conspicuous diel cycle in flight altitude. Most birds regularly flew at 6,000 m and one bird reached 8,700 m, possibly the highest altitude ever recorded for an identified migrating bird. The diel altitude changes took place independently of climate zone, topography, and habitat overflown. Ambient temperature, wind condition, and humidity have no important diel variation at the high altitudes chosen by great snipes. Instead, improved view for orientation by landmarks, predator avoidance, and not least, seeking cold altitudes at day to counteract heating from direct sunlight are the most plausible explanations for the diel altitude cycle. Together with similar recent findings for a small songbird,6 the great snipes' altitudinal performance sheds new light on the complexity and challenges of migratory flights.

Extreme altitudes during diurnal flights in a nocturnal songbird migrant.

Billions of nocturnally migrating songbirds fly across oceans and deserts on their annual journeys. Using multisensor data loggers, we show that great reed warblers (Acrocephalus arundinaceus) regularly prolong their otherwise strictly nocturnal flights into daytime when crossing the Mediterranean Sea and the Sahara Desert. Unexpectedly, when prolonging their flights, they climbed steeply at dawn, from a mean of 2394 meters above sea level to reach extreme cruising altitudes (mean 5367 and maximum 6267 meters above sea level) during daytime flights. This previously unknown behavior of using exceedingly high flight altitudes when migrating during daytime could be caused by diel variation in ambient temperature, winds, predation, vision range, and solar radiation. Our finding of this notable behavior provides new perspectives on constraints in bird flight and might help to explain the evolution of nocturnal migration.

The lunar cycle drives migration of a nocturnal bird.

Every year, billions of seasonal migrants connect continents by transporting nutrients, energy, and pathogens between distant communities and ecosystems. For animals that power their movements by endogenous energy stores, the daily energy intake rates strongly influence the speed of migration. If access to food resources varies cyclically over the season, migrants sensitive to changes in daily energy intake rates may adjust timing of migration accordingly. As an effect, individuals adjusting to a common temporal cycle are expected to approach synchrony in foraging and movement. A large-scale periodic pattern, such as the dark-light cycle of the moon, could thus synchronize migrations across animal populations. However, such cyclic effects on the temporal regulation of migration has not been considered. Here, we show the temporal influence of the lunar cycle on the movement activity and migration tactics in a visual hunting nocturnal insectivore and long-distance migrant, the European nightjar, Caprimulgus europeaus. We found that the daily foraging activity more than doubled during moonlit nights, likely driven by an increase in light-dependent fuelling opportunities. This resulted in a clear cyclicity also in the intensity of migratory movements, with occasionally up to 100% of the birds migrating simultaneously following periods of full moon. We conclude that cyclic influences on migrants can act as an important regulator of the progression of individuals and synchronize pulses of migratory populations, with possible downstream effects on associated communities and ecosystems.

Hypotheses and tracking results about the longest migration: The case of the arctic tern.

The arctic tern Sterna paradisaea completes the longest known annual return migration on Earth, traveling between breeding sites in the northern arctic and temperate regions and survival/molt areas in the Antarctic pack-ice zone. Salomonsen (1967, Biologiske Meddelelser, Copenhagen Danske Videnskabernes Selskab, 24, 1) put forward a hypothetical comprehensive interpretation of this global migration pattern, suggesting food distribution, wind patterns, sea ice distribution, and molt habits as key ecological and evolutionary determinants. We used light-level geolocators to record 12 annual journeys by eight individuals of arctic terns breeding in the Baltic Sea. Migration cycles were evaluated in light of Salomonsen's hypotheses and compared with results from geolocator studies of arctic tern populations from Greenland, Netherlands, and Alaska. The Baltic terns completed a 50,000 km annual migration circuit, exploiting ocean regions of high productivity in the North Atlantic, Benguela Current, and the Indian Ocean between southern Africa and Australia (sometimes including the Tasman Sea). They arrived about 1 November in the Antarctic zone at far easterly longitudes (in one case even at the Ross Sea) subsequently moving westward across 120-220 degrees of longitude toward the Weddell Sea region. They departed from here in mid-March on a fast spring migration up the Atlantic Ocean. The geolocator data revealed unexpected segregation in time and space between tern populations in the same flyway. Terns from the Baltic and Netherlands traveled earlier and to significantly more easterly longitudes in the Indian Ocean and Antarctic zone than terns from Greenland. We suggest an adaptive explanation for this pattern. The global migration system of the arctic tern offers an extraordinary possibility to understand adaptive values and constraints in complex pelagic life cycles, as determined by environmental conditions (marine productivity, wind patterns, low-pressure trajectories, pack-ice distribution), inherent factors (flight performance, molt, flocking), and effects of predation/piracy and competition.

Ecology of animal migration.

Billions of animals are adapted to a travelling life, making regular return migrations between more or less distant living stations on Earth by swimming, flying, running or walking (Figure 1). Extremely long migrations are completed annually by whales between calving areas in warmer waters and feeding areas at higher latitudes in either hemisphere. The longest oceanic migrations among sea turtles and fish are often undertaken by younger immature individuals during a period of several years before they start their more regular return visits to breeding and spawning sites. Among adult leatherback turtles, intervals of several years between successive breeding events leave enough time for extremely long journeys. Famous among bird migrants are arctic terns, showing the longest known annual migration circuit of about 50,000 km. Bar-tailed godwits breed in Alaska and winter in New Zealand and make the longest known non-stop flapping flights, lasting more than two hundred hours and covering up to 12,000 km across the Pacific Ocean. Their total annual migration circuit extends over 30,000 km covered in three main flights (Figure 1). Although diapause with hibernation as egg, pupae, larvae or adult is an important strategy among insects, there are also examples of impressive migrations. Monarch butterflies complete an annual circuit up to 9,000 km in North America in four generations (for more detail, see the review by Steven Reppert in this issue), and the globe skimmer (a dragonfly) presumably exploits the monsoon rains in India and rainy seasons in southern and equatorial Africa in a 15,000 km circuit in four generations (Figure 1). In comparison with swimmers and flyers, animals that migrate by running or walking cover shorter distances. Caribous migrate between boreal forest and tundra over a total distance of not much more than 1000-2000 km per year. Zebras make the longest migrations in Africa, covering at least 500 km, which is just a little bit longer than the well-known wildebeest migration circuit in Serengeti.

Actogram analysis of free-flying migratory birds: new perspectives based on acceleration logging.

The use of accelerometers has become an important part of biologging techniques for large-sized birds with accelerometer data providing information about flight mode, wing-beat pattern, behaviour and energy expenditure. Such data show that birds using much energy-saving soaring/gliding flight like frigatebirds and swifts can stay airborne without landing for several months. Successful accelerometer studies have recently been conducted also for free-flying small songbirds during their entire annual cycle. Here we review the principles and possibilities for accelerometer studies in bird migration. We use the first annual actograms (for red-backed shrike Lanius collurio) to explore new analyses and insights that become possible with accelerometer data. Actogram data allow precise estimates of numbers of flights, flight durations as well as departure/landing times during the annual cycle. Annual and diurnal rhythms of migratory flights, as well as prolonged nocturnal flights across desert barriers are illustrated. The shifting balance between flight, rest and different intensities of activity throughout the year as revealed by actogram data can be used to analyse exertion levels during different phases of the life cycle. Accelerometer recording of the annual activity patterns of individual birds will open up a new dimension in bird migration research.

Annual 10-Month Aerial Life Phase in the Common Swift Apus apus.

The common swift (Apus apus) is adapted to an aerial lifestyle, where food and nest material are captured in the air. Observations have prompted scientists to hypothesize that swifts stay airborne for their entire non-breeding period [1, 2], including migration into sub-Saharan Africa [3-5]. It is mainly juvenile common swifts that occasionally roost in trees or buildings before autumn migration when weather is bad [1, 6]. In contrast, the North American chimney swift (Chaetura pelagica) and Vaux's swift (C. vauxi) regularly settle to roost in places like chimneys and buildings during migration and winter [7, 8]. Observations of common swifts during the winter months are scarce, and roost sites have never been found in sub-Saharan Africa. In the breeding season, non-breeding individuals usually spend the night airborne [9], whereas adult nesting birds roost in the nest [1]. We equipped common swifts with a micro data logger with an accelerometer to record flight activity (years 1-2) and with a light-level sensor for geolocation (year 2). Our data show that swifts are airborne for >99% of the time during their 10-month non-breeding period; some individuals never settled, but occasional events of flight inactivity occurred in most individuals. Apparent flight activity was lower during the daytime than during the nighttime, most likely due to prolonged gliding episodes during the daytime when soaring in thermals. Our data also revealed that twilight ascents, previously observed during the summer [10], occur throughout the year. The results have important implications for understanding physiological adaptations to endure prolonged periods of flight, including the need to sleep while airborne.

Adaptive strategies in nocturnally migrating insects and songbirds: contrasting responses to wind.

Animals that use flight as their mode of transportation must cope with the fact that their migration and orientation performance is strongly affected by the flow of the medium they are moving in, that is by the winds. Different strategies can be used to mitigate the negative effects and benefit from the positive effects of a moving flow. The strategies an animal can use will be constrained by the relationship between the speed of the flow and the speed of the animal's own propulsion in relation to the surrounding air. Here we analyse entomological and ornithological radar data from north-western Europe to investigate how two different nocturnal migrant taxa, the noctuid moth Autographa gamma and songbirds, deal with wind by analysing variation in resulting flight directions in relation to the wind-dependent angle between the animal's heading and track direction. Our results, from fixed locations along the migratory journey, reveal different global strategies used by moths and songbirds during their migratory journeys. As expected, nocturnally migrating moths experienced a greater degree of wind drift than nocturnally migrating songbirds, but both groups were more affected by wind in autumn than in spring. The songbirds' strategies involve elements of both drift and compensation, providing some benefits from wind in combination with destination and time control. In contrast, moths expose themselves to a significantly higher degree of drift in order to obtain strong wind assistance, surpassing the songbirds in mean ground speed, at the cost of a comparatively lower spatiotemporal migratory precision. Moths and songbirds show contrasting but adaptive responses to migrating through a moving flow, which are fine-tuned to the respective flight capabilities of each group in relation to the wind currents they travel within.

Detection of flow direction in high-flying insect and songbird migrants.

Goal-oriented migrants travelling through the sea or air must cope with the effect of cross-flows during their journeys if they are to reach their destination. In order to counteract flow-induced drift from their preferred course, migrants must detect the mean flow direction, and integrate this information with output from their internal compass, to compensate for the deflection. Animals can potentially sense flow direction by two nonexclusive mechanisms: either indirectly, by visually assessing the effect of the current on their movement direction relative to the ground; or directly, via intrinsic properties of the current. Here, we report the first evidence that nocturnal compass-guided insect migrants use a turbulence-mediated mechanism for directly assessing the wind direction hundreds of metres above the ground. By comparison, we find that nocturnally-migrating songbirds do not use turbulence to detect the flow; instead they rely on visual assessment of wind-induced drift to indirectly infer the flow direction.

Three-dimensional tracking of small aquatic organisms using fluorescent nanoparticles.

Tracking techniques are vital for the understanding of the biology and ecology of organisms. While such techniques have provided important information on the movement and migration of large animals, such as mammals and birds, scientific advances in understanding the individual behaviour and interactions of small (mm-scale) organisms have been hampered by constraints, such as the sizes of existing tracking devices, in existing tracking methods. By combining biology, chemistry and physics we here present a method that allows three-dimensional (3D) tracking of individual mm-sized aquatic organisms. The method is based on in-vivo labelling of the organisms with fluorescent nanoparticles, so-called quantum dots, and tracking of the organisms in 3D via the quantum-dot fluorescence using a synchronized multiple camera system. It allows for the efficient and simultaneous study of the behaviour of one as well as multiple individuals in large volumes of observation, thus enabling the study of behavioural interactions at the community scale. The method is non-perturbing - we demonstrate that the labelling is not affecting the behavioural response of the organisms - and is applicable over a wide range of taxa, including cladocerans as well as insects, suggesting that our methodological concept opens up for new research fields on individual behaviour of small animals. Hence, this offers opportunities to focus on important biological, ecological and behavioural questions never before possible to address.

Convergent patterns of long-distance nocturnal migration in noctuid moths and passerine birds.

Vast numbers of insects and passerines achieve long-distance migrations between summer and winter locations by undertaking high-altitude nocturnal flights. Insects such as noctuid moths fly relatively slowly in relation to the surrounding air, with airspeeds approximately one-third of that of passerines. Thus, it has been widely assumed that windborne insect migrants will have comparatively little control over their migration speed and direction compared with migrant birds. We used radar to carry out the first comparative analyses of the flight behaviour and migratory strategies of insects and birds under nearly equivalent natural conditions. Contrary to expectations, noctuid moths attained almost identical ground speeds and travel directions compared with passerines, despite their very different flight powers and sensory capacities. Moths achieved fast travel speeds in seasonally appropriate migration directions by exploiting favourably directed winds and selecting flight altitudes that coincided with the fastest air streams. By contrast, passerines were less selective of wind conditions, relying on self-powered flight in their seasonally preferred direction, often with little or no tailwind assistance. Our results demonstrate that noctuid moths and passerines show contrasting risk-prone and risk-averse migratory strategies in relation to wind. Comparative studies of the flight behaviours of distantly related taxa are critically important for understanding the evolution of animal migration strategies.

Tracking the small with the smallest--using nanotechnology in tracking zooplankton.

A major problem when studying behavior and migration of small organisms is that many of the questions addressed for larger animals are not possible to formulate due to constraints on tracking smaller animals. In aquatic ecosystems, this problem is particularly problematic for zoo- and phytoplankton, since tracking devices are too heavy to allow the organism to act naturally. However, recent advances in nanotechnology have made it possible to track individual animals and thereby to focus on important and urgent questions which previously have not been possible to address. Here we report on a novel approach to track movement and migratory behavior of millimeter sized aquatic animals, particularly Daphnia magna, using the commercially available nanometer sized fluorescent probes known as quantum dots. Experimental trials with and without quantum dots showed that they did not affect behavior, reproduction or mortality of the tested animals. Compared to previously used methods to label small animals, the nano-labeling method presented here offers considerable improvements including: 24 h fluorescence, studies in both light and darkness, much improved optical properties, potential to study large volumes and even track animals in semi-natural conditions. Hence, the suggested method, developed in close cooperation between biologists, chemists and physicists, offers new opportunities to routinely study zooplankton responses to light, food and predation, opening up advancements within research areas such as diel vertical/horizontal migration, partial migration and other differences in intra- and interspecific movements and migration.

Migratory and resident blue tits Cyanistes caeruleus differ in their reaction to a novel object.

Individuals differ consistently in their behavioural reactions towards novel objects and new situations. Reaction to novelty is one part of a suit of individually consistent behaviours called coping strategies or personalities and is often summarised as bold or shy behaviour. Coping strategies could be particularly important for migrating birds exposed to novel environments on their journeys. We compared the average approach latencies to a novel object among migrants and residents in partially migratory blue tits Cyanistes caeruleus. In this test, we found migrating blue tits to have shorter approach latencies than had resident ones. Behavioural reactions to novelty can affect the readiness to migrate and short approach latency may have an adaptive value during migration. Individual behaviour towards novelty might be incorporated among the factors associated with migratory or resident behaviour in a partially migratory population.

A polar system of intercontinental bird migration.

Studies of bird migration in the Beringia region of Alaska and eastern Siberia are of special interest for revealing the importance of bird migration between Eurasia and North America, for evaluating orientation principles used by the birds at polar latitudes and for understanding the evolutionary implications of intercontinental migratory connectivity among birds as well as their parasites. We used tracking radar placed onboard the ice-breaker Oden to register bird migratory flights from 30 July to 19 August 2005 and we encountered extensive bird migration in the whole Beringia range from latitude 64 degrees N in Bering Strait up to latitude 75 degrees N far north of Wrangel Island, with eastward flights making up 79% of all track directions. The results from Beringia were used in combination with radar studies from the Arctic Ocean north of Siberia and in the Beaufort Sea to make a reconstruction of a major Siberian-American bird migration system in a wide Arctic sector between longitudes 110 degrees E and 130 degrees W, spanning one-third of the entire circumpolar circle. This system was estimated to involve more than 2 million birds, mainly shorebirds, terns and skuas, flying across the Arctic Ocean at mean altitudes exceeding 1 km (maximum altitudes 3-5 km). Great circle orientation provided a significantly better fit with observed flight directions at 20 different sites and areas than constant geographical compass orientation. The long flights over the sea spanned 40-80 degrees of longitude, corresponding to distances and durations of 1400-2600 km and 26-48 hours, respectively. The birds continued from this eastward migration system over the Arctic Ocean into several different flyway systems at the American continents and the Pacific Ocean. Minimization of distances between tundra breeding sectors and northerly stopover sites, in combination with the Beringia glacial refugium and colonization history, seemed to be important for the evolution of this major polar bird migration system.

Flight speeds among bird species: allometric and phylogenetic effects.

Flight speed is expected to increase with mass and wing loading among flying animals and aircraft for fundamental aerodynamic reasons. Assuming geometrical and dynamical similarity, cruising flight speed is predicted to vary as (body mass)(1/6) and (wing loading)(1/2) among bird species. To test these scaling rules and the general importance of mass and wing loading for bird flight speeds, we used tracking radar to measure flapping flight speeds of individuals or flocks of migrating birds visually identified to species as well as their altitude and winds at the altitudes where the birds were flying. Equivalent airspeeds (airspeeds corrected to sea level air density, Ue) of 138 species, ranging 0.01-10 kg in mass, were analysed in relation to biometry and phylogeny. Scaling exponents in relation to mass and wing loading were significantly smaller than predicted (about 0.12 and 0.32, respectively, with similar results for analyses based on species and independent phylogenetic contrasts). These low scaling exponents may be the result of evolutionary restrictions on bird flight-speed range, counteracting too slow flight speeds among species with low wing loading and too fast speeds among species with high wing loading. This compression of speed range is partly attained through geometric differences, with aspect ratio showing a positive relationship with body mass and wing loading, but additional factors are required to fully explain the small scaling exponent of Ue in relation to wing loading. Furthermore, mass and wing loading accounted for only a limited proportion of the variation in Ue. Phylogeny was a powerful factor, in combination with wing loading, to account for the variation in Ue. These results demonstrate that functional flight adaptations and constraints associated with different evolutionary lineages have an important influence on cruising flapping flight speed that goes beyond the general aerodynamic scaling effects of mass and wing loading.

Magnetic compass orientation in European robins is dependent on both wavelength and intensity of light.

Magnetic compass orientation in birds has been shown to be light dependent. Results from behavioural studies indicate that magnetoreception capabilities are disrupted under light of peak wavelengths longer than 565 nm, and shifts in orientation have been observed at higher light intensities (43-44x10(15) quanta s(-1) m(-2)). To investigate further the function of the avian magnetic compass with respect to wavelength and intensity of light, we carried out orientation cage experiments with juvenile European robins, caught during their first autumn migration, exposed to light of 560.5 nm (green), 567.5 nm (green-yellow) and 617 nm (red) wavelengths at three different intensities (1 mW m(-2), 5 mW m(-2) and 10 mW m(-2)). We used monochromatic light of a narrow wavelength range (half bandwidth of 9-11 nm, compared with half bandwidths ranging between 30 nm and 70 nm used in other studies) and were thereby able to examine the magnetoreception mechanism in the expected transition zone between oriented and disoriented behaviour around 565 nm in more detail. We show (1) that European robins show seasonally appropriate migratory directions under 560.5 nm light, (2) that they are completely disoriented under 567.5 nm light under a broad range of intensities, (3) that they are able to orient under 617 nm light of lower intensities, although into a direction shifted relative to the expected migratory one, and (4) that magnetoreception is intensity dependent, leading to disorientation under higher intensities. Our results support the hypothesis that birds possess a light-dependent magnetoreception system based on magnetically sensitive, antagonistically interacting spectral mechanisms, with at least one high-sensitive short-wavelength mechanism and one low-sensitive long-wavelength mechanism.

Harmonic oscillatory orientation relative to the wind in nocturnal roosting flights of the swift Apus apus.

Swifts regularly spend the night flying at high altitude. From previous studies based on tracking radar observations, we know that they stay airborne during the night and prefer to orient themselves into the wind direction with an increased angular concentration with increasing wind speed. In this study, we investigated the orientation relative to the wind of individual swifts by frequency (discrete Fourier transform) and autocorrelation analysis based on time series (10s intervals) of the angle between the swifts' heading and the wind direction for radar trackings of long duration (9-60 min). The swifts often showed a significant harmonic oscillation of their heading direction relative to the wind, with a frequency mostly in the range 1-17 mHz, corresponding to cycle periods of 1-16 min. The swifts also sometimes performed circling flights at low wind speeds. Wind speed ranged from 1.3 to 14.8 m s(-1), and we expected to find different patterns of orientation at different wind speeds, assuming that the swifts adapt their orientation to avoid substantial displacement during their nocturnal flights. However, oscillatory orientation was found at all wind speeds with variable frequencies/periods that did not show any consistent relationship with wind speed. It remains to be shown whether cyclic heading changes are a regular feature of bird orientation.