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The efficiency with which flying animals convert metabolic power to mechanical power dictates an individual's flight behaviour and energy requirements. Despite the significance of this parameter, we lack empirical data on conversion efficiency for most species as in vivo measurements are notoriously difficult to obtain. Furthermore, conversion efficiency is often assumed to be constant across flight speeds, even though the components driving flight power are speed-dependent. We show, through direct measurements of metabolic and aerodynamic power, that conversion efficiency in the migratory bat ( Pipistrellus nathusii ) increases from 7.0 to 10.4% with flight speed. Our findings suggest that peak conversion efficiency in this species occurs near maximum range speed, where the cost of transport is minimized. A meta-analysis of 16 bird and 8 bat species revealed a positive scaling relationship between estimated conversion efficiency and body mass, with no discernible differences between bats and birds. This has profound consequences for modelling flight behaviour as estimates assuming 23% efficiency underestimate metabolic costs for P. nathusii by almost 50% on average (36–62%). Our findings suggest that conversion efficiency may vary around an ecologically relevant optimum speed and provide a crucial baseline for investigating whether this drives variation in conversion efficiency between species.

Vertebrates capable of powered flight rely on wings, muscles that drive their flapping and sensory inputs to the brain allowing for control of the motor output. In birds, the wings are formed of arrangements of adjacent flight feathers (remiges), whereas the wings of bats consist of double-layered skin membrane stretched out between the forelimb skeleton, body and legs. Bird feathers become worn from use and brittle from UV exposure, which leads to loss of function; to compensate, they are renewed (moulted) at regular intervals. Bird feathers and the wings of bats can be damaged by accident. Wing damage and loss of wing surface due to moult almost invariably cause reduced flight performance in measures such as take-off angle and speed. During moult in birds, this is partially counteracted by concurrent mass loss and enlarged flight muscles. Bats have sensory hairs covering their wing surface that provide feedback information about flow; thus, wing damage affects flight speed and turning ability. Bats also have thin, thread-like muscles, distributed within the wing membrane and, if these are damaged, the control of wing camber is lost. Here, I review the effects of wing damage and moult on flight performance in birds, and the consequences of wing damage in bats. I also discuss studies of life-history trade-offs that make use of experimental trimming of flight feathers as a way to handicap parent birds feeding their young.

The degree to which avian migrants return to the same stationary sites to mimic routes from previous years has received more and more attention as the possibility of tracking small to medium avian migrants over multiple annual cycles has increased. Repeated measurements of individuals can potentially inform about their navigation and migration strategies and to what extent the degree of variation observed within and among individuals may reflect the selective potential in the population. Here we perform a k-nearest neighbour analysis along with a repeatability measure to distinguish events in the annual cycle with intra-individual spatial convergence and to quantify the degree of individual consistency and repeatability at those events. To demonstrate the usefulness of our approach we analyse the annual space-use of European nightjars (henceforth nightjars) Caprimulgus europaeus tracked in multiple years between northern Europe and southern Africa. We found that the nightjars consistently used the same breeding and wintering sites but that individual route choice during migration were flexible, but significantly repeatable relative to population level variation during the Sahara-crossing. Thus, the nightjars followed individual-specific flyways while allowing for variation of a few hundred kilometres in the actual route in both autumn and spring. The exception was a strong within-individual convergence down to a few tens of kilometres recorded at the initiation of the trans-Saharan flight in spring. Our results suggest that nightjars have incorporated an individual-specific space-use within their annual cycle, but that they allow for a state-dependent flexibility possibly driven by the cost-benefit balance between the use of known stationary sites and an economical route-choice.

Body mass is a commonly used indicator of the energy stores of migratory animals and there is considerable evidence that it is a critical determinant of migration decisions and outcomes. Mean population mass often increases during the post-breeding period in most migratory species. Usually, this increase is interpreted as the result of fuel accumulation for migration based on the assumption that mean population mass mirrors mean individual mass. However, an empirical test of this assumption is lacking, and it is unknown whether the general increase in mean population mass is entirely the result of within-individual mass gain, or if it rather reflects a change in the nature of individuals in the population (mass-dependent turnover). We investigated changes in body mass during the post-fledging period of a migratory bird, the Red-necked Nightjar (Caprimulgus ruficollis), and combined longitudinal and cross-sectional data collected over 9 years to disentangle the relative contribution of individual-level (mass gain) and population-level (selective appearance and disappearance) processes. We found that the average body mass of fully-developed juveniles increased as the season progressed and that both individual mass gain and the selective disappearance of lighter individuals contributed to this increase. Contrary to the general expectations for migrants, the turnover of individuals contributed 3.5 times more to the seasonal increase in average body mass than individual mass gain. On a practical note, this differential contribution implied a discrepancy of over 40% between the time-average rates of mass gain (fuel deposition rates) estimated from population-level and individual-level data. Our study calls for caution in the use of population-level changes in body mass to make inferences about individual fuel deposition rates and, more generally, indicates that longitudinal and cross-sectional approaches need to be combined to uncover phenotype-time correlations in natural populations.

Body-mounted accelerometers provide a new prospect for estimating power use in flying birds, as the signal varies with the two major kinematic determinants of aerodynamic power: wingbeat frequency and amplitude. Yet wingbeat frequency is sometimes used as a proxy for power output in isolation. There is, therefore, a need to understand which kinematic parameter birds vary and whether this is predicted by flight mode (e.g. accelerating, ascending/descending flight), speed or morphology. We investigate this using high-frequency acceleration data from (i) 14 species flying in the wild, (ii) two species flying in controlled conditions in a wind tunnel and (iii) a review of experimental and field studies. While wingbeat frequency and amplitude were positively correlated, R 2 values were generally low, supporting the idea that parameters can vary independently. Indeed, birds were more likely to modulate wingbeat amplitude for more energy-demanding flight modes, including climbing and take-off. Nonetheless, the striking variability, even within species and flight types, highlights the complexity of describing the kinematic relationships, which appear sensitive to both the biological and physical context. Notwithstanding this, acceleration metrics that incorporate both kinematic parameters should be more robust proxies for power than wingbeat frequency alone.

Space-based tracking technology using low-cost miniature tags is now delivering data on fine-scale animal movement at near-global scale. Linked with remotely sensed environmental data, this offers a biological lens on habitat integrity and connectivity for conservation and human health; a global network of animal sentinels of environmental change. Publisher PDF

Abstract A common migratory pattern in birds is that northerly breeding populations migrate to more southerly non-breeding sites compared to southerly breeding populations (leap-frog migration). Not only do populations experience differences in migration distances, but also different environmental conditions, which may vary spatiotemporally within their annual cycles, creating distinctive selective pressures and migratory strategies. Information about such adaptations is important to understand migratory drivers and evolution of migration patterns. We use light-level geolocators and citizen science data on regional spring arrivals to compare two populations of common ringed plover Charadrius hiaticula breeding at different latitudes. We (1) describe and characterize the annual cycles and (2) test predictions regarding speed and timing of migration. The northern breeding population (NBP) wintered in Africa and the southern (SBP) mainly in Europe. The annual cycles were shifted temporally so that the NBP was always later in all stages. The SBP spent more than twice as long time in the breeding area, but there was no difference in winter. The NBP spent more time on migration in general. Spring migration speed was lower in the SBP compared to autumn speed of both populations, and there was no difference in autumn and spring speed in the NBP. We also found a larger variation in spring arrival times across years in the SBP. This suggests that a complex interaction of population specific timing and variation of breeding onset, length of breeding season, and proximity to the breeding area shape the annual cycle and migratory strategies. Significance statement Migration distance, climate, and the resulting composition of the annual cycle are expected to influence migration strategies and timing in birds. Testing theories regarding migration behaviours are challenging, and intraspecific comparisons over the full annual cycle are still rare. Here we compare the spatiotemporal distributions of two latitudinally separated populations of common ringed plovers using light-level geolocators. We found that there was a larger long-term variation in first arrival dates and that migration speed was slower only in spring in a temperate, short-distance migratory population, compared to an Arctic, long-distance migratory population. This suggests that a complex interaction of population specific timing and variation of breeding onset, length of breeding season and proximity to the breeding area shape the annual cycle and migratory behaviours.

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., Highlighted Article: Migrating nightjars show vertical displacements at low rates throughout migratory flights, indicating regular adjustments of flight altitude while maintaining flapping flight.

The flight speeds that animals should adopt to minimise energy expenditure in different scenarios can be predicted by the curve of power against speed. However, relatively few studies have examined how animals respond to the landscape below them, which could affect speed through modifications in climb rate and perceived predation risk. We equipped homing pigeons(Columba livia)with high frequency loggers to examine how flight speed varies in relation to topography and land cover, predicting that these parameters may have a substantial impact on flight speed and hence cost. Pigeons showed mixed evidence for an energy saving strategy, as they minimised climb rates by starting their ascent ahead of hills, but selected rapid speeds in their ascents. Birds did not modify their speed substantially in relation to land cover, but higher speeds were observed during descending flight, highlighting the importance to consider the rate of change in altitude before estimating power use from speed. Finally, we document an unexpected variability in speed and altitude over fine scales; a source of substantial inefficiency. We suggest this may a form of protean behaviour adopted to reduce predation risk when flocking is not an option, and that such a strategy could be widespread.

Spectacular long-distance migration has evolved repeatedly in animals enabling exploration of resources separated in time and space. In birds, these patterns are largely driven by seasonality, cost of migration, and asymmetries in competition leading most often to leapfrog migration, where northern breeding populations winter furthest to the south. Here, we show that the highly aerial common swiftApus apus, spending the nonbreeding period on the wing, instead exhibits a rarely found chain migration pattern, where the most southern breeding populations in Europe migrate to wintering areas furthest to the south in Africa, whereas the northern populations winter to the north. The swifts concentrated in three major areas in sub-Saharan Africa during the nonbreeding period, with substantial overlap of nearby breeding populations. We found that the southern breeding swifts were larger, raised more young, and arrived to the wintering areas with higher seasonal variation in greenness (Normalized Difference Vegetation Index) earlier than the northern breeding swifts. This unusual chain migration pattern in common swifts is largely driven by differential annual timing and we suggest it evolves by prior occupancy and dominance by size in the breeding quarters and by prior occupancy combined with diffuse competition in the winter.

Hovering insects are divided into two categories: ‘normal’ hoverers that move the wing symmetrically in a horizontal stroke plane, and those with an inclined stroke plane. Normal hoverers have been suggested to support their weight during both downstroke and upstroke, shedding vortex rings each half-stroke. Insects with an inclined stroke plane should, according to theory, produce flight forces only during downstroke, and only generate one set of vortices. The type of hovering is thus linked to the power required to hover. Previous efforts to characterize the wake of hovering insects have used low-resolution experimental techniques or simulated the flow using computational fluid dynamics, and so it remains to be determined whether insect wakes can be represented by any of the suggested models. Here, we used tomographic particle image velocimetry, with a horizontal measurement volume placed below the animals, to show that the wake shed by hovering hawkmoths is best described as a series of bilateral, stacked vortex ‘rings’. While the upstroke is aerodynamically active, despite an inclined stroke plane, it produces weaker vortices than the downstroke. In addition, compared with the near wake, the far wake lacks structure and is less concentrated. Both near and far wakes are clearly affected by vortex interactions, suggesting caution is required when interpreting wake topologies. We also estimated induced power (Pind) from downwash velocities in the wake. Standard models predicted a Pind more than double that from our wake measurements. Our results thus question some model assumptions and we propose a reevaluation of the model parameters.

[Figure][1] Colin making observations during field work in Iceland in 1995. Photo credit: Sverrir Thorstensen. Colin Pennycuick died on 9 December 2019 at the age of 86 years. He was a biologist who made ground breaking contributions to the understanding of animal flight performance, not

Many animals have evolved a migratory lifestyle as an adaptation to seasonality

With timely allocated movement phases, mobile organisms can match their space-use with the seasonality of the environment and thereby optimise their resource utilisation over time. Long-distance avian migrants are known to move with the seasonal dynamics on an annual basis, but how individuals respond to seasonality within their tropical non-breeding range has been less studied. Here we analyse the movement pattern of a highly mobile aerial insectivorous bird, the pallid swift Apus pallidus, and its association with the local habitat phenology during the non-breeding period, using individual-based light-level geolocation. We extracted timing and location of 21 birds’ residence periods, as well as characteristics of the intervening movements, such as distance and speed. We used time series of precipitation and vegetation data for each residence area to extract the timing of the local end of the rainy season and the onset of the dry season. The pallid swifts repeatedly upgraded their habitat by undertaking 2–5 intra-tropical migrations correlated with the withdrawal of the rains and the onset of the local dry season. The birds arrived to the sites on average 12 days after rains ended and departed about two weeks after the onset of dry season suggesting that the birds closely tracked a spatiotemporal window presumably timed with optimal foraging conditions. Our results provide insights in the ways Palaearctic–African migrants respond to the asynchronous phenology within their sub-Saharan non-breeding range. We confirmed that pallid swifts actively respond to deteriorating conditions by repeated upgrades in habitat quality, which likely have substantial consequences for an individual's access to an essential, spatiotemporally ephemeral food resource. However, the pallid swifts did not surf an apparent resource wave per se as would be expected in a highly mobile species, indicating that also other factors, such as spatial patchiness of resources, may influence the movement decision. (Less)

The selection of flight speed (airspeed) in migrating birds depends on multiple internal and external factors, such as wing morphology, weight and winds. Adjustment with respect to side winds to maintain an intended track direction may include a shift in heading direction and/or an increase in airspeed. Compensation for cross-winds cannot always be achieved if visual references are lacking or may not even be beneficial if adaptive wind drift is favourable. Flock size is an additional, although often neglected, factor that could influence the airspeed of birds. Here, we show that responses to cross-winds to achieve compensation differed on a small geographical scale (a few kilometres) in migrating shorebirds, where the availability of topographical features such as coastlines may play an important role for the birds' behaviour. We also show that airspeed was significantly influenced by flock size in three species of shorebirds, increasing with increasing flock size. This is contrary to the prediction based on the hypothesis of energy saving by flight in flock formation, but in agreement with empirical findings for migrating terns. The reason why flock size influences airspeed remains unclear, but we propose a mechanistic explanation based on the largest/heaviest individual(s) determining the speed of the flock.

Migratory birds are known to be capable of adjusting their heading direction to compensate for wind drift and their airspeed adaptively with respect to head and tail winds. High-flying nocturnally migrating common swifts, Apus apus, have been shown to compensate for wind drift, but they failed to adjust airspeed as expected (increase in head wind and decrease in tail wind in relation to neutral wind). We report on new measurements of diurnally migrating common swifts at a coastal site in the Baltic, where the birds did adjust airspeed adaptively during spring and autumn migration. During autumn migration, they compensated for lateral wind drift by adjusting heading direction similarly to high-altitude migrants in autumn. We also recorded flight speed and wind compensation during a summer weather-related exodus, when the birds behaved similarly to those during autumn migration, although they showed a small degree of wind drift. Why birds failed to adjust airspeed adaptively at high altitude is discussed, and we argue there is a threshold in the sensory system to detect small changes in optic flow based on visual landmarks.

Cost of flight at various speeds is a crucial determinant of flight behavior in birds. Aerodynamic models, predicting that mechanical power (Pmech) varies with flight speed in a U-shaped manner, have been used together with an energy conversion factor (efficiency) to estimate metabolic power (Pmet). Despite few empirical studies, efficiency has been assumed constant across flight speeds at 23%. Ideally, efficiency should be estimated from measurements of both Pmech and Pmet in un-instrumented flight. Until recently, progress has been hampered by methodological constraints. The main aim of this study was to evaluate recently developed techniques and estimate flight efficiency across flight speeds. We used the 13C-labeled sodium bicarbonate method (NaBi) and Particle Image Velocimetry (PIV) to measure Pmet and Pmech in blackcaps flying in a wind tunnel. We also cross validated measurements made by NaBi with Quantitative Magnetic Resonance (QMR) body composition analysis in yellow-rumped warblers. We found that Pmet estimated by Nabi was ∼12% lower than corresponding values estimated by QMR. Pmet varied in a U-shaped manner across flight speeds in blackcaps, but the pattern was not statistically significant. Pmech could only be reliably measured for two intermediate speeds and estimated efficiency ranged between 14 and 22% (combining the two speeds for raw and weight/lift specific power, with and without correction for the ∼12% difference between NaBi and QMR) were close to the currently used default value. We conclude that NaBi and PIV are viable techniques, allowing researchers to address some of the outstanding questions regarding bird flight energetics.

We develop an optimization model with two decision variables to explore optimal migration mechanisms to facilitate optimal breeding timing in migratory songbirds. In the model, fitness is a function of date-dependent mortality, speed-dependent predation risk, and phenological match at arrival. The model determines the optimal combination of departure date for spring migration and migration speed, which can be mediated either by the power requirement for flight (P) or foraging effort at stopover sites (k). Our model predicts that earlier departure for spring migration should be the primary mechanism underlying earlier breeding timing, with a lesser role for faster migration via lower P or higher k. In contrast, longer migration to breeding areas selects for both earlier departure and faster migration. Empirical data on sex-specific migration traits largely conform to model predictions, since males generally migrate earlier than females but not faster than females. In contrast, empirical data on age-specific migration traits show some disagreement with model predictions, thus implicating additional tradeoffs. In partial agreement with the model, a comparative analysis of 25 songbird species showed that populations with longer migrations migrate more quickly, but do not initiate migration earlier. Our model proves to be a useful framework for interpreting migration strategies in animals making costly seasonal migrations. (Less)

For studies of how birds control their altitude, seabirds are of particular interest because they forage offshore where the visual environment can be simply modelled by a flat world textured by waves then generating only ventral visual cues. This study suggests that optic flow, i.e. the rate at which the sea moves across the eye’s retina, can explain gulls’ altitude control over seas. In particular, a new flight model that includes both energy and optical invariants helps explain the gulls’ trajectories during offshore takeoff and cruising flight. A linear mixed model applied to 352 flights from 16 individual lesser black backed gulls (Larus fuscus) revealed a statistically significant optic flow set-point ofca25° s−1. Thereafter, an optic flow-based flight model was applied to 18 offshore takeoff flights from nine individual gulls. By introducing an upper limit in climb rate on the elevation dynamics, coupled with an optic flow set-point, the predicted altitude gives an optimized fit factor value of 63% on average (30–83% in range) with respect to the GPS data. We conclude that the optic flow regulation principle helps gulls to adjust their altitude over sea without having to directly measure their current altitude.

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., If the fuelling rate of birds varies periodically, it may act as a synchroniser of migrant populations over large spatial scales. Tracking of the nightjar, a long-distance nocturnal migrant bird, reveals that it adjusts its movements in relation to the lunar cycle.

International audience; Sleep is a universal and complex state and it is widely agreed that this state is present in every animal species. However, the evolutionary origins of sleep remain ignored or misunderstood, which has led researchers to study, in various species, this common behaviour of all living organisms. Sleep is commonly studied at various levels under laboratory conditions, using tethered devices which record electroencephalographic or electromyographic readings. These artificial settings tend to induce stress, reduce animal freedom and prevent the use of sleeping shelters. In this paper, we present a novel, implantable instrumentation for a complete characterization of sleep under natural conditions suitable for a wide range of animal species, even for animals as small as pigeons or mice. Several configurations of this system are possible to enable the measurement of up to 16 electrophysiology channels, 3 temperature channels as well as 3-axes accelerometry. With an embedded flash memory card for the storage of data collected, the system can be used as a datalogger for the recording of signals in the field.

This study documents the advancement of laying dates in three species of tits (Paridae) in southernmost Sweden during recent decades, and the absence of a similar response in the pied flycatcher Ficedula hypoleuca. It is based on several different nestbox studies; the oldest one starting in 1969. During 1969 to 2012, mean spring temperatures in the study area increased by between 0.06 and 0.08°C per year, depending on the period considered. Great tits Parus major, blue tits Cyanistes caeruleus and marsh tits Poecile palustris, which generally start egg laying between the last week of April and the first week of May, all advanced laying date at a similar rate during the study period (0.25 d yr–1). This indicates that these species were similarly affected by increasing temperatures. When accounting for mean spring temperature variation, we still found an advancement of laying date over the study period, mostly due to such relationships among marsh and blue tits. This result could reflect ongoing microevolution favouring earlier laying, but could also be a result of other factors such as increased intra- or inter-specific competition for early breeding. Pied flycatchers, which generally lay during the third week of May, did not significantly advance the date of egg laying despite that the long-term trend in the increase in ambient temperature during the 30-d period preceding the start of egg laying was similar for pied flycatchers compared to the tit species.

Wind tunnels, in which birds fly against an artificially generated air flow, have since long been used to evaluate aerodynamic properties of steady bird flight. A new generation of wind tunnels has also allowed the many processes associated with migratory flights to be studied in captivity. We review how wind tunnel studies of aerodynamics and migratory performance together have helped advancing our understanding of bird migration. Current migration theory is based on the power-speed relationship of flight as well as flight range equations, both of which can be evaluated using birds flying in wind tunnels. In addition, and depending on wind tunnel properties, performance during gliding and climbing flight, and effects of air pressure, humidity and turbulence on bird flight has been measured. Long-distance migrant species have been flown repeatedly for up to 16 h non-stop, allowing detailed studies of the energy expenditure, fuel composition, protein turnover, water balance, immunocompetence and stress associated with sustained migratory flights. In addition, wind tunnels allow the fuelling periods between migratory flights to be studied from new angles. We end our review by suggesting several important topics for future wind tunnel studies, ranging from on of the key questions remaining, the efficiency at which chemical power in converted to mechanical power, to new useful avenues, such as improving and calibrating the techniques used for tracking of individual birds in the wild. (Less)

In this study we compare autumn migration strategies in two subspecies of the Common Ringed Plover Charadrius hiaticula in the southern Baltic Sea. This species exhibits a leap-frog migration pattern, whereby northerly breeding populations (C. h. tundrae; henceforth tundrae) migrate past the whole range of the southern population (C. h. hiaticula; henceforth hiaticula). Hiaticula migrates the shortest distance and is hypothesised to minimize energy spent on migration, whereas tundrae is hypothesized to minimize time, because a longer migration imposes time constraints upon other stages of a migrant's life history, such as moult and breeding. We use biometric data collected at Ottenby Bird Observatory, southern Oland, Sweden, between 1946-2012, to test whether each subspecies demonstrates characteristics associated with either an energy- or time-minimized migration. We used the decline in wing and total-head length over the season to distinguish the subspecies. Hiaticula migrated earlier in the season (July-mid August) compared to tundrae (late July-late September). Also the relative timing of age groups between the two subspecies differed. Juvenile and adult hiaticula migrated synchronized in time, whereas tundrae had two main periods of passage for adults (earlier) and juveniles (later). The timing fits that of other studies and gives complementary information about the passage in Europe. Juvenile tundrae showed a positive trend of observed fuel loads as the season progressed, whereas the other groups did not. Daily fuelling rates within the same season were low compared to other wader species that use Ottenby as a stopover and no difference between subspecies was found. However, tundrae stopped over for a shorter time compared with hiaticula. There was no difference in average migration speed between the subspecies, although tundrae had a higher maximum speed. There was large variation in yearly numbers between age classes in the two subspecies. In hiaticula the yearly average was 8 adults and 37 juveniles. The ratio of juvenile to adult tundrae on the other hand was almost 1:1, indicating equal use of Ottenby as stop over site in autumn. No trends in observed fuel loads over the season in hiaticula are consistent with an energy minimizing migration. As for tundrae, a shorter stopover time compared to hiaticula and a positive relationship between observed fuel load and time, are consistent with time minimizing migration. However, there are many uncertainties and more studies of migratory behaviours at several sites along the migration route are needed to understand the differences in migration strategies in this species.

Summary 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.

For studies of how flying animals control their flight, seabirds are of particular interest to track with a biologger because they forage offshore where the visual environment can be simply modeled by a flat world textured by waves. This study suggests that optic flow can explain gull’s altitude control over seas. In particular, a new flight model that includes both energy and optical invariants (called theventral optic flow regulation) explain the dynamics of gulls’ altitude control during offshore takeoff and cruising flight. A linear statistical model applied to 352 flights from 16 individual lesser black backed gulls (Larus fuscus) gave a strong correlation between wind assistance and gulls’ altitude. Thereafter, an optic flow-based flight model was applied to 18 offshore takeoff flights from 1. individual gulls. By introducing an upper limit in climb rate in a non-linear first order parametric model on the gull’s elevation dynamics, coupled with an optic-flow set-point, the predicted altitude gives an optimized fit factor value of 63% on average (min value: 30%, max value: 83%) with respect to GPS data. We conclude that the optic-flow regulation principle (here running close to 25°/s) allows gulls to adjust their altitude over sea without having to directly measure their current altitude.

Flight activity recorders have recently confirmed that alpine and common swifts spend the majority of their non-breeding period on the wing, which may last 6–10 months. Here we test the hypothesis that the closely related pallid swift, a species with a breeding distribution around the Mediterranean, lead a similar aerial life-style during its migration and wintering periods. The pallid swift usually lays two clutches in one season and therefore spends more time in the breeding area than the common swift. We successfully tracked four pallid swifts with data loggers that record light for geolocation and acceleration every 5 min to monitor flight activity. The birds wintered south of the Sahel in west Africa from the Ivory Coast to Cameroon. The pallid swifts spent the majority of their non-breeding time in flight, especially the first two months after leaving the breeding area in autumn, while a few landing events occurred during the winter. The total time grounded was < 1%, similar to that of the common and alpine swifts. The mass specific flight metabolic rate of swifts is similar to the average non-breeding metabolic rate of a long distance terrestrial migrant, suggesting swifts are not more likely to procure oxidative damage as a consequence of continuous flight than other migrants. The open airspace used by swifts may provide a relatively safe habitat that explain the high survival rate found in swifts.

Differential migration by sex, where one sex migrates further than the other, occurs in many bird species. How this pattern evolves is however little understood. The first aim of this study was to investigate the extent of differential migration in the common ringed plover Charadrius hiaticula, breeding in southeast Sweden, and test three main hypotheses (the social dominance, body size and arrival time hypothesis) regarding the evolution of differential migration. Geolocators were used to derive spatiotemporal data and morphometrics were collected from the studied population. Males migrated 800 km farther compared to females, were marginally larger and there was no statistical difference in spring arrival between the sexes. In contrast to other studies none of the previously proposed hypotheses could account for the observed pattern. An additional finding was that both sexes arrive up to 1.5 months before egg laying, but males initiate territorial behavior upon arrival. Based on these observations we suggest that males have a higher energetic demand, and challenges to meet those, early in the breeding season. Therefore we hypothesize that males arrive to the breeding site with residual fuel reserves accumulated at the wintering site to cover at least parts of these demands. Based on this hypothesis we present a simple model to explain the longer migration by males. The model is contingent on a trade-off between site specific fueling rates (which we assume to increase with decreasing latitude), cost of the extra migration distance and predation risk during fueling. This framework may be applicable to other cases of differential migration, especially in temperate breeding species which exhibit long pre-egg laying periods.

It is essential to gain knowledge about the causes and extent of migratory connectivity between stationary periods of migrants to further the understanding of processes affecting populations, and to allow efficient implementation of conservation efforts throughout the annual cycle. Avian migrants likely use optimal routes with respect to mode of locomotion, orientation and migration strategy, influenced by external factors such as wind and topography. In self-powered flapping flying birds, any increases in fuel loads are associated with added flight costs. Energy-minimizing migrants are therefore predicted to trade-off extended detours against reduced travel across ecological barriers with no or limited foraging opportunities. Here, we quantify the extent of detours taken by different populations of European nightjars Caprimulgus europaeus, to test our predictions that they used routes beneficial according to energetic principles and evaluate the effect of route shape on seasonal migratory connectivity. We combined data on birds tracked from breeding sites along a longitudinal gradient from England to Sweden. We analysed the migratory connectivity between breeding and main non-breeding sites, and en route stopover sites just south of the Sahara desert. We quantified each track's route extension relative to the direct route between breeding and wintering sites, respectively, and contrasted it to the potential detour derived from the barrier reduction along the track while accounting for potential wind effects. Nightjars extended their tracks from the direct route between breeding and main non-breeding sites as they crossed the Mediterranean Sea-Sahara desert, the major ecological barrier in the Palaearctic-African migration system. These clockwise detours were small for birds from eastern sites but increased from east to west breeding longitude. Routes of the tracked birds were associated with partial reduction in the barrier crossing resulting in a trade-off between route extension and barrier reduction, as expected in an energy-minimizing migrant. This study demonstrates how the costs of barrier crossings in prevailing winds can disrupt migratory routes towards slightly different goals, and thereby promote migratory connectivity. This is an important link between individual migration strategies in association with an ecological barrier, and both spatially and demographic population patterns. G.N., A.H. and S. A. are grateful to Urban Rundstrom for valuable help during fieldwork. The project was funded by project grants to A. H. and S.A. from the Swedish Research Council (621-2012-3585, 2016-03625; 621-2013-4361). This is a report from the Centre for Animal Movement Research (CAnMove) funded by a Linnaeus grant from the Swedish Research Council (349-2007-8690) and Lund University. R.E. was funded by a BOF mandate (BSFFCMKDK) at Hasselt University. Belgian research equipment was funded by the Agency for Nature and Forest (ANB) (Belgium), and permissions were granted by the ANB and the Royal Belgian Institute of Natural Sciences. R.E. wishes to thank Eddy Ulenaers, Karen Vanmarcke and Fien Evens for their help during fieldwork. G.C. and I.H. are grateful to the Shoreham & District Ornithological Society, a legacy from Clarice Dawson and Forestry Commission England for funding and support.

Animal flight represents a great challenge and model for biomimetic design efforts. Powered flight at low speeds requires not only appropriate lifting surfaces (wings) and actuator (engine), but also an advanced sensory control system to allow maneuvering in confined spaces, and take-off and landing. Millions of years of evolutionary tinkering has resulted in modern birds and bats, which are achieve controlled maneuvering flight as well as hovering and cruising flight with trans-continental non-stop migratory flights enduring several days in some bird species. Unsteady aerodynamic mechanisms allows for hovering and slow flight in insects, birds and bats, such as for example the delayed stall with a leading edge vortex used to enhance lift at slows speeds. By studying animal flight with the aim of mimicking key adaptations allowing flight as found in animals, engineers will be able to design micro air vehicles of similar capacities.

Animal-attached technologies can be powerful means to quantify space use and behavior; however, there are also ethical implications associated with capturing and instrumenting animals. Furthermore, tagging approaches are not necessarily well-suited for examining the movements of multiple individuals within specific, local areas of interest. Here, we assess a method of quantifying animal space use based on a modified theodolite with an inbuilt laser rangefinder. Using a database of >4200 tracks of migrating birds, we show that detection distance increases with bird body mass (range 5 g to >10 kg). The maximum distance recorded to a bird was 5500 m and measurement error was ≤5 m for targets within this distance range: a level comparable to methods such as GPS tagging. We go on to present a case study where this method was used to assess habitat selection in seabirds operating in dynamic coastal waters close to a tidal turbine. Combining positional data with outputs from a hydrographic model revealed that great cormorants (Phalacrocorax carbo) appeared to be highly selective of current characteristics in space and time, exploiting areas where mean current speeds were