Early and accumulated experience shape migration and flight in Egyptian vultures.

Two types of experience affect animals' behavioral proficiencies and, accordingly, their fitness: early-life experience, an animal's environment during its early development, and acquired experience, the repeated practice of a specific task.1,2,3,4,5,6,7,8 Yet, how these two experience types and their interactions affect different proficiencies is still an open question. Here, we study the interactions between these two types of experience during migration, a critical and challenging period.9,10 We do so by comparing migratory proficiencies between birds with different early-life experiences and explain these differences by testing fine-scale flight mechanisms. We used data collected by GPS transmitters during 127 autumn migrations of 65 individuals to study the flight proficiencies of two groups of Egyptian vultures (Neophron percnopterus), a long-distance, soaring raptor.11,12 The two groups differed greatly in their early-life experience, one group being captive bred and the other wild hatched.13 Both groups improved their migratory performance with acquired experience, exhibiting shorter migration times, longer daily progress, and improved flight skills, specifically more efficient soaring-gliding behavior. The observed improvements were mostly apparent for captive-bred vultures, which were the least efficient during their first migration but were able to catch up in their migratory performance already in the second migration. Thus, we show how the strong negative effects of early-life experience were offset by acquired experience. Our findings uncover how the interaction between early-life and acquired experiences may shape animals' proficiencies and shed new light on the ontogeny of animal migration, suggesting possible effects of sensitive periods of learning on the acquisition of migratory skills.

Soaring migrants flexibly respond to sea-breeze in a migratory bottleneck: using first derivatives to identify behavioural adjustments over time.

BACKGROUND: Millions of birds travel every year between Europe and Africa detouring ecological barriers and funnelling through migratory corridors where they face variable weather conditions. Little is known regarding the response of migrating birds to mesoscale meteorological processes during flight. Specifically, sea-breeze has a daily cycle that may directly influence the flight of diurnal migrants. METHODS: We collected radar tracks of soaring migrants using modified weather radar in Latrun, central Israel, in 7 autumns between 2005 and 2016. We investigated how migrating soaring birds adjusted their flight speed and direction under the effects of daily sea-breeze circulation. We analysed the effects of wind on bird groundspeed, airspeed and the lateral component of the airspeed as a function of time of day using Generalized Additive Mixed Models. To identify when birds adjusted their response to the wind over time, we estimated first derivatives. RESULTS: Using data collected during a total of 148 days, we characterised the diel dynamics of horizontal wind flow relative to the migration goal, finding a consistent rotational movement of the wind blowing towards the East (morning) and to the South-East (late afternoon), with highest crosswind speed around mid-day and increasing tailwinds towards late afternoon. Airspeed of radar detected birds decreased consistently with increasing tailwind and decreasing crosswinds from early afternoon, resulting in rather stable groundspeed of 16-17 m/s. In addition, birds fully compensated for lateral drift when crosswinds were at their maximum and slightly drifted with the wind when crosswinds decreased and tailwinds became more intense. CONCLUSIONS: Using a simple and broadly applicable statistical method, we studied how wind influences bird flight through speed adjustments over time, providing new insights regarding the flexible behavioural responses of soaring birds to wind conditions. These adjustments allowed the birds to compensate for lateral drift under crosswind and reduced their airspeed under tailwind. Our work enhances our understanding of how migrating birds respond to changing wind conditions during their long-distance journeys through migratory corridors.

A telemetry study shows that an endangered nocturnal avian species roosts in extremely dry habitats to avoid predation.

Describing animal space use is essential for understanding their ecological needs and for planning effective conservation schemes. Notably, certain biomes and life histories are understudied due to methodological challenges in tracking animals in their natural habitats. Specifically, both arid environments and nocturnal species are not sufficiently researched compared to diurnal species and to other biomes. This knowledge gap hinders our ability to properly prioritize habitats for species protection in areas undergoing human-related development. Here, we investigate the movement ecology of the Egyptian Nightjar (Caprimulgus aegyptius) in the arid Dead-sea region of Israel, the Palestinian Authority (the West Bank) and Jordan. This nocturnal insectivore is a cryptic desert-dweller and was considered locally extinct until it was rediscovered in 2016. For this work we tracked twelve individuals using GPS tags to determine how this resource-poor environment affects their home range, (predicting large areas), habitat use, and day-roost ecology. We found that the tracked Egyptian Nightjars had a much larger home range area than other Nightjar species, commuting nightly between foraging grounds and day-roosts. We found, as expected, intensive foraging activity at agricultural fields, where artificial irrigation likely supports higher resource (insect) density. Additionally, we found that individuals showed very high roosting site fidelity, often returning to the same specific site, located in extremely dry and exposed habitats, presumably for predator avoidance. This finding highlights the ecological value of these barren habitats that are often considered "lifeless" and therefore of lower priority for conservation. Consequently, our research demonstrates the importance of describing the space-use of nocturnal animals in arid habitats for conservation efforts.

Time constraints may pace the ontogeny of movement behaviour.

During early development, juvenile animals need to acquire a diverse behavioural repertoire to interact with their environment. The ontogeny of animal behaviour, is paced by the motivation to improve, e.g. internal clocks, and limited by external constraints, e.g. weather conditions. We here evaluate how naive Egyptian vultures (Neophron percnopterus) improve in locomotor performance, measured as daily maximum displacement, prior to their first migration under three different time constraint regimes: we compared wild hatched vultures, migrating one month after fledging, with captive-hatched vultures, released in spring four months or in winter nine months before migration. We found that the time until migration paced the development of movement behaviour: wild birds rapidly increased displacement distances within the first two weeks after fledging, while spring and winter released vultures delayed movement increases by two and four months, respectively. Under relaxed time constraints captive-hatched vultures displayed diverse functional forms of performance enhancements and therefore great variability in individual ontogeny of movement behaviour. While weather conditions in winter could limit flight movements, some birds indeed moved immediately after their release, indicating that weather may not be limiting. Our findings promote the idea that relaxed ecological constraints could uncover hidden phenotypic flexibility in ontogeny, which could present a greater potential for adaptability under environmental change than currently expected.

TrackUSF, a novel tool for automated ultrasonic vocalization analysis, reveals modified calls in a rat model of autism.

BACKGROUND: Various mammalian species emit ultrasonic vocalizations (USVs), which reflect their emotional state and mediate social interactions. USVs are usually analyzed by manual or semi-automated methodologies that categorize discrete USVs according to their structure in the frequency-time domains. This laborious analysis hinders the effective use of USVs as a readout for high-throughput analysis of behavioral changes in animals. RESULTS: Here we present a novel automated open-source tool that utilizes a different approach towards USV analysis, termed TrackUSF. To validate TrackUSF, we analyzed calls from different animal species, namely mice, rats, and bats, recorded in various settings and compared the results with a manual analysis by a trained observer. We found that TrackUSF detected the majority of USVs, with less than 1% of false-positive detections. We then employed TrackUSF to analyze social vocalizations in Shank3-deficient rats, a rat model of autism, and revealed that these vocalizations exhibit a spectrum of deviations from appetitive calls towards aversive calls. CONCLUSIONS: TrackUSF is a simple and easy-to-use system that may be used for a high-throughput comparison of ultrasonic vocalizations between groups of animals of any kind in any setting, with no prior assumptions.

Understanding the ecological and evolutionary function of stopover in migrating birds.

Global movement patterns of migratory birds illustrate their fascinating physical and physiological abilities to cross continents and oceans. During their voyages, most birds land multiple times to make so-called 'stopovers'. Our current knowledge on the functions of stopover is mainly based on the proximate study of departure decisions. However, such studies are insufficient to gauge fully the ecological and evolutionary functions of stopover. If we study how a focal trait, e.g. changes in energy stores, affects the decision to depart from a stopover without considering the trait(s) that actually caused the bird to land, e.g. unfavourable environmental conditions for flight, we misinterpret the function of the stopover. It is thus important to realise and acknowledge that stopovers have many different functions, and that not every migrant has the same (set of) reasons to stop-over. Additionally, we may obtain contradictory results because the significance of different traits to a migrant is context dependent. For instance, late spring migrants may be more prone to risk-taking and depart from a stopover with lower energy stores than early spring migrants. Thus, we neglect that departure decisions are subject to selection to minimise immediate (mortality risk) and/or delayed (low future reproductive output) fitness costs. To alleviate these issues, we first define stopover as an interruption of migratory endurance flight to minimise immediate and/or delayed fitness costs. Second, we review all probable functions of stopover, which include accumulating energy, various forms of physiological recovery and avoiding adverse environmental conditions for flight, and list potential other functions that are less well studied, such as minimising predation, recovery from physical exhaustion and spatiotemporal adjustments to migration. Third, derived from these aspects, we argue for a paradigm shift in stopover ecology research. This includes focusing on why an individual interrupts its migratory flight, which is more likely to identify the individual-specific function(s) of the stopover correctly than departure-decision studies. Moreover, we highlight that the selective forces acting on stopover decisions are context dependent and are expected to differ between, e.g. K-/r-selected species, the sexes and migration strategies. For example, all else being equal, r-selected species (low survival rate, high reproductive rate) should have a stronger urge to continue the migratory endurance flight or resume migration from a stopover because the potential increase in immediate fitness costs suffered from a flight is offset by the expected higher reproductive success in the subsequent breeding season. Finally, we propose to focus less on proximate mechanisms controlling landing and departure decisions, and more on ultimate mechanisms to identify the selective forces shaping stopover decisions. Our ideas are not limited to birds but can be applied to any migratory species. Our revised definition of stopover and the proposed paradigm shift has the potential to stimulate a fruitful discussion towards a better evolutionary ecological understanding of the functions of stopover. Furthermore, identifying the functions of stopover will support targeted measures to conserve and restore the functionality of stopover sites threatened by anthropogenic environmental changes. This is especially important for long-distance migrants, which currently are in alarming decline.

Evolutionary trade-off between male colouration and feather moult extent also indirectly determines female moult.

Males and females are often influenced by different selective forces, frequently resulting in diverging phenotypes, for example in colouration. Since an animal's colouration may strongly influence its fitness, causes and consequences of sexual dichromatism in birds could aid in understanding important factors affecting sexual and natural selection. Variation in plumage ornamentation may affect mate attraction or intraspecific antagonistic behaviour. In most passerines, body plumage colouration of juveniles is obtained through the process of feather moult. The number of moulted wing and tail feathers, which also influences the bird's appearance, may affect its fitness. Here, we show that body plumage colouration of male, but not female, passerines is correlated with the number of moulted wing and tail feathers in the early stage of the bird's life for both sexes. Thus, the extent of wing and tail moult in females is not modulated by the female's colouration and can prevent females from reaching their sex-specific optima. This result could be explained by high intersexual genetic correlations, which might make it impossible for the sexes to reach their own trait fitness optima. Our findings may indicate that species-specific, rather than sex-specific, internal correlations shaped bird moult strategy, an important avian life-history trait.

Body mass and geographic distribution determined the evolution of the wing flight-feather molt strategy in the Neornithes lineage.

The evolutionary history of many organisms is characterized by major changes in morphology and distribution. Specifically, alterations of body mass and geographic distribution may profoundly influence organismal life-history traits. Here, we reconstructed the evolutionary history of flight-feather molt strategy using data from 1,808 Neornithes species. Our analysis suggests that the ancestral molt strategy of first-year birds was partial or entirely absent, and that complete wing flight-feather molt in first-year birds first evolved in the late Eocene and Oligocene (25-40 Ma), at least 30 Myr after birds first evolved. Complete flight-feather molt occurred mainly at equatorial latitudes and in relatively low body mass species, following a diversification of body mass within the lineage. We conclude that both body mass and geographic distribution shaped the evolution of molt strategies and propose that the evolutionary transition towards complete juvenile molt in the Neornithes is a novel, relatively late adaptation.

Differential survival throughout the full annual cycle of a migratory bird presents a life-history trade-off.

Long-distance migrations are among the most physically demanding feats animals perform. Understanding the potential costs and benefits of such behaviour is a fundamental question in ecology and evolution. A hypothetical cost of migration should be outweighed by higher productivity and/or higher annual survival, but few studies on migratory species have been able to directly quantify patterns of survival throughout the full annual cycle and across the majority of a species' range. Here, we use telemetry data from 220 migratory Egyptian vultures Neophron percnopterus, tracked for 3,186 bird months and across approximately 70% of the species' global distribution, to test for differences in survival throughout the annual cycle. We estimated monthly survival probability relative to migration and latitude using a multi-event capture-recapture model in a Bayesian framework that accounted for age, origin, subpopulation and the uncertainty of classifying fates from tracking data. We found lower survival during migration compared to stationary periods (ß = -0.816; 95% credible interval: -1.290 to -0.318) and higher survival on non-breeding grounds at southern latitudes (<25°N; ß = 0.664; 0.076-1.319) compared to on breeding grounds. Survival was also higher for individuals originating from Western Europe (ß = 0.664; 0.110-1.330) as compared to further east in Europe and Asia, and improved with age (ß = 0.030; 0.020-0.042). Anthropogenic mortalities accounted for half of the mortalities with a known cause and occurred mainly in northern latitudes. Many juveniles drowned in the Mediterranean Sea on their first autumn migration while there were few confirmed mortalities in the Sahara Desert, indicating that migration barriers are likely species-specific. Our study advances the understanding of important fitness trade-offs associated with long-distance migration. We conclude that there is lower survival associated with migration, but that this may be offset by higher non-breeding survival at lower latitudes. We found more human-caused mortality farther north, and suggest that increasing anthropogenic mortality could disrupt the delicate migration trade-off balance. Research to investigate further potential benefits of migration (e.g. differential productivity across latitudes) could clarify how migration evolved and how migrants may persist in a rapidly changing world.

Early-life behaviour predicts first-year survival in a long-distance avian migrant.

Early-life conditions have critical, long-lasting effects on the fate of individuals, yet early-life activity has rarely been linked to subsequent survival of animals in the wild. Using high-resolution GPS and body-acceleration data of 93 juvenile white storks (Ciconia ciconia), we examined the links between behaviour during both pre-fledging and post-fledging (fledging-to-migration) periods and subsequent first-year survival. Juvenile daily activity (based on overall dynamic body acceleration) showed repeatable between-individual variation, the juveniles' pre- and post-fledging activity levels were correlated and both were positively associated with subsequent survival. Daily activity increased gradually throughout the post-fledging period, and the relationship between post-fledging activity and survival was stronger in individuals who increased their daily activity level faster (an interaction effect). We suggest that high activity profiles signified individuals with increased pre-migratory experience, higher individual quality and perhaps more proactive personality, which could underlie their superior survival rates. The duration of individuals' fledging-to-migration periods had a hump-shaped relationship with survival: higher survival was associated with intermediate rather than short or long durations. Short durations reflect lower pre-migratory experience, whereas very long ones were associated with slower increases in daily activity level which possibly reflects slow behavioural development. In accordance with previous studies, heavier nestlings and those that hatched and migrated earlier had increased survival. Using extensive tracking data, our study exposed new links between early-life attributes and survival, suggesting that early activity profiles in migrating birds can explain variation in first-year survival.

Seasonal niche tracking of climate emerges at the population level in a migratory bird.

Seasonal animal migration is a widespread phenomenon. At the species level, it has been shown that many migratory animal species track similar climatic conditions throughout the year. However, it remains unclear whether such a niche tracking pattern is a direct consequence of individual behaviour or emerges at the population or species level through behavioural variability. Here, we estimated seasonal niche overlap and seasonal niche tracking at the individual and population level of central European white storks (Ciconia ciconia). We quantified niche tracking for both weather and climate conditions to control for the different spatio-temporal scales over which ecological processes may operate. Our results indicate that niche tracking is a bottom-up process. Individuals mainly track weather conditions while climatic niche tracking mainly emerges at the population level. This result may be partially explained by a high degree of intra- and inter-individual variation in niche overlap between seasons. Understanding how migratory individuals, populations and species respond to seasonal environments is key for anticipating the impacts of global environmental changes.

Sequential Molt in a Feathered Dinosaur and Implications for Early Paravian Ecology and Locomotion.

Feather molt is an important life-history process in birds, but little is known about its evolutionary history. Here, we report on the first fossilized evidence of sequential wing feather molt, a common strategy among extant birds, identified in the Early Cretaceous four-winged dromaeosaurid Microraptor. Analysis of wing feather molt patterns and ecological properties in extant birds imply that Microraptor maintained its flight ability throughout the entire annual cycle, including the molt period. Therefore, we conclude that flight was essential for either its daily foraging or escaping from predators. Our findings propose that the development of sequential molt is the outcome of evolutionary forces to maintain flight capability throughout the entire annual cycle in both extant birds and non-avialan paravian dinosaurs from 120 mya. VIDEO ABSTRACT.

The effects of long-distance migration on the evolution of moult strategies in Western-Palearctic passerines.

Although feathers are the unifying characteristic of all birds, our understanding of the causes, mechanisms, patterns and consequences of the feather moult process lags behind that of other major avian life-history phenomena such as reproduction and long-distance migration. Migration, which evolved in many species of the temperate and arctic zones, requires high energy expenditure to endure long-distance journeys. About a third of Western-Palearctic passerines perform long-distance migrations of thousands of kilometres each year using various morphological, physiological, biomechanical, behavioural and life-history adaptations. The need to include the largely non-overlapping breeding, long-distance migration and feather moult processes within the annual cycle imposes a substantial constraint on the time over which the moult process can take place. Here, we review four feather-moult-related adaptations which, likely due to time constraints, evolved among long-distance Western-Palearctic migrants: (i) increased moult speed; (ii) increased overlap between moult and breeding or migration; (iii) decreased extent of plumage moult; and (iv) moult of part or all of the plumage during the over-wintering period in the tropics rather than in the breeding areas. We suggest that long-distance migration shaped the evolution of moult strategies and increased the diversity of these strategies among migratory passerines. In contrast to this variation, all resident passerines in the Western Palearctic moult immediately after breeding by renewing the entire plumage of adults and in some species also juveniles, while in other species juvenile moult is partial. We identify important gaps in our current understanding of the moult process that should be addressed in the future. Notably, previous studies suggested that the ancestral moult strategy is a post-breeding summer moult in the Western Palearctic breeding areas and that moult during the winter evolved due to the scheduling of long-distance migration immediately after breeding. We offer an alternative hypothesis based on the notion of southern ancestry, proposing that the ancestral moult strategy was a complete moult during the 'northern winter' in the Afro-tropical region in these species, for both adults and juveniles. An important aspect of the observed variation in moult strategies relates to their control mechanisms and we suggest that there is insufficient knowledge regarding the physiological mechanisms that are involved, and whether they are genetically fixed or shaped by environmental factors. Finally, research effort is needed on how global climate changes may influence avian annual routines by altering the scheduling of major processes such as long-distance migration and feather moult.

Early arrival at breeding grounds: Causes, costs and a trade-off with overwintering latitude.

Early arrival at breeding grounds is of prime importance for migrating birds as it is known to enhance breeding success. Adults, males and higher quality individuals typically arrive earlier, and across years, early arrival has been linked to warmer spring temperatures. However, the mechanisms and potential costs of early arrival are not well understood. To deepen the understanding of arrival date differences between individuals and years, we studied them in light of the preceding spring migration behaviour and atmospheric conditions en route. GPS and body acceleration (ACC) data were obtained for 35 adult white storks (Ciconia ciconia) over five years (2012-2016). ACC records were translated to energy expenditure estimates (overall dynamic body acceleration; ODBA) and to behavioural modes, and GPS fixes were coupled with environmental parameters. At the interindividual level (within years), early arrival was attributed primarily to departing earlier for migration and from more northern wintering sites (closer to breeding grounds), rather than to migration speed. In fact, early-departing birds flew slower, experienced weaker thermal uplifts and expended more energy during flight, but still arrived earlier, emphasizing the cost and the significance of early departure. Individuals that wintered further south arrived later at the breeding grounds but did not produce fewer fledglings, presumably due to positive carry-over effects of advantageous wintering conditions (increased precipitation, vegetation productivity and daylight time). Therefore, early arrival increased breeding success only after controlling for wintering latitude. Males arrived slightly ahead of females. Between years, late arrival was linked to colder temperatures en route through two different mechanisms: stronger headwinds causing slower migration and lower thermal uplifts resulting in longer stopovers. This study showed that distinct migratory properties underlie arrival time variation within and between years. It highlighted (a) an overlooked cost of early arrival induced by unfavourable atmospheric conditions during migration, (b) an important fitness trade-off in storks between arrival date and wintering habitat quality and (c) mechanistic explanations for the negative temperature-arrival date correlation in soaring birds. Such understanding of arrival time can facilitate forecasting migrating species responses to climate changes.

Hovering hummingbird wing aerodynamics during the annual cycle. II. Implications of wing feather moult.

Birds usually moult their feathers in a particular sequence which may incur aerodynamic, physiological and behavioural implications. Among birds, hummingbirds are unique species in their sustained hovering flight. Because hummingbirds frequently hover-feed, they must maintain sufficiently high flight capacities even when moulting their flight feathers. A hummingbird wing consists of 10 primary flight feathers whose absence during moult may strongly affect wing performance. Using dynamic similarity rules, we compared time-accurate aerodynamic loads and flow field measurements over several wing geometries that follow the natural feather moult sequence of Calypte anna, a common hummingbird species in western North America. Our results suggest a drop of more than 20% in lift production during the early stages of the moult sequence in which mid-wing flight feathers are moulted. We also found that the wing's ability to generate lift strongly depended on the morphological integrity of the outer primaries and leading-edge. These findings may explain the evolution of wing morphology and moult attributes. Specifically, the high overlap between adjacent wing feathers, especially at the wing tip, and the slow sequential replacement of the wing feathers result in a relatively small reduction in wing surface area during moult with limited aerodynamic implications. We present power and efficiency analyses for hover flight during moult under several plausible scenarios, suggesting that body mass reduction could be a compensatory mechanism that preserves the energetic costs of hover flight.

Hovering hummingbird wing aerodynamics during the annual cycle. I. Complete wing.

The diverse hummingbird family (Trochilidae) has unique adaptations for nectarivory, among which is the ability to sustain hover-feeding. As hummingbirds mainly feed while hovering, it is crucial to maintain this ability throughout the annual cycle-especially during flight-feather moult, in which wing area is reduced. To quantify the aerodynamic characteristics and flow mechanisms of a hummingbird wing throughout the annual cycle, time-accurate aerodynamic loads and flow field measurements were correlated over a dynamically scaled wing model of Anna's hummingbird (Calypte anna). We present measurements recorded over a model of a complete wing to evaluate the baseline aerodynamic characteristics and flow mechanisms. We found that the vorticity concentration that had developed from the wing's leading-edge differs from the attached vorticity structure that was typically found over insects' wings; firstly, it is more elongated along the wing chord, and secondly, it encounters high levels of fluctuations rather than a steady vortex. Lift characteristics resemble those of insects; however, a 20% increase in the lift-to-torque ratio was obtained for the hummingbird wing model. Time-accurate aerodynamic loads were also used to evaluate the time-evolution of the specific power required from the flight muscles, and the overall wingbeat power requirements nicely matched previous studies.

Age-Dependent Modulation of Songbird Summer Feather Molt by Temporal and Functional Constraints.

Time constraints influence various ecological, life-history, and demographic properties of individuals and populations of many species throughout the annual cycle. Feather molt is a timely undertaking that is considered among the three most energy-demanding processes in the life cycle of birds. To deal with time pressure, passerines may shorten their molt duration, using three non-mutually exclusive mechanisms: (1) replacing only part of the plumage, (2) increasing the speed of molt, and (3) postponing the renewal of some or all the plumage to a later season (i.e., from the summer to the overwintering period). We used a comparative approach by measuring 12,349 individuals from 134 passerine species to explore how feather molt of juvenile and adult passerines is evolutionarily modulated under time constraints. The results indicate that breeding at northern latitudes and long-distance migration limit the time available for molt and that the consequences of time constraints were age dependent. While the duration of adult summer molt decreased, the extent, rather than the duration, of juvenile molt declined under time constraints. This study highlights the importance of considering time constraints in order to enhance the understanding of selective forces that shape life-history processes and their consequences throughout the annual routine.

From Agricultural Benefits to Aviation Safety: Realizing the Potential of Continent-Wide Radar Networks.

Migratory animals provide a multitude of services and disservices-with benefits or costs in the order of billions of dollars annually. Monitoring, quantifying, and forecasting migrations across continents could assist diverse stakeholders in utilizing migrant services, reducing disservices, or mitigating human-wildlife conflicts. Radars are powerful tools for such monitoring as they can assess directional intensities, such as migration traffic rates, and biomass transported. Currently, however, most radar applications are local or small scale and therefore substantially limited in their ability to address large-scale phenomena. As weather radars are organized into continent-wide networks and also detect "biological targets," they could routinely monitor aerial migrations over the relevant spatial scales and over the timescales required for detecting responses to environmental perturbations. To tap these unexploited resources, a concerted effort is needed among diverse fields of expertise and among stakeholders to recognize the value of the existing infrastructure and data beyond weather forecasting.

Mass seasonal bioflows of high-flying insect migrants.

Migrating animals have an impact on ecosystems directly via influxes of predators, prey, and competitors and indirectly by vectoring nutrients, energy, and pathogens. Although linkages between vertebrate movements and ecosystem processes have been established, the effects of mass insect "bioflows" have not been described. We quantified biomass flux over the southern United Kingdom for high-flying (>150 meters) insects and show that ~3.5 trillion insects (3200 tons of biomass) migrate above the region annually. These flows are not randomly directed in insects larger than 10 milligrams, which exploit seasonally beneficial tailwinds. Large seasonal differences in the southward versus northward transfer of biomass occur in some years, although flows were balanced over the 10-year period. Our long-term study reveals a major transport process with implications for ecosystem services, processes, and biogeochemistry.

The challenges of the first migration: movement and behaviour of juvenile vs. adult white storks with insights regarding juvenile mortality.

Migration conveys an immense challenge, especially for juvenile birds coping with enduring and risky journeys shortly after fledging. Accordingly, juveniles exhibit considerably lower survival rates compared to adults, particularly during migration. Juvenile white storks (Ciconia ciconia), which are known to rely on adults during their first fall migration presumably for navigational purposes, also display much lower annual survival than adults. Using detailed GPS and body acceleration data, we examined the patterns and potential causes of age-related differences in fall migration properties of white storks by comparing first-year juveniles and adults. We compared juvenile and adult parameters of movement, behaviour and energy expenditure (estimated from overall dynamic body acceleration) and placed this in the context of the juveniles' lower survival rate. Juveniles used flapping flight vs. soaring flight 23% more than adults and were estimated to expend 14% more energy during flight. Juveniles did not compensate for their higher flight costs by increased refuelling or resting during migration. When juveniles and adults migrated together in the same flock, the juvenile flew mostly behind the adult and was left behind when they separated. Juveniles showed greater improvement in flight efficiency throughout migration compared to adults which appears crucial because juveniles exhibiting higher flight costs suffered increased mortality. Our findings demonstrate the conflict between the juveniles' inferior flight skills and their urge to keep up with mixed adult-juvenile flocks. We suggest that increased flight costs are an important proximate cause of juvenile mortality in white storks and likely in other soaring migrants and that natural selection is operating on juvenile variation in flight efficiency.