Experience reduces route selection for conspecifics by the collectively migrating white stork.

Migration can be an energetically costly behavior with strong fitness consequences in terms of mortality and reproduction.1,2,3,4,5,6,7,8,9,10,11 Migrants should select migratory routes to minimize their costs, but both costs and benefits may change with experience.12,13,14 This raises the question of whether experience changes how individuals select their migratory routes. Here, we investigate the effect of age on route selection criteria in a collectively migrating soaring bird, the white stork (Ciconia ciconia). We perform step-selection analysis on a longitudinal dataset tracking 158 white storks over up to 9 years to quantify how they select their routes based on the social and atmospheric environments and to examine how this selection changes with age. We find clear ontogenetic shifts in route selection criteria. Juveniles choose routes that have good atmospheric conditions and high conspecific densities. Yet, as they gain experience, storks' selection on the availability of social information reduces-after their fifth migration, experienced birds also choose routes with low conspecific densities. Thus, our results suggest that as individuals age, they gradually replace information gleaned from other individuals with information gained from experience, allowing them to shift their migration timing and increasing the timescale at which they select their routes.

Learning shapes the development of migratory behavior.

How animals refine migratory behavior over their lifetime (i.e., the ontogeny of migration) is an enduring question with important implications for predicting the adaptive capacity of migrants in a changing world. Yet, our inability to monitor the movements of individuals from early life onward has limited our understanding of the ontogeny of migration. The exploration-refinement hypothesis posits that learning shapes the ontogeny of migration in long-lived species, resulting in greater exploratory behavior early in life followed by more rapid and direct movement during later life. We test the exploration-refinement hypothesis by examining how white storks (Ciconia ciconia) balance energy, time, and information as they develop and refine migratory behavior during the first years of life. Here, we show that young birds reduce energy expenditure during flight while also increasing information gain by exploring new places during migration. As the birds age and gain more experience, older individuals stop exploring new places and instead move more quickly and directly, resulting in greater energy expenditure during migratory flight. During spring migration, individuals innovated novel shortcuts during the transition from early life into adulthood, suggesting a reliance on spatial memory acquired through learning. These incremental refinements in migratory behavior provide support for the importance of individual learning within a lifetime in the ontogeny of long-distance migration.

The price of being late: short- and long-term consequences of a delayed migration timing.

Choosing the right migration timing is critical for migrants because conditions encountered en route influence movement costs, survival, and, in social migrants, the availability of social information. Depending on lifetime stages, individuals may migrate at different times due to diverging constraints, affecting the composition of migration groups. To examine the consequences of a delayed migration timing, we artificially delayed the migration of juvenile white storks (Ciconia ciconia) and thereby altered their physical and social environment. Using nearly continuous 1 Hz GPS trajectories, we examined their migration behaviour, ranging from sub-second level performance to global long-distance movement, in relation to two control groups. We found that delayed storks experienced suboptimal soaring conditions, but better wind support and thereby achieved higher flight speeds than control storks. Delayed storks had a lower mortality rate than the control storks and wintered closer to the breeding area. In fact, none of the delayed storks reached the traditional African wintering areas. Thus, our results show that juvenile storks can survive migrating at the 'wrong' time. However, this had long-term consequences on migration decisions. We suggest that, when timing their migration, storks balance not just energy and time, but also the availability of social information.

White stork movements reveal the ecological connectivity between landfills and different habitats.

BACKGROUND: Connections between habitats are key to a full understanding of anthropic impacts on ecosystems. Freshwater habitats are especially biodiverse, yet depend on exchange with terrestrial habitats. White storks (Ciconia ciconia) are widespread opportunists that often forage in landfills and then visit wetlands, among other habitats. It is well known that white storks ingest contaminants at landfills (such as plastics and antibiotic resistant bacteria), which can be then deposited in other habitats through their faeces and regurgitated pellets. METHODS: We characterized the role of white storks in habitat connectivity by analyzing GPS data from populations breeding in Germany and wintering from Spain to Morocco. We overlaid GPS tracks on a land-use surface to construct a spatially-explicit network in which nodes were sites, and links were direct flights. We then calculated centrality metrics, identified spatial modules, and quantified overall connections between habitat types. For regional networks in southern Spain and northern Morocco, we built Exponential Random Graph Models (ERGMs) to explain network topologies as a response to node habitat. RESULTS: For Spain and Morocco combined, we built a directed spatial network with 114 nodes and 370 valued links. Landfills were the habitat type most connected to others, as measured by direct flights. The relevance of landfills was confirmed in both ERGMs, with significant positive effects of this habitat as a source of flights. In the ERGM for southern Spain, we found significant positive effects of rice fields and salines (solar saltworks) as sinks for flights. By contrast, in the ERGM for northern Morocco, we found a significant positive effect of marshes as a sink for flights. CONCLUSIONS: These results illustrate how white storks connect landfills with terrestrial and aquatic habitats, some of which are managed for food production. We identified specific interconnected habitat patches across Spain and Morocco that could be used for further studies on biovectoring of pollutants, pathogens and other propagules.

Standardised empirical dispersal kernels emphasise the pervasiveness of long-distance dispersal in European birds.

Dispersal is a key life-history trait for most species and is essential to ensure connectivity and gene flow between populations and facilitate population viability in variable environments. Despite the increasing importance of range shifts due to global change, dispersal has proved difficult to quantify, limiting empirical understanding of this phenotypic trait and wider synthesis. Here, we introduce a statistical framework to estimate standardised dispersal kernels from biased data. Based on this, we compare empirical dispersal kernels for European breeding birds considering age (average dispersal; natal, before first breeding; and breeding dispersal, between subsequent breeding attempts) and sex (females and males) and test whether different dispersal properties are phylogenetically conserved. We standardised and analysed data from an extensive volunteer-based bird ring-recoveries database in Europe (EURING) by accounting for biases related to different censoring thresholds in reporting between countries and to migratory movements. Then, we fitted four widely used probability density functions in a Bayesian framework to compare and provide the best statistical descriptions of the different age and sex-specific dispersal kernels for each bird species. The dispersal movements of the 234 European bird species analysed were statistically best explained by heavy-tailed kernels, meaning that while most individuals disperse over short distances, long-distance dispersal is a prevalent phenomenon in almost all bird species. The phylogenetic signal in both median and long dispersal distances estimated from the best-fitted kernel was low (Pagel's λ < 0.25), while it reached high values (Pagel's λ >0.7) when comparing dispersal distance estimates for fat-tailed dispersal kernels. As expected in birds, natal dispersal was on average 5 km greater than breeding dispersal, but sex-biased dispersal was not detected. Our robust analytical framework allows sound use of widely available mark-recapture data in standardised dispersal estimates. We found strong evidence that long-distance dispersal is common among European breeding bird species and across life stages. The dispersal estimates offer a first guide to selecting appropriate dispersal kernels in range expansion studies and provide new avenues to improve our understanding of the mechanisms and rules underlying dispersal events.

La dispersión es un rasgo clave del ciclo vital de la mayoría de las especies y es esencial para garantizar la conectividad y el flujo genético entre poblaciones y contribuir a la viabilidad de la población en contextos de ambiente variable. A pesar de que la dispersión es clave para estudiar los cambios en el área de distribución de las especies debido al cambio global, la dispersión es difícil de cuantificar, lo que limita la comprensión empírica de este rasgo fenotípico y su síntesis más amplia. Aquí introducimos un marco de trabajo estadístico para estimar de manera estandarizada los kernels de dispersión a partir de datos sesgados. Basándonos en este marco, comparamos los kernels de dispersión empíricos para las aves reproductoras europeas considerando la edad (dispersión media vital; natal, antes de la primera reproducción; y dispersión reproductora, entre los intentos de reproducción posteriores) y el sexo (hembras y machos), además de explorar si las diferentes propiedades de dispersión se conservan filogenéticamente. Estandarizamos y analizamos los datos de una extensa base de datos de anillamiento de aves en Europa (EURING), basada en voluntarios, teniendo en cuenta los sesgos relacionados con los diferentes umbrales de comunicación de las anillas entre países y con los movimientos migratorios. A continuación, ajustamos, en un marco bayesiano, cuatro funciones de probabilidad ampliamente utilizadas para comparar y proporcionar las mejores descripciones estadísticas de los diferentes kernels de dispersión por edad y sexo para cada especie de ave. Los movimientos de dispersión de las 234 especies de aves europeas analizadas se explicaron estadísticamente mejor mediante kernels de cola pesada, lo que significa que, aunque la mayoría de los individuos se dispersan en distancias cortas, la dispersión a larga distancia es un fenómeno prevalente en casi todas las especies de aves. La señal filogenética tanto en las distancias de dispersión medias como en las largas estimadas a partir del kernel mejor ajustado fue baja (λ de Pagel < 0,25), mientras que alcanzó valores altos (λ de Pagel >0,7) al comparar las estimas de distancia de dispersión para los kernels de cola pesada. Como se esperaba en las aves, la dispersión natal fue en promedio 5 km mayor que la dispersión reproductiva, pero no se detectó una dispersión sesgada por sexo. Nuestro robusto marco analítico permite un buen uso de los datos de marcaje y recaptura disponibles para la estimación estandarizada de las distancias de dispersión. Hemos encontrado pruebas sólidas de que la dispersión a larga distancia es común entre las especies de aves reproductoras europeas y en todas las etapas de la vida. Las estimas de dispersión ofrecen un primer paso para seleccionar los kernels de dispersión adecuados para los estudios de expansión del rango de distribución y proporcionar nuevas vías de investigación para mejorar nuestra comprensión de los mecanismos y procesos que subyacen a los eventos de dispersión.

A multi-species evaluation of digital wildlife monitoring using the Sigfox IoT network.

Bio-telemetry from small tags attached to animals is one of the principal methods for studying the ecology and behaviour of wildlife. The field has constantly evolved over the last 80 years as technological improvement enabled a diversity of sensors to be integrated into the tags (e.g., GPS, accelerometers, etc.). However, retrieving data from tags on free-ranging animals remains a challenge since satellite and GSM networks are relatively expensive and or power hungry. Recently a new class of low-power communication networks have been developed and deployed worldwide to connect the internet of things (IoT). Here, we evaluated one of these, the Sigfox IoT network, for the potential as a real-time multi-sensor data retrieval and tag commanding system for studying fauna across a diversity of species and ecosystems. We tracked 312 individuals across 30 species (from 25 g bats to 3 t elephants) with seven different device concepts, resulting in more than 177,742 successful transmissions. We found a maximum line of sight communication distance of 280 km (on a flying cape vulture [Gyps coprotheres]), which sets a new documented record for animal-borne digital data transmission using terrestrial infrastructure. The average transmission success rate amounted to 68.3% (SD 22.1) on flying species and 54.1% (SD 27.4) on terrestrial species. In addition to GPS data, we also collected and transmitted data products from accelerometers, barometers, and thermometers. Further, we assessed the performance of Sigfox Atlas Native, a low-power method for positional estimates based on radio signal strengths and found a median accuracy of 12.89 km (MAD 5.17) on animals. We found that robust real-time communication (median message delay of 1.49 s), the extremely small size of the tags (starting at 1.28 g without GPS), and the low power demands (as low as 5.8 µAh per transmitted byte) unlock new possibilities for ecological data collection and global animal observation.

New frontiers in bird migration research.

Bird migrations are impressive behavioral phenomena, representing complex spatiotemporal strategies to balance costs of living while maximizing fitness. The field of bird migration research has made great strides over the past decades, yet fundamental gaps remain. Technologies have sparked a transformation in the study of bird migration research by revealing remarkable insights into the underlying behavioral, cognitive, physiological and evolutionary mechanisms of these diverse journeys. Here, we aim to encourage broad discussions and promote future studies by highlighting research fields that are characterized by major knowledge gaps or conflicting evidence, namely the fields of navigation, social learning, individual development, energetics and conservation. We approach each topic by summarizing the current state of knowledge and provide a future outlook of ideas and state-of-the-art methods to further advance the field. Integrating knowledge across these disciplines will allow us to understand the adaptive abilities of different species and to develop effective conservation strategies in a rapidly changing world.

Scatter-hoarding birds disperse seeds to sites unfavorable for plant regeneration.

Scatter-hoarding birds provide effective long-distance seed dispersal for plants. Transporting seeds far promotes population spread, colonization of new areas, and connectivity between populations. However, whether seeds transported over long distances are deposited in habitats favorable to plant regeneration has rarely been investigated, mainly due to methodological constraints. To investigate dispersal patterns and distances of Swiss stone pine (Pinus cembra) seeds we utilized advances in tracking technology to track the movements of their sole disperser, the spotted nutcracker (Nucifraga caryocatactes). We found routine individual movements between single seed harvesting and seed caching site. Harvesting sites of individual birds overlapped, whereas seed caching sites were separated and located on average 5.3 km away from the harvesting site. Interestingly, most distant caching sites were located at low elevations and in spruce forest, where Swiss stone pine does not naturally occur. This suggests that nutcrackers disperse seeds over long distances but that a large portion of these seeds are cached outside the known pine habitat. Therefore, we conclude that the implications of such long-distance seed dispersal movements for plant populations should be carefully considered in combination with the effects of habitat quality on plant recruitment.

Environmental and seasonal correlates of capercaillie movement traits in a Swedish wind farm.

Animals continuously interact with their environment through behavioral decisions, rendering the appropriate choice of movement speed and directionality an important phenotypic trait. Anthropogenic activities may alter animal behavior, including movement. A detailed understanding of movement decisions is therefore of great relevance for science and conservation alike. The study of movement decisions in relation to environmental and seasonal cues requires continuous observation of movement behavior, recently made possible by high-resolution telemetry. We studied movement traits of 13 capercaillie (Tetrao urogallus), a mainly ground-moving forest bird species of conservation interest, over two summer seasons in a Swedish windfarm using high-resolution GPS tracking data (5-min sampling interval). We filtered and removed unreliable movement steps using accelerometer data and step characteristics. We explored variation in movement speed and directionality in relation to environmental and seasonal covariates using generalized additive mixed models (GAMMs). We found evidence for clear daily and seasonal variation in speed and directionality of movement that reflected behavioral adjustments to biological and environmental seasonality. Capercaillie moved slower when more turbines were visible and faster close to turbine access roads. Movement speed and directionality were highest on open bogs, lowest on recent clear-cuts (<5 y.o.), and intermediate in all types of forest. Our results provide novel insights into the seasonal and environmental correlates of capercaillie movement patterns and supplement previous behavioral observations on lekking behavior and wind turbine avoidance with a more mechanistic understanding.

Smell of green leaf volatiles attracts white storks to freshly cut meadows.

Finding food is perhaps the most important task for all animals. Birds often show up unexpectedly at novel food sources such as freshly tilled fields or mown meadows. Here we test whether wild European white storks primarily use visual, social, auditory or olfactory information to find freshly cut farm pastures where insects and rodents abound. Aerial observations of an entire local stork population documented that birds could not have become aware of a mown field through auditory, visual or social information. Only birds within a 75° downwind cone over 0.4-16.6 km approached any mown field. Placing freshly cut grass from elsewhere on selected unmown fields elicited similarly immediate stork approaches. Furthermore, uncut fields that were sprayed with a green leaf volatile organic compound mix ((Z)-3-hexenal, (Z)-3-hexenol, hexenyl acetate), the smell of freshly cut grass, immediately attracted storks. The use of long-distance olfactory information for finding food may be common in birds, contrary to current perception.

Health monitoring in birds using bio-loggers and whole blood transcriptomics.

Monitoring and early detection of emerging infectious diseases in wild animals is of crucial global importance, yet reliable ways to measure immune status and responses are lacking for animals in the wild. Here we assess the usefulness of bio-loggers for detecting disease outbreaks in free-living birds and confirm detailed responses using leukocyte composition and large-scale transcriptomics. We simulated natural infections by viral and bacterial pathogens in captive mallards (Anas platyrhynchos), an important natural vector for avian influenza virus. We show that body temperature, heart rate and leukocyte composition change reliably during an acute phase immune response. Using genome-wide gene expression profiling of whole blood across time points we confirm that immunostimulants activate pathogen-specific gene regulatory networks. By reporting immune response related changes in physiological and behavioural traits that can be studied in free-ranging populations, we provide baseline information with importance to the global monitoring of zoonotic diseases.

Longer days enable higher diurnal activity for migratory birds.

Seasonal geophysical cycles strongly influence the activity of life on Earth because they affect environmental conditions like temperature, precipitation and day length. An increase in daylight availability during summer is especially enhanced when animals migrate along a latitudinal gradient. Yet, the question of how day length (i.e. daylight availability) influences the activity patterns of long-distance, latitudinal migrants is still unclear. Here, we ask whether migration provides benefits to long-distance migrants by enabling them to increase their diurnal movement activities due to an increase in daylight availability. To answer this question, we tested whether four vastly different species of long-distance migratory birds-two arctic migrants and two mid-latitude migrants-can capitalise on day length changes by adjusting their daily activity. We quantified the relationship between daily activity (measured using accelerometer data) and day length, and estimated each species' daily activity patterns. In addition, we evaluated the role of day length as an ultimate driver of bird migration. All four species exhibited longer activity periods during days with more daylight hours, showing a strong positive relationship between total daily activity and day length. The slope of this relationship varied between the different species, with activity increasing 1.5-fold on average when migrating from wintering to breeding grounds. Underlying mechanisms of these relationships reveal two distinct patterns of daily activity. Flying foragers showed increasing activity patterns, that is, their daytime activities rose uniformly up to solar noon and decreased until dusk, thereby exhibiting a season-specific activity slope. In contrast, ground foragers showed a constant activity pattern, whereby they immediately increased their activity to a certain level and maintained this level throughout the day. Our study reveals that long days allow birds to prolong their activity and increase their total daily activity. These findings highlight that daylight availability could be an additional ultimate cause of bird migration and act as a selective agent for the evolution of migration.

Los ciclos geofísicos estacionales influyen fuertemente la actividad de la vida en la Tierra ya que afectan diversas condiciones ambientales como la temperatura, la precipitación y la duración del día. El aumento de la disponibilidad de luz solar durante el verano favorece especialmente a las especies que migran a lo largo de un gradiente latitudinal. Sin embargo, el efecto de la duración del día (es decir, la disponibilidad de luz solar) en los patrones de actividad de las especies que migran latitudinalmente largas distancias, aún no es claro. Aquí nos preguntamos si un aumento de la disponibilidad de luz solar representa un beneficio para los animales que migran largas distancias, al prolongar el periodo de actividad diurna. Para responder a esta pregunta, investigamos si cuatro especies diferentes de aves migratorias de larga distancia, dos migrantes árticos y dos migrantes de latitudes medias, pueden ajustar su actividad diaria y aprovechar los cambios en la duración del día. Cuantificamos la relación entre la actividad diaria (medida con datos de acelerómetro) y la duración del día, y estimamos los patrones de actividad de cada especie. Además, evaluamos el papel de la duración del día como mecanismo último subyacente a la migración de las aves. Las cuatro especies mostraron períodos de actividad más largos durante los días con más horas de luz, mostrando una fuerte relación positiva entre la actividad diaria total y la duración del día. Esta relación varió entre las especies evaluadas. La actividad aumentó en promedio 1.5 veces durante la migración desde las áreas de invernada a las áreas de reproducción. Los mecanismos subyacentes a estas relaciones revelan dos patrones de actividad diaria. Las especies que forrajean en vuelo mostraron un aumento en su patrón de actividad. En este caso, la actividad diurna aumentó uniformemente hasta el mediodía y disminuyó hasta el atardecer, mostrando una pendiente de actividad específica para la estación. De otro lado, las especies que forrajean en tierra mostraron un patrón de actividad constante. Según este patrón, la actividad diurna aumenta hasta un determinado nivel, a partir del cual se mantiene durante el resto del día. Nuestro estudio revela que el aumento en la longitud del día le permite a las aves prolongar su actividad e incrementar su actividad diaria total. Estos resultados señalan que la disponibilidad de luz diurna podría ser otro mecanismo último subyacente a la migración de las aves y puede actuar como un factor de selección en la evolución de la migración.

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.

Overland and oversea migration of white storks through the water barriers of the straits of Gibraltar.

Soaring landbirds typically exploit atmospheric uplift as they fly overland, displaying a highly effective energy-saving locomotion. However, large water bodies lack thermal updrafts, potentially becoming ecological barriers that hamper migration. Here we assessed the effects of a sea surface on the migratory performance of GPS-tagged white storks (Ciconia ciconia) before, during and after they crossed the straits of Gibraltar. Oversea movements involved only flapping and gliding and were faster, traversed in straighter, descending trajectories and resulted in higher movement-related energy expenditure levels than overland, supporting the water barrier hypothesis. Overland movements at both sides of the sea straits resulted in tortuous routes and ascending trajectories with pre-crossing flights showing higher elevations and more tortuous routes than post-crossing, thus supporting the barrier negotiation hypothesis. Individual positions at both ends of the sea narrow were predicted by zonal winds and storks´ location at entry in the European hinterland, and birds did not show compensational movements overland in anticipation to subsequent wind displacements oversea. The length of the water narrow at departure shore, the elevation therein and the winds on route affected major components of sea crossing performance (such as distances and times overwater, minimum elevations, climb angles, speeds and energy expenditure), supporting the departure position and oversea winds hypotheses. In summary, our study provides a prime example at high temporal resolution of how birds adjust their behavior and physiology as they interact with the changing conditions of the travelling medium, reallocating resources and modifying their movement to overcome an ecological barrier.

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.

Daily energy expenditure in white storks is lower after fledging than in the nest.

Many juvenile birds turn into long-distance migrants within weeks of fledging. This transition involves upheavals in their energy management as major changes in growth and activity occur. Understanding such ontogenetic transitions in energy allocation has been difficult because collecting continuous data on energy costs in wild developing birds was previously largely impossible. Here, we continuously measured heart rate and fine-scale movements of 20 free-living juvenile white storks (Ciconia ciconia) using on-board bio-loggers to explore individual and environmental factors relating to daily mean heart rate. In addition, we explored which specific energy management strategy storks use during these crucial early life stages. We found that daily mean heart rate increased with overall movement activity, and increasing body temperature, but that it decreased with age. Further, we found that during the nestling period, when growth costs are high, activity costs are low, and post-fledging that activity costs are increased while maintenance costs are low, indicating a constraint on overall energy use in both phases. Our observations are consistent with the hypothesis that individuals invested more energy per unit time while still in the nest than after fledging despite the high costs of flight.

"Closer-to-home" strategy benefits juvenile survival in a long-distance migratory bird.

Human-induced changes in the climate and environment that occur at an unprecedented speed are challenging the existence of migratory species. Faced with these new challenges, species with diverse and flexible migratory behaviors may suffer less from population decline, as they may be better at responding to these changes by altering their migratory behavior. At the individual level, variations in migratory behavior may lead to differences in fitness and subsequently influence the population's demographic dynamics. Using lifetime GPS bio-logging data from 169 white storks (Ciconia ciconia), we explore whether the recently shortened migration distance of storks affects their survival during different stages of their juvenile life. We also explore how other variations in migratory decisions (i.e., time, destination), movement activity (measured using overall body dynamic acceleration), and early life conditions influence juvenile survival. We observed that their first autumn migration was the riskiest period for juvenile white storks. Individuals that migrated shorter distances and fledged earlier experienced lower mortality risks. In addition, higher movement activity and overwintering "closer-to-home" (with 84.21% of the tracked individuals stayed Europe or North Africa) were associated with higher survival. Our study shows how avian migrants can change life history decisions over only a few decades, and thus it helps us to understand and predict how migrants respond to the rapidly changing world.

Fly with the flock: immersive solutions for animal movement visualization and analytics.

Understanding the movement of animals is important for a wide range of scientific interests including migration, disease spread, collective movement behaviour and analysing motion in relation to dynamic changes of the environment such as wind and thermal lifts. Particularly, the three-dimensional (3D) spatial-temporal nature of bird movement data, which is widely available with high temporal and spatial resolution at large volumes, presents a natural option to explore the potential of immersive analytics (IA). We investigate the requirements and benefits of a wide range of immersive environments for explorative visualization and analytics of 3D movement data, in particular regarding design considerations for such 3D immersive environments, and present prototypes for IA solutions. Tailored to biologists studying bird movement data, the immersive solutions enable geo-locational time-series data to be investigated interactively, thus enabling experts to visually explore interesting angles of a flock and its behaviour in the context of the environment. The 3D virtual world presents the audience with engaging and interactive content, allowing users to 'fly with the flock', with the potential to ascertain an intuitive overview of often complex datasets, and to provide the opportunity thereby to formulate and at least qualitatively assess hypotheses. This work also contributes to ongoing research efforts to promote better understanding of bird migration and the associated environmental factors at the global scale, thereby providing a visual vehicle for driving public awareness of environmental issues and bird migration patterns.

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.

From local collective behavior to global migratory patterns in white storks.

Soaring migrant birds exploit columns of rising air (thermals) to cover large distances with minimal energy. Using social information while locating thermals may benefit such birds, but examining collective movements in wild migrants has been a major challenge for researchers. We investigated the group movements of a flock of 27 naturally migrating juvenile white storks by using high-resolution GPS and accelerometers. Analyzing individual and group movements on multiple scales revealed that a small number of leaders navigated to and explored thermals, whereas followers benefited from their movements. Despite this benefit, followers often left thermals earlier and at lower height, and consequently they had to flap considerably more. Followers also migrated less far annually than did leaders. We provide insights into the interactions between freely flying social migrants and the costs and benefits of collective movement in natural populations.