Behavioral plasticity shapes population aging patterns in a long-lived avian scavenger.

Studying the mechanisms shaping age-related changes in behavior ("behavioral aging") is important for understanding population dynamics in our changing world. Yet, studies that capture within-individual behavioral changes in wild populations of long-lived animals are still scarce. Here, we used a 15-y GPS-tracking dataset of a social obligate scavenger, the griffon vulture (Gyps fulvus), to investigate age-related changes in movement and social behaviors, and disentangle the role of behavioral plasticity and selective disappearance in shaping such patterns. We tracked 142 individuals for up to 12 y and found a nonlinear increase in site fidelity with age: a sharp increase in site fidelity before sexual maturity (<5 y old), stabilization during adulthood (6 to 15 y), and a further increase at old age (>15 y). This pattern resulted from individuals changing behavior throughout their life (behavioral plasticity) and not from selective disappearance. Mature vultures increased the predictability of their movement routines and spent more nights at the most popular roosting sites compared to younger individuals. Thus, adults likely have a competitive advantage over younger conspecifics. These changes in site fidelity and movement routines were mirrored in changes to social behavior. Older individuals interacted less with their associates (decreasing average strength with age), particularly during the breeding season. Our results reveal a variety of behavioral aging patterns in long-lived species and underscore the importance of behavioral plasticity in shaping such patterns. Comprehensive longitudinal studies are imperative for understanding how plasticity and selection shape the persistence of wild animal populations facing human-induced environmental changes.

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.

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 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.

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.

Linking migration and microbiota at a major stopover site in a long-distance avian migrant.

Migration is one of the most physical and energetically demanding periods in an individual bird's life. The composition of the bird's gut or cloacal microbiota can temporarily change during migration, likely due to differences in diets, habitats and other environmental conditions experienced en route. However, how physiological condition, migratory patterns, and other drivers interact to affect microbiota composition of migratory birds is still unclear. We sampled the cloacal bacterial microbiota of a long-distance migrant, the steppe buzzard (Buteo buteo vulpinus), at an important spring stopover bottleneck in Eilat, Israel, after crossing the ca. 1800 km Sahara Desert. We examined whether diversity and composition of the cloacal microbiota varied with body condition, sex, movement patterns (i.e., arrival time and migration distance), and survival. Early arrival to Eilat was associated with better body condition, longer post-Eilat spring migration distance, higher microbial α-diversity, and differences in microbiota composition. Specifically, early arrivals had higher abundance of the phylum Synergistota and five genera, including Jonquetella and Peptococcus, whereas the phylum Proteobacteria and genus Escherichia-Shigella (as well as three other genera) were more abundant in later arrivals. While the differences in α-diversity and Escherichia-Shigella seem to be mainly driven by body condition, other compositional differences associated with arrival date could be indicators of longer migratory journeys (e.g., pre-fueling at wintering grounds or stopover habitats along the way) or migratory performance. No significant differences were found between the microbiota of surviving and non-surviving individuals. Overall, our results indicate that variation in steppe buzzard microbiota is linked to variation in migratory patterns (i.e., capture/arrival date) and body condition, highlighting the importance of sampling the microbiota of GPS-tracked individuals on multiple occasions along their migration routes to gain a more detailed understanding of the links between migration, microbiota, and health in birds.

Unravelling migration connectivity reveals unsustainable hunting of the declining ortolan bunting.

In France, illegal hunting of the endangered ortolan bunting Emberiza hortulana has been defended for the sake of tradition and gastronomy. Hunters argued that ortolan buntings trapped in southwest France originate from large and stable populations across the whole of Europe. Yet, the European Commission referred France to the Court of Justice of the European Union (EU) in December 2016 for infringements to legislation (IP/16/4213). To better assess the impact of hunting in France, we combined Pan-European data from archival light loggers, stable isotopes, and genetics to determine the migration strategy of the species across continents. Ortolan buntings migrating through France come from northern and western populations, which are small, fragmented and declining. Population viability modeling further revealed that harvesting in southwest France is far from sustainable and increases extinction risk. These results provide the sufficient scientific evidence for justifying the ban on ortolan harvesting in France.