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.

Wind effects on the migration routes of trans-Saharan soaring raptors: geographical, seasonal, and interspecific variation.

Wind is among the most important environmental factors shaping birds' migration patterns. Birds must deal with the displacement caused by crosswinds and their behavior can vary according to different factors such as flight mode, migratory season, experience, and distance to goal areas. Here we analyze the relationship between wind and migratory movements of three raptor species which migrate by soaring-gliding flight: Egyptian vulture Neophron percnopterus, booted eagle Aquila pennata, and short-toed snake eagle Circaetus gallicus. We analyzed daily migratory segments (i.e., the path joining consecutive roosting locations) using data recorded by GPS satellite telemetry. Daily movements of Egyptian vultures and booted eagles were significantly affected by tailwinds during both autumn and spring migrations. In contrast, daily movements of short-toed eagles were only significantly affected by tailwinds during autumn migration. The effect of crosswinds was significant in all cases. Interestingly, Egyptian vultures and booted eagles showed latitudinal differences in their behavior: both species compensated more frequently at the onset of autumn migration and, at the end of the season when reaching their wintering areas, the proportion of drift segments was higher. In contrast, there was a higher drift at the onset of spring migration and a higher compensation at the end. Our results highlight the effect of wind patterns on the migratory routes of soaring raptors, with different outcomes in relation to species, season, and latitude, ultimately shaping the loop migration patterns that current tracking techniques are showing to be widespread in many long distance migrants.

Food predictability determines space use of endangered vultures: implications for management of supplementary feeding.

Understanding space use of free-living endangered animals is key to informing management decisions for conservation planning. Like most scavengers, vultures have evolved under a context of unpredictability of food resources (i.e., exploiting scattered carcasses that are intermittently available). However, the role of predictable sources of food in shaping spatial ecology of vultures has seldom been studied in detail. Here, we quantify the home range of the Egyptian Vulture (Neophron percnopterus), a long-lived raptor that has experienced severe population decline throughout its range and is qualified as endangered worldwide. To this end, six adults were tracked by satellite telemetry in Spain during the breeding season, from 2007 to 2012, recording 10360 GPS locations. Using Resource Utilization Functions, we assessed the topology of the Utilization Distribution, a three-dimensional measure that shows the probability of finding an animal within the home range. Our results showed how food availability, and principally, how food predictability, determines ranging behavior of this species. Egyptian Vultures showed consistent site fidelity across years, measured as the two- and three-dimensional overlap in their home ranges. Space use varied considerably within the home range and remarkably, places located far from nesting sites were used more frequently than some areas located closer. Therefore, traditional conservation measures based on establishing restrictive rules within a fixed radius around nesting sites could be biologically meaningless if other areas within the home range are not protected too. Finally, our results emphasize the importance of anthropogenic predictable sources of food (mainly vulture restaurants) in shaping the space use of scavengers, which is in agreement with recent findings. Hence, measures aimed at ensuring food availability are essential to preserve this endangered vulture, especially in the present context of limiting carrion dumping in the field due to sanitary regulations according to European legislation.

Scavengers on the move: behavioural changes in foraging search patterns during the annual cycle.

BACKGROUND: Optimal foraging theory predicts that animals will tend to maximize foraging success by optimizing search strategies. However, how organisms detect sparsely distributed food resources remains an open question. When targets are sparse and unpredictably distributed, a Lévy strategy should maximize foraging success. By contrast, when resources are abundant and regularly distributed, simple brownian random movement should be sufficient. Although very different groups of organisms exhibit Lévy motion, the shift from a Lévy to a brownian search strategy has been suggested to depend on internal and external factors such as sex, prey density, or environmental context. However, animal response at the individual level has received little attention. METHODOLOGY/PRINCIPAL FINDINGS: We used GPS satellite-telemetry data of Egyptian vultures Neophron percnopterus to examine movement patterns at the individual level during consecutive years, with particular interest in the variations in foraging search patterns during the different periods of the annual cycle (i.e. breeding vs. non-breeding). Our results show that vultures followed a brownian search strategy in their wintering sojourn in Africa, whereas they exhibited a more complex foraging search pattern at breeding grounds in Europe, including Lévy motion. Interestingly, our results showed that individuals shifted between search strategies within the same period of the annual cycle in successive years. CONCLUSIONS/SIGNIFICANCE: Results could be primarily explained by the different environmental conditions in which foraging activities occur. However, the high degree of behavioural flexibility exhibited during the breeding period in contrast to the non-breeding period is challenging, suggesting that not only environmental conditions explain individuals' behaviour but also individuals' cognitive abilities (e.g., memory effects) could play an important role. Our results support the growing awareness about the role of behavioural flexibility at the individual level, adding new empirical evidence about how animals in general, and particularly scavengers, solve the problem of efficiently finding food resources.

Interspecific comparison of the performance of soaring migrants in relation to morphology, meteorological conditions and migration strategies.

BACKGROUND: Performance of migrating birds can be affected by a number of intrinsic and extrinsic factors like morphology, meteorological conditions and migration strategies. We compared travel speeds of four raptor species during their crossing of the Sahara desert. Focusing the analyses on this region allows us to compare different species under equivalent conditions in order to disentangle which factors affect migratory performance. METHODOLOGY/PRINCIPAL FINDING: We tracked raptors using GPS satellite transmitters from Sweden, Spain and Italy, and evaluated their migratory performance at both an hourly and a daily scale. Hourly data (flight speed and altitude for intervals of two hours) were analyzed in relation to time of day, species and season, and daily data (distance between roosting sites) in relation to species, season, day length and tailwind support. CONCLUSIONS/SIGNIFICANCE: Despite a clear variation in morphology, interspecific differences were generally very small, and did only arise in spring, with long-distance migrants (>5000 km: osprey and Western marsh-harrier) being faster than species that migrate shorter distances (Egyptian vulture and short-toed eagle). Our results suggest that the most important factor explaining hourly variation in flight speed is time of day, while at a daily scale, tailwind support is the most important factor explaining variation in daily distance, raising new questions about the consequences of possible future changes in worldwide wind patterns.