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The northern wheatear Oenanthe oenanthe has an almost circumpolar breeding distribution in the Northern Hemisphere, but all populations migrate to sub‐Saharan Africa in winter. Currently, tracking data suggest two main access routes to the northern continents via the Middle East and the Iberian Peninsula. These routes would require detours for birds breeding in the European Alps. Our aim was to map the migration routes and determine annual schedules for birds breeding in Switzerland and Austria, using light level geolocators. We compared their migration patterns with birds from a lowland breeding population in Germany. Birds from the Alps cross the Mediterranean Sea directly heading straight to their non‐breeding sites. In contrast, birds from Germany travelled further west via the Iberian Peninsula. While the German population initiated autumn migration relatively early, arrival on the wintering sites was nearly synchronous across the three populations. During spring migration, German birds arrived earlier at their breeding grounds than birds from the Alps. A comparison with the literature indicated that the breeding populations in the Alps use their own route and are among the latest to arrive in spring, showing resemblance to the phenology of Arctic breeding populations. Our results indicate that the annual cycle of Alps‐breeding wheatears is influenced primarily by breeding ground conditions, and not solely by migration distance.

Organisms living in high‐elevation habitats are usually habitat specialists who occupy a narrow ecological niche. To envision the response of alpine species to a changing environment, it is fundamental to understand their habitat preferences on multiple spatial and temporal scales. However, information on small‐scale habitat use is still widely lacking. We investigated the foraging habitat preferences of the migratory northern wheatear Oenanthe oenanthe during the entire presence at a breeding site in the central Alps. We repeatedly observed 121 adult and juvenile individuals. We applied Bayesian logistic regression models to investigate which habitat characteristics influenced foraging habitat selection on a fine spatial scale, and how habitat use varied temporally. Throughout their presence on the breeding grounds, northern wheatears showed a consistent preference for a mosaic of stones and bare ground patches with slow‐growing, short vegetation. The proximity of marmot burrows was preferred, whereas dense and low woody vegetation was avoided. After arrival at the breeding site, short vegetation, preferably close to the snow, was favored. The preference for open habitat patches that provide access to prey underlines the critical role of small‐scale habitat heterogeneity for northern wheatears. The strong and consistent preference for a habitat that is under pressure from land‐use and climate change suggests that this alpine bird species may be sensitive to habitat loss, leading to a potential range contraction. We highlight the need to conserve habitat diversity on a small spatial scale to ensure the long‐term availability of suitable habitat for northern wheatears in the Alps. Repeated observations of northern wheatears (Oenanthe oenanthe) in their Alpine breeding range underline the importance of small‐scale habitat heterogeneity on the availability of suitable foraging habitat for this alpine passerine. The study revealed that a mosaic of short vegetation and bare ground patches within more productive areas are favored throughout the presence of the species at the breeding site. Conservation of structural heterogeneity that is affected by climate and land‐use change is crucial, as Alpine areas are becoming increasingly important for this habitat specialist.

Background To understand the ecology of long-distance migrant bird species, it is necessary to study their full annual cycle, including migratory routes and stopovers. This is especially important for species in high-elevation habitats that are particularly vulnerable to environmental change. Here, we investigated both local and global movements during all parts of the annual cycle in a small trans-Saharan migratory bird breeding at high elevation. Methods Recently, multi-sensor geolocators have opened new research opportunities in small-sized migratory organisms. We tagged Northern Wheatears Oenanthe oenanthe from the central-European Alpine population with loggers recording atmospheric pressure and light intensity. We modelled migration routes and identified stopover and non-breeding sites by correlating the atmospheric pressure measured on the birds with global atmospheric pressure data. Furthermore, we compared barrier-crossing flights with other migratory flights and studied the movement behaviour throughout the annual cycle. Results All eight tracked individuals crossed the Mediterranean Sea, using islands for short stops, and made longer stopovers in the Atlas highlands. Single non-breeding sites were used during the entire boreal winter and were all located in the same region of the Sahel. Spring migration was recorded for four individuals with similar or slightly different routes compared to autumn. Migratory flights were typically nocturnal and characterized by fluctuating altitudes, frequently reaching 2000 to 4000 m a.s.l, with a maximum of up to 5150 m. Barrier-crossing flights, i.e., over the sea and the Sahara, were longer, higher, and faster compared to flights above favourable stopover habitat. In addition, we detected two types of altitudinal movements at the breeding site. Unexpected regular diel uphill movements were undertaken from the breeding territories towards nearby roosting sites at cliffs, while regional scale movements took place in response to local meteorological conditions during the pre-breeding period. Conclusion Our data inform on both local and global scale movements, providing new insights into migratory behaviour and local movements in small songbirds. This calls for a wider use of multi-sensor loggers in songbird migration research, especially for investigating both local and global movements in the same individuals.

Sex-specific dispersal behavior has been documented in a wide range of different species. Avoidance of inbreeding and kin competition as well as different benefits of philopatry have been invoked as explanations for these patterns. All of these factors have, however, focused on explaining why dispersal behavior differs between the sexes. In this paper, we make the case that dispersal causes an increase in spatial variability in the sex ratio which can reduce the local availability of mates, and thus feed back to influence the evolution of sex-specific dispersal and lead to more, rather than less, similar dispersal behavior in the sexes. We investigate this mechanism in two different models, first in a conceptually simple case showing why the coevolutionary effect arises, second in an individual-based model where we model a population in explicit space with dispersal implemented as dispersal kernels. While our mechanism is not expected to completely remove sex-bias in dispersal, it can act alongside other selection pressures to reduce such biases. Our model thus shows that dispersal of one sex can have an effect on the selective pressures on the opposite sex, without implementing inbreeding avoidance or differential benefits or costs of dispersal.

Light‐level geolocator tags use ambient light recordings to estimate the whereabouts of an individual over the time it carried the device. Over the past decade, these tags have emerged as an important tool and have been used extensively for tracking animal migrations, most commonly small birds. Analysing geolocator data can be daunting to new and experienced scientists alike. Over the past decades, several methods with fundamental differences in the analytical approach have been developed to cope with the various caveats and the often complicated data. Here, we explain the concepts behind the analyses of geolocator data and provide a practical guide for the common steps encompassing most analyses – annotation of twilights, calibration, estimating and refining locations, and extraction of movement patterns – describing good practices and common pitfalls for each step. We discuss criteria for deciding whether or not geolocators can answer proposed research questions, provide guidance in choosing an appropriate analysis method and introduce key features of the newest open‐source analysis tools. We provide advice for how to interpret and report results, highlighting parameters that should be reported in publications and included in data archiving. Finally, we introduce a comprehensive supplementary online manual that applies the concepts to several datasets, demonstrates the use of open‐source analysis tools with step‐by‐step instructions and code and details our recommendations for interpreting, reporting and archiving. Over the past decade, light‐level geolocator tags have emerged as an important tool for tracking animals. However, due to various caveats, the translation of light recordings into location estimates can be daunting. In this 'How to…' paper, we provide a practical guide for the analysis of geolocator data by describing good practices and discussing common pitfalls.

Aim Knowledge of broad‐scale biogeographical patterns of animal migration is important for understanding ecological drivers of migratory behaviours. Here, we present a flyway‐scale assessment of the spatial structure and seasonal dynamics of the Afro‐Palaearctic bird migration system and explore how phenology of the environment guides long‐distance migration. Location Europe and Africa. Time period 2009–2017. Major taxa studied Birds. Methods We compiled an individual‐based dataset comprising 23 passerine and near‐passerine species of 55 European breeding populations, in which a total of 564 individuals were tracked during migration between Europe and sub‐Saharan Africa. In addition, we used remotely sensed primary productivity data (the normalized difference vegetation index) to estimate the timing of vegetation green‐up in spring and senescence in autumn across Europe. First, we described how individual breeding and non‐breeding sites and the migratory flyways link geographically. Second, we examined how the timing of migration along the two major Afro‐Palaearctic flyways is tuned with vegetation phenology at the breeding sites. Results We found the longitudes of individual breeding and non‐breeding sites to be related in a strongly positive manner, whereas the latitudes of breeding and non‐breeding sites were related negatively. In autumn, migration commenced ahead of vegetation senescence, and the timing of migration was 5–7 days earlier along the Western flyway compared with the Eastern flyway. In spring, the time of arrival at breeding sites was c. 1.5 days later for each degree northwards and 6–7 days later along the Eastern compared with the Western flyway, reflecting the later spring green‐up at higher latitudes and more eastern longitudes. Main conclusions Migration of the Afro‐Palaearctic landbirds follows a longitudinally parallel leapfrog migration pattern, whereby migrants track vegetation green‐up in spring but depart before vegetation senescence in autumn. The degree of continentality along migration routes and at the breeding sites of the birds influences the timing of migration on a broad scale.

Swifts (Apodidae) are an unusual group of birds that spend most of their lives in flight, landing only when breeding. Although this aerial lifestyle greatly reduces their likelihood of being bitten by vectors and infected by vector-born parasites, swifts can still be heavily infested during breeding by nest-based vectors such as louse flies (Hippoboscidae). Here, we investigated host, vector, and vector-borne parasite relationships in the three most widespread swift species in the Western Palearctic (WP): common swifts ( Apus apus ), pallid swifts ( A. pallidus ), and alpine swifts ( Tachymarptis melba ), their nest-based louse flies ( Crataerina pallida and C. melbae ) and avian haemosporidians (genera Haemoproteus , Plasmodium , and Leucocytozoon ). Studies of haemosporidian infections in Apodidae remain limited, with clear evidence of infection found to date in just four Neotropical and one Australasian species. The possible role of louse flies in transmitting haemosporidian infections has never been tested in swifts. We assessed the occurrence of haemosporidian infection by PCR screenings of DNA from blood samples from 34 common swifts and 44 pallid swifts from Italy, and 45 alpine swifts from Switzerland. We also screened 20 ectoparasitic louse flies present on 20 birds and identified them by both morphological features and cytochrome oxidase subunit 1 (COI) barcodes. Our results provide no evidence of haemosporidian infection in the 123 swifts tested or in the two louse fly species we identified. Our findings are consistent with available knowledge showing no haemosporidian occurrence in WP swift species and that the most likely infection route for these highly aerial species (via louse fly ectoparasites during nesting) is unlikely.

Studying individual flight behaviour throughout the year is indispensable to understand the ecology of a bird species. Recent development in technology allows now to track flight behaviour of small long-distance bird migrants throughout its annual cycle. The specific flight behaviour of twilight ascents in birds has been documented in a few studies, but only during a short period of the year, and never quantified on the individual level. It has been suggested that twilight ascents might be a role in orientation and navigation. Previous studies had reported the behaviour only near the breeding site and during migration. We investigated yearround flight behaviour of 34 individual Alpine swifts (Apus melba) of four different populations in relation to twilight ascents. We recorded twilight ascents all around the year and found a twofold higher frequency in ascents during the non-breeding residence phase in Africa compared to all other phases of the year. Dawn ascents were twice as common as dusk ascents and occurred mainly when atmospheric conditions remained stable over a 24-h period. We found no conclusive support that twilight ascents are essential for recalibration of compass cues and landmarks. Data on the wing flapping intensity revealed that high activity at twilight occurred more regularly than the ascents. We therefore conclude that alpine swift generally increase flight activity—also horizontal flight—during the twilight period and we suppose that this increased flight activity, including ascents, might be part of social interactions between individuals.

In many taxa, the most common form of sex-biased migration timing is protandry—the earlier arrival of males at breeding areas. Here we test this concept across the annual cycle of long-distance migratory birds. Using more than 350 migration tracks of small-bodied trans-Saharan migrants, we quantify differences in male and female migration schedules and test for proximate determinants of sex-specific timing. In autumn, males started migration about 2 days earlier, but this difference did not carry over to arrival at the non-breeding sites. In spring, males on average departed from the African non-breeding sites about 3 days earlier and reached breeding sites ca 4 days ahead of females. A cross-species comparison revealed large variation in the level of protandry and protogyny across the annual cycle. While we found tight links between individual timing of departure and arrival within each migration season, only for males the timing of spring migration was linked to the timing of previous autumn migration. In conclusion, our results demonstrate that protandry is not exclusively a reproductive strategy but rather occurs year-round and the two main proximate determinants for the magnitude of sex-biased arrival times in autumn and spring are sex-specific differences in departure timing and migration duration.

Currently, the deployment of tracking devices is one of the most frequently used approaches to study movement ecology of birds. Recent miniaturization of light‐level geolocators enabled studying small bird species whose migratory patterns were widely unknown. However, geolocators may reduce vital rates in tagged birds and may bias obtained movement data. There is a need for a thorough assessment of the potential tag effects on small birds, as previous meta‐analyses did not evaluate unpublished data and impact of multiple life‐history traits, focused mainly on large species and the number of published studies tagging small birds has increased substantially. We quantitatively reviewed 549 records extracted from 74 published and 48 unpublished studies on over 7,800 tagged and 17,800 control individuals to examine the effects of geolocator tagging on small bird species (body mass <100 g). We calculated the effect of tagging on apparent survival, condition, phenology and breeding performance and identified the most important predictors of the magnitude of effect sizes. Even though the effects were not statistically significant in phylogenetically controlled models, we found a weak negative impact of geolocators on apparent survival. The negative effect on apparent survival was stronger with increasing relative load of the device and with geolocators attached using elastic harnesses. Moreover, tagging effects were stronger in smaller species. In conclusion, we found a weak effect on apparent survival of tagged birds and managed to pinpoint key aspects and drivers of tagging effects. We provide recommendations for establishing matched control group for proper effect size assessment in future studies and outline various aspects of tagging that need further investigation. Finally, our results encourage further use of geolocators on small bird species but the ethical aspects and scientific benefits should always be considered. Tagging slightly reduces only apparent survival of treated birds. The authors found stronger tagging effects when relatively heavier tags were used and no differences between published and unpublished studies. Finally, they call for the control group establishment in all future studies and provide guidelines for the selection of control individuals.