The association between haemosporidian infection and non-breeding moult location in great reed warblers revisited by combining feather stable isotope profiles and geolocator data.

Stable isotope analysis provides valuable insights into the ecology of long-distance migratory birds during periods spent away from a specific study site. In a previous study, Swedish great reed warblers (Acrocephalus arundinaceus) infected with haemosporidian parasites differed in feather isotope ratios compared to non-infected birds, suggesting that infected and non-infected birds spent the non-breeding season in different locations or habitats. Here, we use a novel dataset comprising geolocator data, isotopes, and haemosporidian infection status of 92 individuals from four Eurasian populations to investigate whether parasite transmission varies with geography or habitats. We found that the probability of harbouring Plasmodium and Leucocytozoon parasites was higher in birds moulting in the eastern region of the non-breeding grounds. However, no geographic pattern occurred for Haemoproteus infections or overall infection status. In contrast to the previous study, we did not find any relationship between feather isotope ratios and overall haemosporidian infection for the entire current dataset. Plasmodium-infected birds had lower feather δ15N values indicating that they occupied more mesic habitats. Leucocytozoon-infected birds had higher feather δ34S values suggesting more coastal sites or wetlands with anoxic sulphate reduction. As the composition and prevalence of haemosporidian parasites differed between the old and the current dataset, we suggest that the differences might be a consequence of temporal dynamics of haemosporidian parasites. Our results emphasize the importance of replicating studies conducted on a single population over a restricted time period, as the patterns can become more complex for data from wider geographical areas and different time periods.

Weak effects of geolocators on small birds: A meta-analysis controlled for phylogeny and publication bias.

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.

Arthropod biomass increase in spring correlates with NDVI in grassland habitat.

Data from remote sensing are often used as proxies to quantify biological processes, especially at large geographical scales. The normalized difference vegetation index (NDVI) is the most frequently used proxy for primary productivity. Assuming a direct, positive interrelation between primary and secondary production in terrestrial habitats, NDVI is often used to predict food availability for higher trophic levels. However, the relationship between NDVI and arthropod biomass has rarely been tested. In this study, we analyzed extensive datasets of arthropod communities from semi-natural grasslands in central Europe to test the relationship between arthropod biomass of consumer trophic levels ("herbivores," "mixed," and "carnivores") in grassland communities and NDVI during the spring season. We found that arthropod biomass generally increased with NDVI. The same positive relationship between biomass and NDVI was observed for each individual trophic level. Cross-correlation analyses did not show statistically significant lags between the NDVI and biomass of herbivores and carnivores. All in all, our study provides correlational evidence for the positive relation of primary and secondary productivity in temperate terrestrial habitats during spring. Moreover, it supports the applicability of NDVI data as a suitable habitat-specific proxy for the food availability of insectivores during spring.

A full annual perspective on sex-biased migration timing in long-distance migratory birds.

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.

Low intensity blood parasite infections do not reduce the aerobic performance of migratory birds.

Blood parasites (Haemosporidia) are thought to impair the flight performance of infected animals, and therefore, infected birds are expected to differ from their non-infected counterparts in migratory capacity. Since haemosporidians invade host erythrocytes, it is commonly assumed that infected individuals will have compromised aerobic capacity, but this has not been examined in free-living birds. We tested if haemosporidian infections affect aerobic performance by examining metabolic rates and exercise endurance in migratory great reed warblers (Acrocephalus arundinaceus) experimentally treated with Plasmodium relictum pGRW04 and in naturally infected wild birds over consecutive life-history stages. We found no effect of acute or chronic infections on resting metabolic rate, maximum metabolic rate or exercise endurance in either experimentally treated or free-living birds. Oxygen consumption rates during rest and while undergoing maximum exercise as well as exercise endurance increased from breeding to migration stages in both infected and non-infected birds. Importantly, phenotypic changes associated with preparation for migration were similarly unaffected by parasitaemia. Consequently, migratory birds experiencing parasitaemia levels typical of chronic infection do not differ in migratory capacity from their uninfected counterparts. Thus, if infected hosts differ from uninfected conspecifics in migration phenology, other mechanisms besides aerobic capacity should be considered.

Host migration strategy and blood parasite infections of three sparrow species sympatrically breeding in Southeast Europe.

Mobile hosts like birds occupy a wide array of habitats in which they encounter various vector and parasite faunas. If the infection probability for vector-borne parasites varies among seasons and biomes, a migratory life can critically influence the infections of a host. The growing body of literature on avian blood parasites suggests that host migrations do not only influence prevalence of infection but can also evoke higher infection intensities and increased parasite diversity in migrant compared to resident host species. We investigated the prevalence, intensity and diversity of Plasmodium and Haemoproteus infections in three closely-related and sympatrically breeding sparrow species with different migration strategies ranging from residential house sparrow and partially migratory tree sparrow to the obligate migratory Spanish sparrow. With a prevalence of 49%, the migratory Spanish sparrows were significantly less frequently infected than the resident house sparrows (82%). The partially migratory tree sparrow showed an intermediate prevalence of 60%. The parasitaemias were similar in all three host species and indicated mostly chronic but also few acute infections. While we found Plasmodium parasites in all three sparrow species, only Spanish sparrows were infected with Haemoproteus parasites in our study. With nine clearly identified parasite lineages in our study and the highest number of lineages per infected individuals (i.e. relative diversity), Spanish sparrows harboured the most diverse parasite fauna. Our results suggest that migration strategies can affect Plasmodium and Haemoproteus infections of sparrows resulting in a lower parasite prevalence and higher parasite diversity in migratory hosts-at least during our host's breeding period. A general scope for all annual cycle periods and across various bird taxa remains to be elucidated.

Individual migration timing of common nightingales is tuned with vegetation and prey phenology at breeding sites.

BACKGROUND: The timing of migration substantially influences individual fitness. To match peak requirements with peak resource availability, we hypothesized that individual migrants schedule spring migration in close relation to seasonal changes in environmental conditions along the route and particularly, at the breeding destination.To test this hypothesis, we investigated the timing of spring migration in male common nightingales Luscinia megarhynchos, a small Palearctic-African long-distance migrant, by linking spring migration timing to the phenology of local environmental conditions at non-breeding migratory stopover and breeding sites. In particular, we related individual migration decisions (i.e. departure and arrival) of nine males to site-specific vegetation phenology (based on remotely sensed vegetation index) and a proxy of food availability (based on insects' thermal requirements). RESULTS: We found weak relation of departures from non-breeding and no relation of stopover timing with local phenology. However, our results showed that individuals, which departed early from their non-breeding sites and arrived early at the breeding site closely matched spring green-up there. Early arrival at the breeding site meant also a close match with peak food availability for adults and in a time-lagged manner, for offspring. CONCLUSION: Our findings suggest that male nightingale used cues other than local phenology for their departure decisions from non-breeding grounds and that there is some evidence for equalizing late departures during the course of migration.

Avian Haemosporidians Infecting Short- and Long-Distance Migratory Old World Flycatcher Species and the Variation in Parasitaemia After Endurance Flights.

PURPOSE: Avian haemosporidians are widespread parasites, occurring in many bird families and causing pathologies ranging from rather benign infections to highly virulent diseases. The state of knowledge about lineage-specific intensities of haemosporidian infection (i.e., parasitaemia) is mainly based on infection experiments conducted under laboratory conditions. The levels and range of parasitaemia in natural host-parasite associations as well as their influencing factor remain largely unexplored. METHODS: Thus, we explored the parasitaemia of four songbird species (i.e., European Robins, Black and Common Redstarts and Whinchats) during migration by screening individuals upon landing on an insular passage site after extensive endurance flights to (1) describe their natural host-parasite associations, (2) quantify parasitaemia and (3) explore potential host- and parasite-related factors influencing parasitaemia. RESULTS: We found 68% of Whinchats to be infected with haemosporidians, which is more frequent than any other of the studied host species (30-34%). Furthermore, we confirmed that parasitaemia of Haemoproteus infections was higher than average Plasmodium infections. Median parasitaemia levels were rather low (parasite cells in 0.01% of hosts' red blood cells) and varied largely among the different parasite lineages. However, we found four individuals hosting infections with parasitaemia higher than typical chronic infections. CONCLUSIONS: Based on the known transmission areas of the respective lineages, we argue that these higher intensity infections might be relapses of consisting infections rather than acute phases of recent primary infections.

Short- and long-distance avian migrants differ in exercise endurance but not aerobic capacity.

BACKGROUND: Migratory birds differ markedly in their migration strategies, particularly those performing short- versus long-distance migrations. In preparation for migration, all birds undergo physiological and morphological modifications including enlargement of fat stores and pectoral muscles to fuel and power their flights, as well as cardiovascular and biochemical adjustments that improve lipid and oxygen delivery and uptake by flight muscles. While the magnitude of these changes varies in relation to migration strategy, the consequence of these variations on aerobic performance is unknown. We tested whether the aerobic performance of four Old-world flycatcher species (Muscicapidae) varied according to migration strategy by comparing minimum resting metabolic rates (RMRmin), exercise-induced maximum metabolic rates (MMR), and exercise endurance times of short-distance and long-distance migratory birds. RESULTS: As expected, RMRmin did not vary between short-distance and long-distance migrants but differed between the species within a migration strategy and between sexes. Unexpectedly, MMR did not vary with migration strategy, but MMR and blood haemoglobin content were positively related among the birds tested. Exercise endurance times differed substantially between migration strategies with long-distance migrants sustaining exercise for > 60% longer than short-distance migrants. Blood haemoglobin content had a significant positive effect on endurance among all birds examined. CONCLUSIONS: The lack of difference in RMRmin and MMR between long- and short-distance migrants during this stage of migration suggests that the attributes favouring the greater aerobic endurance of long-distance migrants did not come at the expense of increased maintenance costs or require greater aerobic capacity.

Effects of blood parasite infections on spatiotemporal migration patterns and activity budgets in a long-distance migratory passerine.

How blood parasite infections influence the migration of hosts remains a lively debated issue as past studies found negative, positive, or no response to infections. This particularly applies to small birds, for which monitoring of detailed migration behavior over a whole annual cycle has been technically unachievable so far. Here, we investigate how bird migration is influenced by parasite infections. To this end, we tracked great reed warblers (Acrocephalus arundinaceus) with multisensor loggers, characterized general migration patterns as well as detailed flight bout durations, resting times and flight heights, and related these to the genus and intensity of their avian haemosporidian infections. We found migration distances to be shorter and the onset of autumn migration to be delayed with increasing intensity of blood parasite infection, in particular for birds with Plasmodium and mixed-genus infections. Additionally, the durations of migratory flight bout were prolonged for infected compared to uninfected birds. But since severely infected birds and particularly birds with mixed-genus infections had shorter resting times, initial delays seemed to be compensated for and the timing in other periods of the annual cycle was not compromised by infection. Overall, our multisensor logger approach revealed that avian blood parasites have mostly subtle effects on migratory performance and that effects can occur in specific periods of the year only.

Spatially different annual cycles but similar haemosporidian infections in distant populations of collared sand martins.

BACKGROUND: Populations of long-distance migratory birds experience different environments and are consequently exposed to different parasites throughout their annual cycles. Though, specific whereabouts and accompanied host-parasite interactions remain unknown for most migratory passerines. Collared sand martins (Riparia riparia) breeding in the western Palaearctic spend the nonbreeding period in Africa, but it is not yet clear whether specific populations differ in overwintering locations and whether these also result in varying infections with vector-transmitted endoparasites. RESULTS: Geolocator tracking revealed that collared sand martins from northern-central and central-eastern Europe migrate to distant nonbreeding sites in West Africa and the Lake Chad basin in central Africa, respectively. While the ranges of these populations were clearly separated throughout the year, they consistently spent up to 60% of the annual cycle in Africa. Ambient light recorded by geolocators further indicated unsheltered roosting during the nonbreeding season in Africa compared to the breeding season in Europe. We found 5-26% prevalence of haemosporidian parasites in three breeding populations and one migratory passage population that was only sampled but not tracked. In total, we identified seven Plasmodium and nine Haemoproteus lineages (incl. two and seven new lineages, respectively), the latter presumably typical for swallows (Hirundinae) hosts. 99.5% of infections had a low intensity, typical for chronic infection stages, whereas three individuals (0.5%) showed high parasitaemia typical for acute infections during spring migration and breeding. CONCLUSIONS: Our study shows that blood parasite infections are common in several western Palaearctic breeding populations of collared sand martins who spent the nonbreeding season in West Africa and the lake Chad region. Due to long residency at the nonbreeding grounds blood parasite transmissions may mainly occur at host population-specific residences sites in Europe and Africa; the latter being likely facilitated by unsheltered roosting and thus high vulnerability to hematophagous insects. The rare cases of high parasitaemia during spring migration and breeding further indicates either relapses of chronic infection or primary infections which occurred shortly before migration and during breeding.

Population- and age-specific patterns of haemosporidian assemblages and infection levels in European bee-eaters (Merops apiaster).

Amongst other factors, host behaviour critically determines the patterns with which blood parasites occur in wild host populations. In particular, migratory hosts that sequentially occupy distant sites within and across years are expected to show distinct patterns of blood parasitism depending on their population-specific schedules and whereabouts. Here, we monitored haemosporidian parasitism in two populations of European bee-eaters (Merops apiaster), breeding in Portugal and Germany, with fundamentally different spatiotemporal migration patterns and colonisation histories. We describe and compare the composition of their parasite fauna as well as host population-, age- and sex-specific patterns in the frequency and intensity of infections. We found haemosporidian prevalence to be higher in Portugal compared with Germany and the prevalence generally increased with host age in both populations. Bee-eaters breeding in Portugal and wintering in western Africa mostly hosted parasites of the genus Haemoproteus, while Plasmodium lineages prevailed in birds breeding in Germany and wintering in central Africa. We found 18 genetic lineages, of which nine uniquely occurred in Germany, three uniquely in Portugal and six occurred in both breeding populations. The infection intensities (= % infected per inspected erythrocytes) ranged from 0.002% up to maximally 2.5% in Portugal and 9.6% in Germany. The intensity was higher in Germany compared with Portugal, vastly varied between the parasite genera (Haemoproteus > Plasmodium), but also differed between lineages of the same genus. Our results suggest that populations from different parts of a host's breeding range differ in prevalence and the composition of their haemosporidian assemblages, rather than in the intensity of their infections. Whether these patterns are mainly caused by differential habitat use throughout the annual cycle and/or the population-specific co-evolutionary backgrounds of a host species in range expansion remains to be elucidated.

Moult-related reduction of aerobic scope in passerine birds.

It is well established that the nutrient and energy requirements of birds increase substantially during moult, but it is not known if these increased demands affect their aerobic capacity. We quantified the absolute aerobic scope of house and Spanish sparrows, Passer domesticus and P. hispaniolensis, respectively, before and during sequential stages of their moult period. The absolute aerobic scope (AAS) is the difference between maximum metabolic rate (MMR) during peak locomotor activities and minimum resting metabolic rate (RMRmin), thus representing the amount of aerobic power above that committed to maintenance needs available for other activities. As expected, RMRmin increased over the moult period by up to 40 and 63% in house and Spanish sparrows, respectively. Surprisingly, the maximum metabolic rates also decreased during moult in both species, declining as much as 25 and 38% compared with pre-moult values of house and Spanish sparrows, respectively. The concurrent changes in RMRmin and MMR during moult resulted in significant decreases in AAS, being up to 32 and 47% lower than pre-moult levels of house and Spanish sparrows, respectively, during moult stages having substantial feather replacement. We argue that the combination of reduced flight efficiency due to loss of wing feathers and reduced aerobic capacity places moulting birds at greater risk of predation. Such performance constraints likely contribute to most birds temporally separating moult from annual events requiring peak physiological capacity such as breeding and migration.

Intra-tropical movements as a beneficial strategy for Palearctic migratory birds.

Migratory birds often move significantly within their non-breeding range before returning to breed. It remains unresolved under which circumstances individuals relocate, whether movement patterns are consistent between populations and to what degree the individuals benefit from the intra-tropical movement (ITM). We tracked adult great reed warblers Acrocephalus arundinaceus from a central and a southeastern European breeding population, which either stay at a single non-breeding site, or show ITM, i.e. move to a second site. We related ITM to the normalized difference vegetation index (NDVI) describing vegetation conditions and probably reflecting food abundance for these insectivorous birds. Three-quarters of birds showed ITM across the non-breeding range. We found no difference in range values and mean values of NDVI between the single non-breeding sites of stationary birds and the two sites of moving birds. The vegetation conditions were better at the second sites compared to the first sites during the period which moving birds spent at the first sites. Vegetation conditions further deteriorated at the first sites during the period the moving birds resided at their second sites. Our study provides evidence that birds probably benefit from improved conditions after ITM compared to the conditions at the sites from where they departed.

Miniaturized multi-sensor loggers provide new insight into year-round flight behaviour of small trans-Sahara avian migrants.

BACKGROUND: Over the past decade, the miniaturisation of animal borne tags such as geolocators and GPS-transmitters has revolutionized our knowledge of the whereabouts of migratory species. Novel light-weight multi-sensor loggers (1.4 g), which harbour sensors for measuring ambient light intensity, atmospheric pressure, temperature and acceleration, were fixed to two long-distance migrant bird species - eurasian hoopoe (Upupa epops) and great reed warbler (Acrocephalus arundinaceus). Using acceleration and atmospheric pressure data recorded every 5 and 30 min, respectively, we aimed at reconstructing individual diurnal and seasonal patterns of flight activity and flight altitude and thereby, at describing basic, yet hitherto unknown characteristics of migratory flight behaviour. Furthermore, we wanted to characterise the variability in these migration characteristics between individuals, species and migration periods. RESULTS: The flight duration from breeding to sub-Saharan African non-breeding sites and back was more variable within than between the species. Great reed warblers were airborne for a total of 252 flight hours and thus, only slightly longer than eurasian hoopoes with 232 h. With a few exceptions, both species migrated predominantly nocturnally - departure around dusk and landing before dawn. Mean flight altitudes were higher during pre- than during post-breeding migration (median 1100 to 1600 m a.s.l.) and flight above 3000 m occurred regularly with a few great reed warblers exceeding 6000 m a.s.l. (max. 6458 m a.s.l.). Individuals changed flight altitudes repeatedly during a flight bout, indicating a continuous search for (more) favourable flight conditions. CONCLUSIONS: We found high variation between individuals in the flight behaviour parameters measured - a variation that surprisingly even exceeded the variation between the species. More importantly, our results have shown that multi-sensor loggers have the potential to provide detailed insights into many fundamental aspects of individual behaviour in small aerial migrants. Combining the data recorded on the multiple sensors with, e.g., remote sensing data like weather and habitat quality on the spatial and temporal scale will be a great step forward to explore individual decisions during migration and their consequences.

Spatiotemporal Group Dynamics in a Long-Distance Migratory Bird.

Thousands of species migrate [1]. Though we have some understanding of where and when they travel, we still have very little insight into who migrates with whom and for how long. Group formation is pivotal in allowing individuals to interact, transfer information, and adapt to changing conditions [2]. Yet it is remarkably difficult to infer group membership in migrating animals without being able to directly observe them. Here, we use novel lightweight atmospheric pressure loggers to monitor group dynamics in a small migratory bird, the European bee-eater (Merops apiaster). We present the first evidence of a migratory bird flying together with non-kin of different ages and sexes at all stages of the life cycle. In fact, 49% stay together throughout the annual cycle, never separating longer than 5 days at a time despite the ∼14,000-km journey. Of those that separated for longer, 89% reunited within less than a month with individuals they had previously spent time with, having flown up to 5,000 km apart. These birds were not only using the same non-breeding sites, but also displayed coordinated foraging behaviors-these are unlikely to result from chance encounters in response to the same environmental conditions alone. Better understanding of migratory group dynamics, using the presented methods, could help improve our understanding of collective decision making during large-scale movements.

Barrier crossing in small avian migrants: individual tracking reveals prolonged nocturnal flights into the day as a common migratory strategy.

Over decades it has been unclear how individual migratory songbirds cross large ecological barriers such as seas or deserts. By deploying light-level geolocators on four songbird species weighing only about 12 g, we found that these otherwise mainly nocturnal migrants seem to regularly extend their nocturnal flights into the day when crossing the Sahara Desert and the Mediterranean Sea. The proportion of the proposed diurnally flying birds gradually declined over the day with similar landing patterns in autumn and spring. The prolonged flights were slightly more frequent in spring than in autumn, suggesting tighter migratory schedules when returning to breeding sites. Often we found several patterns for barrier crossing for the same individual in autumn compared to the spring journey. As only a small proportion of the birds flew strictly during the night and even some individuals might have flown non-stop, we suggest that prolonged endurance flights are not an exception even in small migratory species. We emphasise an individual's ability to perform both diurnal and nocturnal migration when facing the challenge of crossing a large ecological barrier to successfully complete a migratory journey.

Longer wings for faster springs - wing length relates to spring phenology in a long-distance migrant across its range.

In migratory birds, morphological adaptations for efficient migratory flight often oppose morphological adaptations for efficient behavior during resident periods. This includes adaptations in wing shape for either flying long distances or foraging in the vegetation and in climate-driven variation of body size. In addition, the timing of migratory flights and particularly the timely arrival at local breeding sites is crucial because fitness prospects depend on site-specific phenology. Thus, adaptations for efficient long-distance flights might be also related to conditions at destination areas. For an obligatory long-distance migrant, the common nightingale, we verified that wing length as the aerodynamically important trait, but not structural body size increased from the western to the eastern parts of the species range. In contrast with expectation from aerodynamic theory, however, wing length did not increase with increasing migration distances. Instead, wing length was associated with the phenology at breeding destinations, namely the speed of local spring green-up. We argue that longer wings are beneficial for adjusting migration speed to local conditions for birds breeding in habitats with fast spring green-up and thus short optimal arrival periods. We suggest that the speed of spring green-up at breeding sites is a fundamental variable determining the timing of migration that fine tune phenotypes in migrants across their range.

Variable detours in long-distance migration across ecological barriers and their relation to habitat availability at ground.

Migration detours, the spatial deviation from the shortest route, are a widespread phenomenon in migratory species, especially if barriers must be crossed. Moving longer distances causes additional efforts in energy and time, and to be adaptive, this should be counterbalanced by favorable condition en route. We compared migration patterns of nightingales that travelled along different flyways from their European breeding sites to the African nonbreeding sites. We tested for deviations from shortest routes and related the observed and expected routes to the habitat availability at ground during autumn and spring migration. All individuals flew detours of varying extent. Detours were largest and seasonally consistent in western flyway birds, whereas birds on the central and eastern flyways showed less detours during autumn migration, but large detours during spring migration (eastern flyway birds). Neither migration durations nor the time of arrival at destination were related to the lengths of detours. Arrival at the breeding site was nearly synchronous in birds flying different detours. Flying detours increased the potential availability of suitable broad-scale habitats en route only along the western flyway. Habitat availability on observed routes remained similar or even decreased for individuals flying detours on the central or the eastern flyway as compared to shortest routes. Thus, broad-scale habitat distribution may partially explain detour performance, but the weak detour-habitat association along central and eastern flyways suggests that other factors shape detour extent regionally. Prime candidate factors are the distribution of small suitable habitat patches at local scale as well as winds specific for the region and altitude.