Climatic effects on timing of spring migration and breeding in a long-distance migrant, the pied flycatcher Ficedula hypoleuca

Climate change has advanced the breeding dates of many bird species, but for few species we know whether this advancement is sufficient to track the advancement of the underlying levels of the food chain. For the long-distance migratory pied flycatcher Ficedula hypoleuca the advancement in breeding time has been insufficient to maintain the synchrony with their main food sources. The timing of arrival in the breeding areas from their African wintering grounds is likely to constrain the advancement of breeding date. We hypothesise that this is because in Africa they cannot predict the advancement of spring in their breeding habitat. However, long-distance migrants may advance their arrival time by migrating faster when circumstances en route are favourable. In this study we show that both arrival and breeding date depend on temperatures at their main North African staging grounds, as well as on temperature at the breeding grounds. Male arrival and average laying date were not correlated, but the positive effect of temperature in North Africa on breeding dates suggests that breeding date is indeed constrained by arrival of females. Long-distance migrants thus are able to adjust arrival and hence breeding by faster spring migration, but the degree of adjustment is probably limited as timing schedules in spring are tight. Furthermore, as climate change is affecting temperatures differently along the migratory flyway and the breeding areas, it is unlikely that arrival dates are advanced at the same rate as the timing of breeding should advance, given the advancement of the underlying levels of the food chain.     

Spring migration phenology of birds in the Northern Prairie region is correlated with local climate change

ABSTRACT In apparent response to recent periods of global warming, some migratory birds now arrive earlier at stopover sites and breeding grounds. However, the effects of this warming on arrival times vary among locations and species. Migration timing is generally correlated with temperature, with earlier arrival during warm years than during cold years, so local variation in climate change might produce different effects on migration phenology in different geographic regions. We examined trends in first spring arrival dates (FADs) for 44 species of common migrant birds in South Dakota (1971–2006) and Minnesota (1964–2005) using observations compiled by South Dakota and Minnesota Ornithologists’ Unions. We found significant trends in FAD over time for 20 species (18 arriving earlier and two later) in South Dakota and 16 species (all earlier) in Minnesota. Of these species, 10 showed similar significant trends for both states. All 10 of these species exhibited significantly earlier arrival, and all were early spring migrants, with median FADs before 10 April in both states. Eighteen of the 44 species showed significant negative correlations of FADs with either winter (December–February) or spring (arrival month plus previous month) temperatures in one or both states. Interestingly, spring temperatures in both South Dakota and Minnesota did not warm significantly from 1971–2006, but winter temperatures in both states warmed significantly over the same time period. This suggests that the warmer winters disproportionately affected early spring migrants, especially those associated with aquatic habitats (seven of the 10 species showing significantly earlier spring arrival in both states). The stronger response to climate change by early spring migrants in our study is consistent with the results of several other studies, and suggests that migrants, especially early migrants, are capable of responding to local temperature conditions experienced on wintering grounds or along the migration route.     

Arrival and onset of breeding of three passerine birds in eastern Finland tracks climatic variation and phenology of insects

Anthropogenic climate change poses a challenge to the annual cycles of migratory birds. It has become urgent to understand whether migratory birds are able to advance their spring phenology when the climate is warming and whether they are able to adjust these phenological phases to the spring phenology in their breeding areas. In this work, we studied long‐term trends in first arrival and onset of breeding for three passerine birds in eastern Finland; the pied flycatcher Ficedula hypoleuca, the common redstart Phoenicurus phoenicurus and the great tit Parus major. The pied flycatcher and the common redstart are long‐distance migrants while the great tit is a partial migrant in Finland. We asked what environmental variables best explain the first arrival or onset of breeding, if there is evidence of ‘thermal delay’ (long‐term increase in the accumulated temperatures) at arrival or onset of breeding and if the interannual variation in the onset of breeding correlates with variation in spring phenology of local insects. We found that the pied flycatcher and the common redstart had advanced their first arrival (explained by increased temperatures at the migration route), but we found no long‐term change in the onset of breeding (explained by local temperatures). Also, the onset of breeding of the great tit is tracking local temperatures. We found no or only weak evidence of thermal delay at arrival or onset of breeding for any of the species. The onsets of breeding for the pied flycatcher and the great tit are also closely tracking the spring phenology of the local insects. The stable or increasing population sizes of all three species in Finland could be a result from their ability to effectively track climatic and environmental variation.     

Forage plants of an Arctic‐nesting herbivore show larger warming response in breeding than wintering grounds,potentially disrupting migration phenology

During spring migration, herbivorous waterfowl breeding in the Arctic depend on peaks in the supply of nitrogen‐rich forage plants, following a “green wave” of grass growth along their flyway to fuel migration and reproduction. The effects of climate warming on forage plant growth are expected to be larger at the Arctic breeding grounds than in temperate wintering grounds, potentially disrupting this green wave and causing waterfowl to mistime their arrival on the breeding grounds. We studied the potential effect of climate warming on timing of food peaks along the migratory flyway of the Russian population of barnacle geese using a warming experiment with open‐top chambers. We measured the effect of 1.0–1.7°C experimental warming on forage plant biomass and nitrogen concentration at three sites along the migratory flyway (temperate wintering site, temperate spring stopover site, and Arctic breeding site) during 2 months for two consecutive years. We found that experimental warming increased biomass accumulation and sped up the decline in nitrogen concentration of forage plants at the Arctic breeding site but not at temperate wintering and stop‐over sites. Increasing spring temperatures in the Arctic will thus shorten the food peak of nitrogen‐rich forage at the breeding grounds. Our results further suggest an advance of the local food peak in the Arctic under 1–2°C climate warming, which will likely cause migrating geese to mistime their arrival at the breeding grounds, particularly considering the Arctic warms faster than the temperate regions. The combination of a shorter food peak and mistimed arrival is likely to decrease goose reproductive success under climate warming by reducing growth and survival of goslings after hatching.     

Climate change and fitness components of a migratory bird breeding in the Mediterranean region

The increase in spring temperatures in temperate regions over the last two decades has led to an advancing spring phenology, and most resident birds have responded to it by advancing their onset of breeding. The pied flycatcher (Ficedula hypoleuca) is a long‐distance migrant bird with a relatively late onset of breeding with respect to both resident birds and spring phenology in Europe. In the present correlational study, we show that some fitness components of pied flycatchers are suffering from climate change in two of the southernmost European breeding populations. In both montane study areas, temperature during May increased between 1980 and 2000 and an advancement of oak leafing was detected by using the normalized difference vegetation index (NDVI) to assess tree phenology. This might result in an advancement of the peak in availability of caterpillars, the main prey during the nestling stage. Over the past 18 yr, the time of egg laying and clutch size of pied flycatchers were not affected by the increase in spring temperatures in these Mediterranean populations. However, this increase seems to have an adverse effect on the reproductive output of pied flycatchers over the same period. Our data suggest that the mismatch between the timing of peak food supply and nestling demand caused by recent climate change might result in a reduction of parental energy expenditure that is reflected in a reduction of nestling growth and survival of fledged young in our study populations. The data seem to indicate that the breeding season has not shifted and it is the environment that has shifted away from the timing of the pied flycatcher breeding season. Mediterranean pied flycatchers were not able to advance their onset of breeding, probably, because they are constrained by their late arrival date and their restricted high altitude breeding habitat selection near the southern border of their range.     

Central European Greylag Geese Anser anser show a shortening of migration distance and earlier spring arrival over 60 years

Global climate change can cause pronounced changes in species? migratory behaviour. Numerous recent studies have demonstrated climate‐driven changes in migration distance and spring arrival date in waterbirds, but detailed studies based on long‐term records of individual recapture or re‐sighting events are scarce. Using re‐sighting data from 430 marked individuals spanning a 60‐year period (winters 1956/1957 to 2015/2016), we assessed patterns in migration distance and spring arrival date, wintering‐site fidelity and survival in the increasing central European breeding population of Greylag Geese Anser anser. We demonstrate a long‐term decrease in migration distance, changes in the wintering range caused by winter partial short‐stopping, and the earlier arrival of geese on their breeding grounds. Greylag Geese marked on central Europe moulting grounds have not been recorded wintering in Spain since 1986 or in Tunisia and Algeria since 2004. The migration distance and spring arrival of geese indicated an effect of temperature at the breeding site and values of the NAO index. Greylag Geese migrate shorter distances and arrive earlier in milder winters. We suggest that shifts in the migratory behaviour of Central European Greylag Geese are individual temperature‐dependent decisions to take advantage of wintering grounds becoming more favourable closer to their breeding grounds, allowing birds to acquire breeding territories earlier.     

Tracking climate impacts on the migratory monarch butterfly

Understanding the impacts of climate on migratory species is complicated by the fact that these species travel through several climates that may be changing in diverse ways throughout their complete migratory cycle. Most studies are not designed to tease out the direct and indirect effects of climate at various stages along the migration route. We assess the impacts of spring and summer climate conditions on breeding monarch butterflies, a species that completes its annual migration cycle over several generations. No single, broad‐scale climate metric can explain summer breeding phenology or the substantial year‐to‐year fluctuations observed in population abundances. As such, we built a Poisson regression model to help explain annual arrival times and abundances in the Midwestern United States. We incorporated the climate conditions experienced both during a spring migration/breeding phase in Texas as well as during subsequent arrival and breeding during the main recruitment period in Ohio. Using data from a state‐wide butterfly monitoring network in Ohio, our results suggest that climate acts in conflicting ways during the spring and summer seasons. High spring precipitation in Texas is associated with the largest annual population growth in Ohio and the earliest arrival to the summer breeding ground, as are intermediate spring temperatures in Texas. On the other hand, the timing of monarch arrivals to the summer breeding grounds is not affected by climate conditions within Ohio. Once in Ohio for summer breeding, precipitation has minimal impacts on overall abundances, whereas warmer summer temperatures are generally associated with the highest expected abundances, yet this effect is mitigated by the average seasonal temperature of each location in that the warmest sites receive no benefit of above average summer temperatures. Our results highlight the complex relationship between climate and performance for a migrating species and suggest that attempts to understand how monarchs will be affected by future climate conditions will be challenging.     

Climate change, migratory connctivity and changes in laying date and clutch size of the pied flycatcher

We examined long-term (1943–2003) variability in laying dates and clutch sizes in a Finnish population of the pied flycatcher Ficedula hypoleuca Pallas, and analysed whether potential changes were explained by changes in climatic factors at the wintering area in Africa, at migration route or at breeding grounds. Among-year variation in both mean and skewness of laying dates increased, which for mean laying date appeared to be explained by variability of temperatures at the breeding grounds and for skewness by variable temperature trends along the migration route. Pied flycatchers bred earlier in warm springs, but despite a warming trend in pre-laying temperatures, the laying dates tended to delay. Laying dates became continuously later in relation to the phenology of the environment. Mean clutch size decreased with time when mean laying date was controlled for, but the climatic factors did not appear to explain the decrease. The advancement of spring phenology may have shifted some food sources needed for egg-laying, thus leading to later laying and smaller clutches. Variation in clutch size increased when wintering conditions were favourable so that clutch size distribution was skewed with a tail of small clutches when there had been lot of rainfall (more vegetation and insects) in the wintering area. We suggest that when ecological conditions during winter were good, the tail of small clutches represented low-quality individuals that were not able to breed after bad winters. Our analyses demonstrate that measures of spread and symmetry give different information about population level changes than means, and thus complement the understanding of the potential influences of climate change on populations.     

Global change and the distributional dynamics of migratory bird populations wintering in Central America

Understanding the susceptibility of highly mobile taxa such as migratory birds to global change requires information on geographic patterns of occurrence across the annual cycle. Neotropical migrants that breed in North America and winter in Central America occur in high concentrations on their non‐breeding grounds where they spend the majority of the year and where habitat loss has been associated with population declines. Here, we use eBird data to model weekly patterns of abundance and occurrence for 21 forest passerine species that winter in Central America. We estimate species’ distributional dynamics across the annual cycle, which we use to determine how species are currently associated with public protected areas and projected changes in climate and land‐use. The effects of global change on the non‐breeding grounds is characterized by decreasing precipitation, especially during the summer, and the conversion of forest to cropland, grassland, or peri‐urban. The effects of global change on the breeding grounds are characterized by increasing winter precipitation, higher temperatures, and the conversion of forest to peri‐urban. During spring and autumn migration, species are projected to encounter higher temperatures, forests that have been converted to peri‐urban, and increased precipitation during spring migration. Based on current distributional dynamics, susceptibility to global change is characterized by the loss of forested habitats on the non‐breeding grounds, warming temperatures during migration and on the breeding grounds, and declining summer rainfall on the non‐breeding grounds. Public protected areas with low and medium protection status are more prevalent on the non‐breeding grounds, suggesting that management opportunities currently exist to mitigate near‐term non‐breeding habitat losses. These efforts would affect more individuals of more species during a longer period of the annual cycle, which may create additional opportunities for species to respond to changes in habitat or phenology that are likely to develop under climate change.     

Climate warming,ecological mismatch at arrival and population decline in migratory birds

Climate is changing at a fast pace, causing widespread, profound consequences for living organisms. Failure to adjust the timing of life-cycle events to climate may jeopardize populations by causing ecological mismatches to the life cycle of other species and abiotic factors. Population declines of some migratory birds breeding in Europe have been suggested to depend on their inability to adjust migration phenology so as to keep track of advancement of spring events at their breeding grounds. In fact, several migrants have advanced their spring arrival date, but whether such advancement has been sufficient to compensate for temporal shift in spring phenophases or, conversely, birds have become ecologically mismatched, is still an unanswered question, with very few exceptions. We used a novel approach based on accumulated winter and spring temperatures (degree-days) as a proxy for timing of spring biological events to test if the progress of spring at arrival to the breeding areas by 117 European migratory bird species has changed over the past five decades. Migrants, and particularly those wintering in sub-Saharan Africa, now arrive at higher degree-days and may have therefore accumulated a ‘thermal delay’, thus possibly becoming increasingly mismatched to spring phenology. Species with greater ‘thermal delay’ have shown larger population decline, and this evidence was not confounded by concomitant ecological factors or by phylogenetic effects. These findings provide general support to the largely untested hypotheses that migratory birds are becoming ecologically mismatched and that failure to respond to climate change can have severe negative impacts on their populations. The novel approach we adopted can be extended to the analysis of ecological consequences of phenological response to climate change by other taxa.     

Fitness Consequences of Timing of Migration and Breeding in Cormorants

In most bird species timing of breeding affects reproductive success whereby early breeding is favoured. In migratory species migration time, especially arrival at the breeding grounds, and breeding time are expected to be correlated. Consequently, migration time should also have fitness consequences. However, in contrast to breeding time, evidence for fitness consequences of migration time is much more limited. Climate change has been shown to negatively affect the synchrony between trophic levels thereby leading to directional selection on timing but again direct evidence in avian migration time is scarce. We here analysed fitness consequences of migration and breeding time in great cormorants and tested whether climate change has led to increased selection on timing using a long-term data set from a breeding colony on the island of Vorsø (Denmark). Reproductive success, measured as number of fledglings, correlated with breeding time and arrival time at the colony and declined during the season. This seasonal decline became steeper during the study period for both migration and breeding time and was positively correlated to winter/spring climate, i.e. selection was stronger after warmer winters/springs. However, the increasing selection pressure on timing seems to be unrelated to climate change as the climatic variables that were related to selection strength did not increase during the study period. There is indirect evidence that phenology or abundances of preferred prey species have changed which could have altered selection on timing of migration and breeding.     

Impact of temperature on the breeding performance and selection patterns in lesser kestrels Falco naumanni

Adjusting breeding phenology to climate fluctuations can be problematic for migratory birds as they have to account for local environmental conditions on the breeding grounds while migrating from remote wintering areas. Predicting general responses to climate change is not straightforward, because these responses vary between migrant species due to the species‐specific ecological drivers of breeding behaviour. Therefore more information is needed on species with different ecological requirements, including data on heritability of migration, factors driving phenological changes and how climate affects selection pressures. Here, we measure heritability in settlement dates and the effect of local climate at the breeding grounds on settlement dates, reproductive success and selection patterns in a French population of a trans‐Saharan migratory insectivorous raptor, the lesser kestrel Falco naumanni, monitored and ringed since 1996. Heritability of settlement dates was low (0.07 ± 0.03), indicating a weak evolutionary potential. Nevertheless, plasticity in settlement dates in response to temperatures allowed earlier settlement when early spring was warmer than average. Reproductive success and selection patterns were strongly affected by temperature during settlement and chick rearing respectively. Warmer spring decreased selection for earlier settling and warmer early summer increased reproductive success. Interestingly, selection for earlier settling was more intense in cooler springs, contrasting with patterns from passerines lagging behind food peaks. Altogether, these results suggest a positive effect of warmer temperatures on breeding performances of lesser kestrels most likely because the French population is at the coolest boundary of the species European breeding range.     

Differences in shifts of wintering and breeding ranges lead to changing migration distances in European birds

Studies on the impact of climate change on the distributions of bird species in Europe have largely focused on one season at a time, especially concerning summer breeding ranges. We investigated whether migratory bird species show consistent range shifts over the past 55 yr in both breeding and wintering areas or if these shifts are independent. We then analyzed whether patterns in changing migration distances of Finnish breeding birds could be explained by habitat use, phylogeny or body size. We used long‐term datasets from the Finnish ringing centre to analyze the mean wintering latitudes of 29 species of Finnish breeding birds, then used breeding distribution data to make predictions as to whether certain species were migrating shorter or longer distances based on the comparative shifts in the wintering and breeding grounds. Our data reveal species‐specific differences in changing migration distances. We show that for many species, long‐term shifts in wintering ranges have not followed the same patterns as those in the breeding range, leading to differences in migration distances over time. We conclude that species are not adjusting predictably to climate change in their wintering grounds, leading to changing migration distances in some, but not all, species breeding in Finland. This research fills an important gap in the current climate change biology literature, focusing on individuals’ entire life histories and revealing new complexities in range shift patterns.     

Patterns of bird migration phenology in South Africa suggest northern hemisphere climate as the most consistent driver of change

Current knowledge of phenological shifts in Palearctic bird migration is largely based on data collected on migrants at their breeding grounds; little is known about the phenology of these birds at their nonbreeding grounds, and even less about that of intra‐African migrants. Because climate change patterns are not uniform across the globe, we can expect regional disparities in bird phenological responses. It is also likely that they vary across species, as species show differences in the strength of affinities they have with particular habitats and environments. Here, we examine the arrival and departure of nine Palearctic and seven intra‐African migratory species in the central Highveld of South Africa, where the former spend their nonbreeding season and the latter their breeding season. Using novel analytical methods based on bird atlas data, we show phenological shifts in migration of five species – red‐backed shrike, spotted flycatcher, common sandpiper, white‐winged tern (Palearctic migrants), and diederik cuckoo (intra‐African migrant) – between two atlas periods: 1987–1991 and 2007–2012. During this time period, Palearctic migrants advanced their departure from their South African nonbreeding grounds. This trend was mainly driven by waterbirds. No consistent changes were observed for intra‐African migrants. Our results suggest that the most consistent drivers of migration phenological shifts act in the northern hemisphere, probably at the breeding grounds.     

Photoperiodic response may facilitate adaptation to climatic change in long-distance migratory birds

Recent climatic change is causing spring events in northern temperate regions to occur earlier in the year. As a result, migratory birds returning from tropical wintering sites may arrive too late to take full advantage of the food resources on their breeding grounds. Under these conditions, selection will favour earlier spring arrival that could be achieved by overwintering closer to the breeding grounds. However, it is unknown how daylength conditions at higher latitudes will affect the timing of life cycle stages. Here, we show in three species of Palaearctic-African migratory songbirds that a shortening of migration distance induces an advancement of springtime activities. Birds exposed to daylengths simulating migration to and wintering in southern Europe considerably advanced their spring migratory activity and testicular development. This response to the novel photoperiodic environment will enable birds wintering further north to advance spring arrival and to start breeding earlier. Thus, phenotypic flexibility in response to the photoperiod may reinforce selection for shorter migration distance if spring temperatures continue to rise.     

Climate change leads to differential shifts in the timing of annual cycle stages in a migratory bird

Shifts in reproductive phenology due to climate change have been well documented in many species but how, within the same species, other annual cycle stages (e.g. moult, migration) shift relative to the timing of breeding has rarely been studied. When stages shift at different rates, the interval between stages may change resulting in overlaps, and as each stage is energetically demanding, these overlaps may have negative fitness consequences. We used long‐term data of a population of European pied flycatchers (Ficedula hypoleuca) to investigate phenological shifts in three annual cycle stages: spring migration (arrival dates), breeding (egg‐laying and hatching dates) and the onset of postbreeding moult. We found different advancements in the timing of breeding compared with moult (moult advances faster) and no advancement in arrival dates. To understand these differential shifts, we explored which temperatures best explain the year‐to‐year variation in the timing of these stages, and show that they respond differently to temperature increases in the Netherlands, causing the intervals between arrival and breeding and between breeding and moult to decrease. Next, we tested the fitness consequences of these shortened intervals. We found no effect on clutch size, but the probability of a fledged chick to recruit increased with a shorter arrival‐breeding interval (earlier breeding). Finally, mark–recapture analyses did not detect an effect of shortened intervals on adult survival. Our results suggest that the advancement of breeding allows more time for fledgling development, increasing their probability to recruit. This may incur costs to other parts of the annual cycle, but, despite the shorter intervals, there was no effect on adult survival. Our results show that to fully understand the consequences of climate change, it is necessary to look carefully at different annual cycle stages, especially for organisms with complex cycles, such as migratory birds.     

Climate change can alter competitive relationships between resident and migratory birds

Climate change could affect resource competition between resident and migratory bird species by changing the interval between their onsets of breeding or by altering their population densities. We studied interspecific nest-hole competition between resident great tits and migrant pied flycatchers in South-Western Finland over the past five decades (1953-2005). We found that appearance of fatal take-over trials, the cases where a pied flycatcher tried to take over a great tit nest but was killed by the tit, increased with a reduced interspecific laying date interval and with increasing densities of both tits and flycatchers. The probability of pied flycatchers taking over great tit nests increased with the density of pied flycatchers. Laying dates of the great tit and pied flycatcher are affected by the temperatures of different time periods, and divergent changes in these temperatures could consequently modify their competitive interactions. Densities are a result of reproductive success and survival, which can be affected by separate climatic factors in the resident great tit and trans-Saharan migrant pied flycatcher. On these bases we conclude that climate change has a great potential to alter the competitive balance between these two species.     

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.     

Light‐level geolocators reveal migratory connectivity in European populations of pied flycatchers Ficedula hypoleuca

Understanding what drives or prevents long‐distance migrants to respond to environmental change requires basic knowledge about the wintering and breeding grounds, and the timing of movements between them. Both strong and weak migratory connectivity have been reported for Palearctic passerines wintering in Africa, but this remains unknown for most species. We investigated whether pied flycatchers Ficedula hypoleuca from different breeding populations also differ in wintering locations in west‐Africa. Light‐level geolocator data revealed that flycatchers from different breeding populations travelled to different wintering sites, despite similarity in routes during most of the autumn migration. We found support for strong migratory connectivity showing an unexpected pattern: individuals breeding in Fennoscandia (S‐Finland and S‐Norway) wintered further west compared to individuals breeding at more southern latitudes in the Netherlands and SW‐United Kingdom. The same pattern was found in ring recovery data from sub‐Saharan Africa of individuals with confirmed breeding origin. Furthermore, population‐specific migratory connectivity was associated with geographical variation in breeding and migration phenology: birds from populations which breed and migrate earlier wintered further east than birds from ‘late’ populations. There was no indication that wintering locations were affected by geolocation deployment, as we found high repeatability and consistency in δ¹³C and δ¹⁵N stable isotope ratios of winter grown feathers of individuals with and without a geolocator. We discuss the potential ecological factors causing such an unexpected pattern of migratory connectivity. We hypothesise that population differences in wintering longitudes of pied flycatchers result from geographical variation in breeding phenology and the timing of fuelling for spring migration at the wintering grounds. Future research should aim at describing how temporal dynamics in food availability across the wintering range affects migration, wintering distribution and populations’ capacity to respond to environmental changes.     

Climate change and timing of spring migration in the long-distance migrant Ficedula hypoleuca in central Europe: the role of spatially different temperature changes along migration routes

Coinciding with increasing spring temperatures in Europe, many migrants have advanced their arrival or passage times over the last decades. However, some species, namely long-distance migrants, could be constrained in their arrival dates due to their largely inherited migratory behaviour and thus a likely inflexibility in their response to exogenous factors. To examine this hypothesis for pied flycatchers (Ficedula hypoleuca), we tested the effects of the temperature regimes along their migration routes north of the Sahara on their arrival times in central Europe. To do so, we developed a site-independent large-scale approach based on temperature data available on the Internet. Temperature regimes along the migration routes of pied flycatchers within Europe convincingly correlate with their first arrival times. It can be concluded that the progression of spring migration in this species is strongly influenced by temperature en route. Because of the recent inconsistent climatic changes in various parts of Europe, we hypothesize that individuals migrating along different routes will be unequally affected by further climatic changes.