Ecological conditions in wintering and passage areas as determinants of timing of spring migration in trans-Saharan migratory birds

1.?Climate change has been associated with shifts in the timing of biological events, including the spring arrival of migratory birds. Early arrival at breeding sites is an important life-history trait, usually associated with higher breeding success and therefore, susceptible to selection and evolution in response to changing climatic conditions. 2.?Here, we examine the effect of changes in the environmental conditions of wintering and passage areas on the mean passage time of 13 trans-Saharan passerines during their spring migration through the western Mediterranean over the 15 years from 1993 to 2007. 3.?We found that most of the species studied have been advancing the timing of their passage in recent years. However, annual variation in the mean date of passage was positively correlated with vegetation growth (measured as the normalized difference vegetation index [NDVI]) both in the Sahel (the region of departure) and in northern Africa (the passage area). Thus, migration dates were delayed in years with high primary productivity in passage and wintering zones. All species seem to respond similarly to NDVI in the Sahel; however, late migrants were less affected by ecological conditions in northern Africa than those migrating earlier, suggesting differences based on species ecology. 4.?Mean timing of passage was not related to the North Atlantic Oscillation (NAO), El Ni?o-Southern Oscillation (ENSO), temperature or NDVI in the species-specific wintering areas (the overwintering region) when analysed in combination with the other covariates. 5.?Our findings show that ecological conditions in the winter quarters (specifically the Sahel) and en route are relevant factors influencing trends in the passage dates of trans-Saharan migratory birds on the southern fringe of Europe. Possible long-term consequences for late arriving spring migrants are discussed.     

Ecological conditions during winter predict arrival date at the breeding quarters in a trans-Saharan migratory bird

We studied variation in arrival date to the breeding colonies in Italy of a trans‐Saharan migratory bird, the barn swallow Hirundo rustica, in relation to variation in ecological conditions, as reflected by the normalized difference vegetation index (NDVI), in the winter quarters. Arrival date of old but not young individuals captured during consecutive breeding seasons was earlier after winters with favourable conditions. Change in arrival date in relation to change in NDVI was similar in the two sexes. Change in arrival date significantly and positively predicted change in breeding date. As a result of increased frequency of second broods determined by earlier arrival, the number of fledged offspring per season was larger after African winters with good in comparison to poor ecological conditions for barn swallows. This is the first study demonstrating phenotypic plasticity in migration phenology of a long‐distance migratory bird in relation to ecological conditions during wintering.     

Climate anomalies affect annual survival rates of swifts wintering in sub‐Saharan Africa

Several species of migratory swifts breed in the Western Palearctic, but they differ in reproductive traits and nonbreeding areas explored in Africa. We examined survival and recapture probabilities of two species of swifts by capture–mark–recapture data collected in northern Italy (Pallid Swift Apus pallidus in Carmagnola, Turin, and Common Swift Apus apus in Guiglia, Modena) in the breeding season (May–July). Apparent survival rates were relatively high (>71%), comparable to other studies of European swifts, but showed marked annual variations. We used geolocators to establish the exact wintering areas of birds breeding in our study colonies. Common Swifts explored the Sahel zone during migration and spent the winter in SE Africa, while the Pallid Swifts remained in the Sahel zone for a longer time, shifting locations southeast down to Cameroun and Nigeria later in winter. These movements followed the seasonal rains from north to south (October to December). In both species, we found large yearly differences in survival probabilities related to different climatic indices. In the Pallid Swift, wintering in Western Africa, the Sahel rainfall index best explained survival, with driest seasons associated with reduced survival. In the Common Swift, wintering in SE Africa, the El Niño–Southern Oscillation (ENSO) cycle performed significantly better than Sahel rainfall or North Atlantic Oscillation (NAO). Extreme events and precipitation anomalies in Eastern Africa during La Niña events resulted in reduced survival probabilities in Common Swifts. Our study shows that the two species of swifts have similar average annual survival, but their survival varies between years and is strongly affected by different climatic drivers associated with their respective wintering areas. This finding could suggest important ecological diversification that should be taken into account when comparing survival and area use of similar species that migrate between temperate breeding areas and tropical wintering areas.     

Local temperature fine-tunes the timing of spring migration in birds

Evidence for climate-driven phenological changes is rapidly increasing at all trophic levels. Our current poor knowledge of the detailed control of bird migration from the level of genes and hormonal control to direct physiological and behavioral responses hampers our ability to understand and predict consequences of climatic change for migratory birds. In order to better understand migration phenology and adaptation in environmental changes, we here assess the scale at which weather affects timing of spring migration in passerine birds. We use three commonly used proxies of spring-time climatic conditions: (1) vegetation "greenness" (NDVI) in Europe, (2) local spring temperatures in northern Europe, and (3) the North Atlantic Oscillation Index (NAO) as predictors of the phenology of avian migration as well as the strength of their effect on different subsets of populations and the dependence of correlations on species-specific migratory strategy. We analyze phenological patterns of the entire spring migration period in 12 Palaearctic passerine species, drawing on long-term data collected at three locations along a longitudinal gradient situated close to their northern European breeding area. Local temperature was the best single predictor of phenology with the highest explanatory power achieved in combination with NAO. Furthermore, early individuals are more affected by climatic variation compared to individuals on later passage, indicating that climatic change affects subsets of migratory populations differentially. Species wintering closer to the breeding areas were affected more than were those travelling longer distances and this pattern was strongest for the earliest subsets of the population. Overall, our results suggest that at least early subsets of the population are affected by local conditions and early birds use local conditions to fine-tune the date of their spring arrival while individuals arriving later are driven by other factors than local conditions e.g. endogenous control. Understanding what cues migratory organisms use to arrive at an optimum time is important for increasing our knowledge of fundamental issues like decision making in organisms during migration and is crucial for future protection of migratory organisms.     

Weekend bias in Citizen Science data reporting: implications for phenology studies

Studies of bird phenology can help elucidate the effects of climate change on wildlife species but observations over broad spatial scales are difficult without a network of observers. Recently, networks of citizen volunteers have begun to report first arrival dates for many migratory species. Potential benefits are substantial (e.g., understanding ecological processes at broad spatial and temporal scales) if known biases of citizen data reporting are identified and addressed. One potential source of bias in bird phenology studies is the tendency for more “first” migratory arrivals to be reported on weekends than on weekdays. We investigated weekend bias in data reporting for five common bird species in North America (Baltimore Oriole, Icterus galbula; Barn Swallow, Hirundo rustica; Chimney Swift, Chaetura pelagica; Purple Martin, Progne subis; and Ruby-throated Hummingbird, Archilochus colubris), and assessed whether this bias affected mean arrival dates reported using data from historical (1880–1969; N?=?25,555) and recent (1997–2010; N?=?63,149) Citizen Science databases. We found a greater percentage of first arrivals reported on weekends and small but significant differences in mean arrival dates (approximately 0.5 days) for four of five species. Comparing time periods, this weekend bias decreased from 33.7 % and five species in the historical time period to 32 % and three species in the recent, perhaps related to changes in human activity patterns. Our results indicate that weekend bias in citizen data reporting is decreasing over time in North America and including a ‘day of week’ term in models examining changes in phenology could help make conclusions more robust.     

Spring arrival patterns of the Cuckoo Cuculus canorus,Nightingale Luscinia megarhynchos and Spotted Flycatcher Musciapa striata in Britain

Data on the arrival dates of Cuckoo, Nightingale and Spotted Flycatcher published in the Royal Meteorological Society phenological reports are presented. From these records, contour maps showing the pattern of progression through Britain were produced for each species. Of the three species, the Cuckoo arrived earliest and progressed the slowest from south to north; it entered Britain by the southeast. The Nightingale arrived later than the Cuckoo and progressed along a southeast to northwest axis. The Spotted Flycatcher was the last to arrive, but progressed with the greatest speed from south to north. Arrival dates of all three species were influenced by the temperature in southern Europe in the month preceding arrival in Britain and by the difference between these temperatures in southern Europe and those a month later in France or Britain. The diminishing population levels of Nightingale may have had a masking effect on its arrival dates. Although mean arrival dates for the three species were strongly influenced by temperature, long-term trend effects should be interpreted with caution as other factors also influence mean arrival dates.     

Long‐term changes in the migration phenology of UK breeding birds detected by large‐scale citizen science recording schemes

The timing of migration is one of the key life‐history parameters of migratory birds. It is expected to be under strong selection, to be sensitive to changing environmental conditions and to have implications for population dynamics. However, most phenological studies do not describe arrival and departure phenologies for a species in a way that is robust to potential biases, or that can be clearly related to breeding populations. This hampers our ability to understand more fully how climate change may affect species’ migratory strategies, their life histories and ultimately their population dynamics. Using generalized additive models (GAMs) and extensive large‐scale data collected in the UK over a 40‐year period, we present standardized measures of migration phenology for common migratory birds, and examine how the phenology of bird migration has changed in the UK since the 1960s. Arrival dates for 11 of 14 common migrants became significantly earlier, with six species advancing their arrival by more than 10 days. These comprised two species, Blackcap Sylvia atricapilla and Chiffchaff Phylloscopus collybita, which winter closest to Britain in southern Europe and the arid northern zone of Africa, Common Redstart Phoenicurus phoenicurus, which winters in the arid zone, and three hirundines (Sand Martin Riparia riparia, House Martin Delichon urbicum and Barn Swallow Hirundo rustica), which winter in different parts of Africa. Concurrently, departure dates became significantly later for four of the 14 species and included species that winter in southern Europe (Blackcap and Chiffchaff) and in humid zones of Africa (Garden Warbler Sylvia borin and Whinchat Saxicola rubetra). Common Swift Apus apus was the exception in departing significantly earlier. The net result of earlier arrival and later departure for most species was that length of stay has become significantly longer for nine of the 14 species. Species that have advanced their timing of arrival showed the most positive trends in abundance, in accordance with previous studies. Related in part to earlier arrival and the relationship above, we also show that species extending their stay in Great Britain have shown the most positive trends. Further applications of our modelling approach will provide opportunities for more robust tests of relationships between phenological change and population dynamics than have been possible previously.     

Earlier arrival of some farmland migrants in western Poland

A dataset consisting of observations on the first arrival dates (FAD) of 16 species of birds from a locality in western Poland during the period 19131996 is presented and analysed. For 14 of the 16 species, there is evidence of a trend towards earliness in recent years. In the 19701996 period this trend was significant for Pied Wagtail, Wood Pigeon, Black Redstart and Barn Swallow, and marginally significant for a further four species: European Serin, Blackcap, Common Nightingale and Red-backed Shrike. Only two species showed the opposite trend in the 19701996 period: Lesser Whitethroat and Whinchat, of which only the former is (marginally) significant. The trend towards earliness was more pronounced for short-distance (European) migrants than for long-distance (African) migrants. Short-distance migrants generally displayed a correlation with spring temperatures in the breeding locality, a feature less obvious in the long-distance migrants.     

Population-scale drivers of individual arrival times in migratory birds

1. In migratory species, early arrival on the breeding grounds can often enhance breeding success. Timing of spring migration is therefore a key process that is likely to be influenced both by factors specific to individuals, such as the quality of winter and breeding locations and the distance between them, and by annual variation in weather conditions before and during migration. 2. The Icelandic black-tailed godwit Limosa limosa islandica population is currently increasing and, throughout Iceland, is expanding into poorer quality breeding areas. Using a unique data set of arrival times in Iceland in different years for individuals of known breeding and wintering locations, we show that individuals breeding in lower quality, recently occupied and colder areas arrive later than those from traditionally occupied areas. The population is also expanding into new wintering areas, and males from traditionally occupied winter sites also arrive earlier than those occupying novel sites. 3. Annual variation in timing of migration of individuals is influenced by large-scale weather systems (the North Atlantic Oscillation), but between-individual variation is a stronger predictor of arrival time than the NAO. Distance between winter and breeding sites does not influence arrival times. 4. Annual variation in timing of migration is therefore influenced by climatic factors, but the pattern of individual arrival is primarily related to breeding and winter habitat quality. These habitat effects on arrival patterns are likely to operate through variation in individual condition and local-scale density-dependent processes. Timing of migration thus appears to be a key component of the intricate relationship between wintering and breeding grounds in this migratory system.     

Influences of the atmospheric patterns on unstable climate-growth associations of western Mediterranean forests

Quantifying climate-growth associations is needed to evaluate how forest productivity will respond to climate change. Year-to-year fluctuations in forest productivity and radial growth are partly explained by local climatic conditions driven by large-scale atmospheric patterns. This is illustrated by Iberian forests in the western Mediterranean Basin, which are subjected to complex climatic and atmospheric influences such as Atlantic and Mediterranean cyclogenesis. The North Atlantic Oscillation (NAO) is one of the major atmospheric circulation patterns affecting Iberian forests since positive winter NAO phases lead to dry and warm conditions. The Western Mediterranean Oscillation (WeMO) may also explain Iberian forest growth in some areas since this index captures Mediterranean cyclogenesis and WeMO negative phases are linked to warm and wet spring to summer conditions. Here, we analyze the associations between atmospheric patterns, climate and tree growth and we determine if they are changing through time. We use dendrochronology to relate radial growth of four tree species (Pyrenean oak, Sweet chestnut, Maritime pine and Scots pine) growing in western Spain to climate conditions and the NAO and WeMO indices. Winter and early spring temperatures increased since the 1950s in the area whereas the negative association between winter precipitation and the NAO strengthened since then. However, mean temperature rise was particularly evident since the 1970s. Growth was reduced by dry conditions during the growing season (spring and summer), but also by cold and dry conditions during the previous autumn and winter. This explains why the NAO January and the WeMo April indices were negative to growth of three species excluding Pyrenean oak. The early 1970s reflected an inflection point in the instability of climate-growth associations in the study area. We conclude that the winter NAO is a relevant driver of forest growth in the western Iberian Peninsula forests but additional atmospheric patterns (WeMO) also affect, albeit to a minor extent, these forests.     

Do migratory pathways affect the regional abundance of wintering birds? A test in northern Spain

Aim The abundance distribution of organisms at regional scales is commonly interpreted as the result of spatial variation in habitat suitability. However, the possibility that geography itself may affect patterns of distribution has received less attention. For example, the abundance of wintering bird populations might be influenced by the cost of reaching areas located far away from the main migratory pathways. We studied the abundance distribution of three common migratory passerines (meadow pipits, Anthus pratensis; common chaffinches, Fringilla coelebs; and European robins, Erithacus rubecula) wintering in farmlands located in the 600‐km long Cantabrian coastal sector of northern Spain, roughly perpendicular to the west Pyrenean migratory pathway that drives European migrant birds into the Iberian Peninsula. Location The study area occupies a belt located between the Atlantic coast and the Cantabrian Mountains in northern Spain. Methods We counted wintering and breeding birds and measured the structure of vegetation and environmental variables (altitude, rainfall, temperature) in 68 farmlands distributed at different distances from the west Pyrenean migratory flyway. We also studied the distribution of birds ringed in central and northern Europe and recovered in the study area between October and February. Analyses were based on single univariate statistics (chi‐square tests), ordination by principal components analysis and multiple regression. Results Controlling for the effects of climate, vegetation structure and local abundance of breeding conspecifics, the winter abundance of all three species decreased with the distance from their main migratory route in the western Pyrenees. Such patterns fitted well to the observed distribution of ringing recoveries. Main conclusions Our results support a link between the movements of birds along the Pyrenean migratory pathway and their winter abundance in northern Spain. According to this view, the sectors located near the migratory pathway seem to be more easily occupied by migrants, supporting the idea that proximity to passage areas may explain the fine‐grain regional patterning of species abundance in wintering grounds.     

Climatic patterns in the establishment of wintering areas by North American migratory birds

Long‐distance migration in birds is relatively well studied in nature; however, one aspect of this phenomenon that remains poorly understood is the pattern of distribution presented by species during arrival to and establishment of wintering areas. Some studies suggest that the selection of areas in winter is somehow determined by climate, given its influence on both the distribution of bird species and their resources. We analyzed whether different migrant passerine species of North America present climatic preferences during arrival to and departure from their wintering areas. We used ecological niche modeling to generate monthly potential climatic distributions for 13 migratory bird species during the winter season by combining the locations recorded per month with four environmental layers. We calculated monthly coefficients of climate variation and then compared two GLM (generalized linear models), evaluated with the AIC (Akaike information criterion), to describe how these coefficients varied over the course of the season, as a measure of the patterns of establishment in the wintering areas. For 11 species, the sites show nonlinear patterns of variation in climatic preferences, with low coefficients of variation at the beginning and end of the season and higher values found in the intermediate months. The remaining two species analyzed showed a different climatic pattern of selective establishment of wintering areas, probably due to taxonomic discrepancy, which would affect their modeled winter distribution. Patterns of establishment of wintering areas in the species showed a climatic preference at the macroscale, suggesting that individuals of several species actively select wintering areas that meet specific climatic conditions. This probably gives them an advantage over the winter and during the return to breeding areas. As these areas become full of migrants, alternative suboptimal sites are occupied. Nonrandom winter area selection may also have consequences for the conservation of migratory bird species, particularly under a scenario of climate change.     

Climate and the complexity of migratory phenology: sexes,migratory distance,and arrival distributions

The intra- and inter-season complexity of bird migration has received limited attention in climatic change research. Our phenological analysis of 22 species collected in Chicago, USA, (1979–2002) evaluates the relationship between multi-scalar climate variables and differences (1) in arrival timing between sexes, (2) in arrival distributions among species, and (3) between spring and fall migration. The early migratory period for earliest arriving species (i.e., short-distance migrants) and earliest arriving individuals of a species (i.e., males) most frequently correlate with climate variables. Compared to long-distance migrant species, four times as many short-distance migrants correlate with spring temperature, while 8 of 11 (73%) of long-distance migrant species’ arrival is correlated with the North Atlantic Oscillation (NAO). While migratory phenology has been correlated with NAO in Europe, we believe that this is the first documentation of a significant association in North America. Geographically proximate conditions apparently influence migratory timing for short-distance migrants while continental-scale climate (e.g., NAO) seemingly influences the phenology of Neotropical migrants. The preponderance of climate correlations is with the early migratory period, not the median of arrival, suggesting that early spring conditions constrain the onset or rate of migration for some species. The seasonal arrival distribution provides considerable information about migratory passage beyond what is apparent from statistical analyses of phenology. A relationship between climate and fall phenology is not detected at this location. Analysis of the within-season complexity of migration, including multiple metrics of arrival, is essential to detect species’ responses to changing climate as well as evaluate the underlying biological mechanisms.     

Does wintering north or south of the Sahara correlate with timing and breeding performance in black‐tailed godwits?

Migrating long distances requires time and energy, and may interact with an individual's performance during breeding. These seasonal interactions in migratory animals are best described in populations with disjunct nonbreeding distributions. The black‐tailed godwit (Limosa limosa limosa), which breeds in agricultural grasslands in Western Europe, has such a disjunct nonbreeding distribution: The majority spend the nonbreeding season in West Africa, while a growing number winters north of the Sahara on the Iberian Peninsula. To test whether crossing the Sahara has an effect on breeding season phenology and reproductive parameters, we examined differences in the timing of arrival, breeding habitat quality, lay date, egg volume, and daily nest survival among godwits (154 females and 157 males), individually marked in a breeding area in the Netherlands for which wintering destination was known on the basis of resightings. We also examined whether individual repeatability in arrival date differed between birds wintering north or south of the Sahara. Contrary to expectation, godwits wintering south of the Sahara arrived two days earlier and initiated their clutch six days earlier than godwits wintering north of the Sahara. Arrival date was equally repeatable for both groups, and egg volume larger in birds wintering north of the Sahara. Despite these differences, we found no association between wintering location and the quality of breeding habitat or nest survival. This suggests that the crossing of an important ecological barrier and doubling of the migration distance, twice a year, do not have clear negative reproductive consequences for some long‐distance migrants.     

Geographical and environmental factors affecting the distribution of wintering black storks Ciconia nigra in the Iberian Peninsula

Here we explore the environmental and geographical factors affecting the winter distribution of the black stork Ciconia nigra in the Iberian Peninsula, where an increasing number of individuals have remained to winter in the last two decades. We recorded 179 locations of 54 ringed individuals between 1988 and 2011 to map the species habitat suitability with MaxEnt, a machine‐learning technique based on the principle of maximum entropy. The migratory movements of 25 birds equipped with satellite transmitters were used to define the autumnal migratory flyway used by most storks crossing the Peninsula as well as to define the wintering period. The aim was to test if the number of wintering storks was positively correlated to habitat suitability and negatively correlated to the flyway distance. Data provided by an extensive count across Portugal and Spain during the 2012–2013 winter supported the findings that black storks were more abundant in areas of high habitat suitability close to the migratory flyway. This agrees with previous evidence on the role of migratory flyways in determining the distribution of some wintering birds in Iberia. A gap analysis reflected that just 12.3% of the suitable areas and 18.8% of individuals recorded during the 2012–2013 winter were included within the Special Protection Areas network of Portugal and Spain. Most of these birds were crowded in unprotected areas covered by rice fields (68% of individuals), a key habitat for the species.     

Wing morphology,winter ecology,and fecundity selection: evidence for sex-dependence in barn swallows (<Emphasis Type="Italic">Hirundo rustica</Emphasis>)

Variation in wing morphology results from the combination of diverse selection pressures. Wing feather morphology within species varies with sex and ontogenetic effects, and also with ecological factors. Yet, the direction of causation for the wing morphology–ecology association remains to be elucidated. Under the ‘ecology-dependence’ hypothesis, wing morphology covaries with ecological conditions, because the latter affect feather molt. Alternatively, the ‘habitat choice’ hypothesis posits that individuals with different wing morphology choose different habitats because of the habitat-dependent advantages of a specific wing morphology. We tested these competing hypotheses in the migratory, aerially insectivorous barn swallow (Hirundo rustica). We quantified wing morphology (isometric size, pointedness, and convexity) on the same individuals during consecutive breeding seasons (i.e., before and after molt in sub-Saharan wintering areas) and located wintering areas using light-level geolocators. Wing pointedness of females but not males during 1 year negatively correlated with vegetation vigor (gauged by the Normalized Difference Vegetation Index; NDVI) in the African area where individuals spent the next winter. Partial least-squares path modelling showed that the association between wing morphology and NDVI was sex-dependent. Conversely, NDVI during wintering did not predict wing morphology in the next breeding season. Because wing morphology can have carry-over effects on subsequent performance, we investigated selection on wing traits and found strong positive fecundity selection on wing size of females. Our results suggest that female barn swallows choose their wintering habitat depending on their wing morphology. In addition, directional fecundity selection operates on females, suggesting sex-dependence of current selection on the flight apparatus.     

Identification of current ecosystem functional types in the Iberian Peninsula

Aim To examine the geographical patterns of the interception of photosynthetically active radiation by vegetation and to describe its spatial heterogeneity through the definition of ecosystem functional types (EFTs) based on the annual dynamics of the Normalized Difference Vegetation Index (NDVI), a spectral index related to carbon gains. Location The Iberian Peninsula. Methods EFTs were derived from three attributes of the NDVI obtained from NOAA/AVHRR sensors: the annual integral (NDVI‐I), as a surrogate of primary production, an integrative indicator of ecosystem functioning; and the intra‐annual relative range (RREL) and month of maximum NDVI (MMAX), which represent key features of seasonality. Results NDVI‐I decreased south‐eastwards. The highest values were observed in the Eurosiberian Region and in the highest Mediterranean ranges. Low values occurred in inner plains, river basins and in the southeast. The Eurosiberian Region and Mediterranean mountains presented the lowest RREL, while Eurosiberian peaks, river basins, inner‐agricultural plains, wetlands and the southeastern part of Iberia presented the highest. Eurosiberian ecosystems showed a summer maximum of NDVI, as did high mountains, wetlands and irrigated areas in the Mediterranean Region. Mediterranean mountains had autumn–early‐winter maxima, while semi‐arid zones, river basins and continental plains had spring maxima. Based on the behaviour in the functional traits, 49 EFTs were defined. Main conclusions The classification, based on only the NDVI dynamics, represents the spatial heterogeneity in ecosystem functioning by means of the interception of radiation by vegetation in the Iberian Peninsula. The patterns of the NDVI attributes may be used as a reference in evaluating the impacts of environmental changes. Iberia had a high spatial variability: except for biophysically impossible combinations (high NDVI‐I and high seasonality), almost any pattern of seasonal dynamics of radiation interception was represented in the Peninsula. The approach used to define EFTs opens the possibility of monitoring and comparing ecosystem functioning through time.     

Spring arrival response to climate change in birds: a case study from eastern Europe

This paper analyses the dependence of the first spring arrival dates of short/medium- and long-distance migrant bird species on climate warming in eastern Europe. The timing of arrival of the selected species at the observation site correlates with the North Atlantic Oscillation (NAO) index, air temperature, atmospheric pressure, precipitation and wind characteristics. A positive correlation of fluctuations in winter and spring air temperatures with variations in the NAO index has been established in eastern Europe. Positive winter NAO index values are related to earlier spring arrival of birds in the eastern Baltic region and vice versa—arrival is late when the NAO index is negative. The impact of climate warming on the bird’s life cycle depends on local or regional climate characteristics. We tested the hypothesis that differences in climate indices between North Africa and Europe can influence the timing of spring arrival. Our results support the hypothesis that differences in first spring arrival dates between European populations occur after individuals cross the Sahara. We assume that the endogenous programme of migration control in short/medium-distance migrants synchronises with the changing environment on their wintering grounds and along their migration routes, whereas in long-distance migrants it is rather with environmental changes in the second part of their migratory route in Europe. Our results strongly indicate that the mechanism of dynamic balance in the interaction between the endogenous regulatory programme and environmental factors determines the pattern of spring arrival, as well as migration timing.     

The importance of traditional farmland areas for steppe birds: a case study of migrant female Great Bustards Otis tarda in Spain

A detailed knowledge of the habitat requirements of steppe birds living in farmland habitats is necessary to identify agricultural practices compatible with their conservation. The globally threatened Great Bustard Otis tarda is a partial migrant in central Iberia, but factors affecting its winter habitat use have not been identified. We assessed habitat differences between breeding and wintering areas and winter habitat selection of radiotagged migrant female Great Bustards in central Spain. Of 68 tagged females, 35% moved to wintering areas located 64.3 ± 24.0 km south of their breeding areas, and 80% wintered in a single area of c. 236 km². A census of the population in this area identified it as one of the most important wintering areas of this species in the world, holding c. 1500 individuals. There were significant differences between breeding and wintering habitats of individually marked migrant females. Compared with breeding areas, wintering areas of migrant females were located further from roads and urban nuclei, had lower human population densities and area of urban developments, and a higher diversity of land‐use types, with less cover of cereals and more vineyards and olive groves. Within this area, radiotracked migrant females preferred sites with more vineyards and a lower land‐use diversity. Our results highlight the importance of traditional Mediterranean dry farmland mosaics, and suggest that different conservation strategies are needed for migrant and resident populations in winter to secure the conservation of suitable wintering habitat for Great Bustards in the Iberian Peninsula.