A matter of timing: changes in the first date of arrival and last date of departure of Australian migratory birds

Although there is substantial evidence that Northern Hemisphere species have responded to climatic change over the last few decades, there is little documented evidence that Southern Hemisphere species have responded in the same way. Here, we report that Australian migratory birds have undergone changes in the first arrival date (FAD) and last date of departure (LDD) of a similar magnitude as species from the Northern Hemisphere. We compiled data on arrival and departure of migratory birds in south‐east Australia since 1960 from the published literature, Bird Observer Reports, and personal observations from bird watchers. Data on the FAD for 24 species and the LDD for 12 species were analyzed. Sixteen species were short‐ to middle‐distance species arriving at their breeding grounds, seven were long‐distance migrants arriving at their nonbreeding grounds, and one was a middle‐distance migrant also arriving at its nonbreeding ground. For 12 species, we gathered data from more than one location, enabling us to assess the consistency of intraspecific trends at different locations. Regressions of climate variables against year show that across south‐east Australia average annual maximum and minimum temperatures have increased by 0.17°C and 0.13°C decade^?1 since 1960, respectively. Over this period there has been an average advance in arrival of 3.5 days decade^?1; 16 of the 45 time‐series (representing 12 of the 24 species studied) showed a significant trend toward earlier arrival, while only one time‐series showed a significant delay. Conversely, there has been an average delay in departure of 5.1 days decade^?1; four of the 21 departure time‐series (four species) showed a significant trend toward later departure, while one species showed a significant trend toward earlier departure. However, differences emerge between the arrival and departure of short‐ to middle‐distance species visiting south‐east Australia to breed compared with long‐distance species that spend their nonbreeding period here. On average, short‐ to middle‐distance migrants have arrived at their breeding grounds 3.1 days decade^?1 earlier and delayed departure by 8.1 days decade^?1, thus extending the time spent in their breeding grounds by ～11 days decade^?1. The average advance in arrival at the nonbreeding grounds of long‐distance migrants is 6.8 days decade^?1. These species, however, have also advanced departure by an average of 6.9 days decade^?1. Hence, the length of stay has not changed but rather, the timing of events has advanced. The patterns of change in FAD and LDD of Australian migratory birds are of a similar magnitude to changes undergone by Northern Hemisphere species, and add further evidence that the modest warming experienced over the past few decades has already had significant biological impacts on a global scale.     

Niche partitioning of two Palearctic passerine migrants with Afrotropical residents in their West African winter quarters

In bird migration systems, the question of coexistence and competitionof migrants with residents in the nonbreeding season and theirrole in shaping the evolution of present avian communities ismuch debated. However, conclusions are often drawn in a speculativeway. In the Palearctic-African bird migration system, many studieshave addressed the question of coexistence of Palearctic breedingbirds with Afrotropical species in the former's wintering grounds.These studies have led to some generalizations concerning habitatselection and foraging ecology about the traits that might enablemigrants to coexist with residents. Migrants were thereforeassumed to forage in more open habitats, in more peripheralparts of the vegetation and with a higher foraging speed thanresidents. Furthermore, they were also assumed to be more flexiblein foraging behavior by using a wider range of foraging tactics,but some studies revealed contradicting results. We studiedthe ecology of Palearctic migrants in Comoé NationalPark, Ivory Coast, West Africa, during three successive wintersto explore the factors of niche partitioning between migrantsand residents and to test the hypothesis of whether there arecommon behavioral traits in migrants. Therefore, we comparedthe ecology of two Palearctic breeding species: pied flycatcher,Ficedula hypoleuca, and willow warbler, Phylloscopus trochilus,with resident members of the respective guilds. With respectto use of microhabitat, foraging speed, and intake rates, wecould not confirm the above-mentioned generalizations, showingthat care has to be taken into account when drawing conclusionsfrom few studies for a whole migration system on a huge continent.However, both migrating species were more flexible in foragingbehavior than were their Afrotropical counterparts. As thereare hints that this is also the case when migrants are comparedwith residents on their breeding grounds, we suggest that thisflexibility enables migrants to partition resources with residentsand, therefore, coexist with Afrotropical species. We discuss,however, whether this flexibility is an adaptation to migratorybehavior or a prerequisite for the evolution of migration. Therole that competition plays in present communities cannot besolved with a few observational studies because of the followingproblems. First, it is difficult to detect competition in thefield. Second, there are constraints of performing field experimentsthat have not previously been performed in Africa, and third,there are several possible hypothetical scenarios about therole of competition in shaping present communities, includingfactors that might have been important in the past and are thereforeimpossible to detect at present.     

Novel methods reveal shifts in migration phenology of barn swallows in South Africa

Many migratory bird species, including the barn swallow (Hirundo rustica), have advanced their arrival date at Northern Hemisphere breeding grounds, showing a clear biotic response to recent climate change. Earlier arrival helps maintain their synchrony with earlier springs, but little is known about the associated changes in phenology at their non-breeding grounds. Here, we examine the phenology of barn swallows in South Africa, where a large proportion of the northern European breeding population spends its non-breeding season. Using novel analytical methods based on bird atlas data, we show that swallows first arrive in the northern parts of the country and gradually appear further south. On their north-bound journey, they leave South Africa rapidly, resulting in mean stopover durations of 140 days in the south and 180 days in the north. We found that swallows are now leaving northern parts of South Africa 8 days earlier than they did 20 years ago, and so shortened their stay in areas where they previously stayed the longest. By contrast, they did not shorten their stopover in other parts of South Africa, leading to a more synchronized departure across the country. Departure was related to environmental variability, measured through the Southern Oscillation Index. Our results suggest that these birds gain their extended breeding season in Europe partly by leaving South Africa earlier, and thus add to scarce evidence for phenology shifts in the Southern Hemisphere.     

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.     

Time of emergence of novel climates for North American migratory bird populations

To better understand the ecological implications of global climate change for species that display geographically and seasonally dynamic life‐history strategies, we need to determine where and when novel climates are projected to first emerge. Here, we use a multivariate approach to estimate time of emergence (ToE) of novel climates based on three climate variables (precipitation, minimum and maximum temperature) at a weekly temporal resolution within the Western Hemisphere over a 280‐yr period (2021–2300) under a high emissions scenario (RCP8.5). We intersect ToE estimates with weekly estimates of relative abundance for 77 passerine bird species that migrate between temperate breeding grounds in North America and southern tropical and subtropical wintering grounds using observations from the eBird citizen‐science database. During the non‐breeding season, migrants that winter within the tropics are projected to encounter novel climates during the second half of this century. Migrants that winter in the subtropics are projected to encounter novel climates during the first half of the next century. During the beginning of the breeding season, migrants on their temperate breeding grounds are projected to encounter novel climates during the first half of the next century. During the end of the breeding season, migrants are projected to encounter novel climates during the second half of this century. Thus, novel climates will first emerge ca 40–50 yr earlier during the second half of the breeding season. These results emphasize the large seasonal and spatial variation in the formation of novel climates, and the pronounced challenges migratory birds are likely to encounter during this century, especially on their tropical wintering grounds and during the transition from breeding to migration. When assessing the ecological implications of climate change, our findings emphasize the value of applying a full annual cycle perspective using standardized metrics that promote comparisons across space and time.     

Long‐term changes in autumn migration dates at the Strait of Gibraltar reflect population trends of soaring birds

A growing body of work shows that climate change is the cause of a number of directional shifts in the spring phenology of migratory birds. However, changes in autumn phenology are well studied and their consistency across species, as well as their link with population trends, remains uncertain. We investigate changes in the autumn migration dates of 11 species of soaring birds over the Strait of Gibraltar over a 16‐year period. Using models corrected for phylogeny, we assessed whether ecological and morphological characteristics, as well as population trends, account for interspecific shifts in migration times. We recorded different phenological changes in different periods of the migration season and suggest that these differences are due to age‐dependent responses. The variable best predicting advances in migration dates was population trend: species that did not advance their autumn migration dates were those showing a decline in their European breeding populations. We repeated our tests on a dataset representing the migration date of soaring birds across the Pyrenees Mountains and found that population trends at this site also predicted phenological shifts. Our results suggest that flexibility in migratory strategy and population trends may be related, such that different adaptive capacity in migration timing may be more relevant than other ecological traits in determining the conservation status of migratory birds in Europe and perhaps other regions.     

Why fly the extra mile? Latitudinal trend in migratory fuel deposition rate as driver of trans‐equatorial long‐distance migration

Trans‐equatorial long‐distance migrations of high‐latitude breeding animals have been attributed to narrow ecological niche widths. We suggest an alternative hypothesis postulating that trans‐equatorial migrations result from a possible increase in the rate at which body stores to fuel migration are deposited with absolute latitude; that is, longer, migrations away from the breeding grounds surpassing the equator may actually enhance fueling rates on the nonbreeding grounds and therewith the chance of a successful, speedy and timely migration back to the breeding grounds. To this end, we first sought to confirm the existence of a latitudinal trend in fuel deposition rate in a global data set of free‐living migratory shorebirds and investigated the potential factors causing this trend. We next tested two predictions on how this trend is expected to impact the migratory itineraries on northward migration under the time‐minimization hypothesis, using 56 tracks of high‐latitude breeding shorebirds migrating along the East Asian‐Australasian Flyway. We found a strong positive effect of latitude on fuel deposition rate, which most likely relates to latitudinal variations in primary productivity and available daily foraging time. We next confirmed the resulting predictions that (1) when flying from a stopover site toward the equator, migrants use long jumps that will take them to an equivalent or higher latitude at the opposite hemisphere; and (2) that from here onward, migrants will use small steps, basically fueling only enough to make it to the next suitable staging site. These findings may explain why migrants migrate “the extra mile” across the equator during the nonbreeding season in search of better fueling conditions, ultimately providing secure and fast return migrations to the breeding grounds in the opposite hemisphere.     

Do changes in climate patterns in wintering areas affect the timing of the spring arrival of trans‐Saharan migrant birds?

The life cycles of plants and animals are changing around the world in line with the predictions originated from hypotheses concerning the impact of global warming and climate change on biological systems. Commonly, the search for ecological mechanisms behind the observed changes in bird phenology has focused on the analysis of climatic patterns from the species breeding grounds. However, the ecology of bird migration suggests that the spring arrival of long‐distance migrants (such as trans‐Saharan birds) is more likely to be influenced by climate conditions in wintering areas given their direct impact on the onset of migration and its progression. We tested this hypothesis by analysing the first arrival dates (FADs) of six trans‐Saharan migrants (cuckoo Cuculus canorus, swift Apus apus, hoopoe Upupa epops, swallow Hirundo rustica, house martin Delichon urbica and nightingale Luscinia megarhynchos), in a western Mediterranean area since from 1952 to 2003. By means of multiple regression analyses, FADs were analysed in relation to the monthly temperature and precipitation patterns of five African climatic regions south of the Sahara where species are thought to overwinter and from the European site from where FADs were collected. We obtained significant models for five species explaining 9–41% of the variation in FADs. The interpretation of the models suggests that: (1) The climate in wintering quarters, especially the precipitation, has a stronger influence on FADs than that in the species' potential European breeding grounds. (2) The accumulative effects of climate patterns prior to migration onset may be of considerable importance since those climate variables that served to summarize climate patterns 12 months prior to the onset of migration were selected by final models. (3) Temperature and precipitation in African regions are likely to affect departure decision in the species studied through their indirect effects on food availability and the build‐up of reserves for migration. Our results concerning the factors that affect the arrival times of trans‐Saharan migrants indicate that the effects of climate change are more complex than previously suggested, and that these effects might have an interacting impact on species ecology, for example by reversing ecological pressures during species' life cycles.     

A rare study from the wintering grounds provides insight into the costs of malaria infection for migratory birds

Malaria parasites can have strong effects on the population dynamics and evolution of migratory bird species. In many species, parasite transmission occurs on the wintering grounds, but studies to determine the consequences of infection have taken place during the breeding season, when malaria parasites circulate at chronic levels. We examined the predictors of malarial infections for great reed warblers during the northern winter in Africa, where active parasite transmission is thought to occur and naïve individuals experience acute infections. Counter to expectations, we found that winter infection intensities were lower than those encountered on the breeding grounds. One potential explanation is that reduced immune function during breeding allows parasites to persist at higher chronic intensities. We found no relationships between the incidence or intensity of infection on condition (as measured by scaled mass index, plasma metabolites, and feather corticosterone), spring migration departure dates, or home range sizes. We also tested a prediction of the Hamilton–Zuk hypothesis and found that male ornament (song) quality was unrelated to parasitic infection status. Overall, our results provide the first evidence that long‐distance migrants captured on their wintering grounds are in the chronic stage of infection, and suggest that winter studies may fare no better than breeding studies at determining the costs of acute malarial infection for great reed warblers.     

Climate variability and the timing of spring raptor migration in eastern North America

Many birds have advanced their spring migration and breeding phenology in response to climate change, yet some long‐distance migrants appear constrained in their adjustments. In addition, bird species with long generation times and those in higher trophic positions may also be less able to track climate‐induced shifts in food availability. Migratory birds of prey may therefore be particularly vulnerable to climate change because: 1) most are long‐lived and have relatively low reproductive capacity, 2) many feed predominately on insectivorous passerines, and 3) several undertake annual migrations totaling tens of thousands of kilometers. Using multi‐decadal datasets for 14 raptor species observed at six sites across the Great Lakes region of North America, we detected phenological shifts in spring migration consistent with decadal climatic oscillations and global climate change. While the North Atlantic and El Niño Southern Oscillations exerted heterogeneous effects on the phenology of a few species, arrival dates more generally advanced by 1.18 d per decade, a pattern consistent with the effects of global climate change. After accounting for heterogeneity across observation sites, five of the 10 most abundant species advanced the bulk of their spring migration phenology. Contrary to expectations, we found that long‐distance migrants and birds with longer generation times tended to make the greatest advancements to their spring migration. Such results may indicate that phenotypic plasticity can facilitate climatic responses among these long‐lived predators.     

Intra‐African movements of the African cuckoo Cuculus gularis as revealed by satellite telemetry

Despite many bird species migrating regularly within the African continent, in response to rainfall and breeding opportunities, documented evidence of the spatiotemporal patterns of such movements is scarce. We use satellite telemetry to document the year round movement of an intra‐African migrant breeding in the savannah zone of sub‐Saharan Africa, the African cuckoo. After breeding in central Nigeria, the birds migrated to more forested sites in the Adamawa region of Cameroon (n = 2) and western Central African Republic (n = 1). Departure from the breeding ground coincided with deteriorating environmental conditions whereas arrival at the non‐breeding sites matched period of increasing vegetation greenness. Migratory movements generally occurred during dark hours. In total, an average distance of 748 km in 66 d was covered during the post‐breeding migration and 744 km in 27 d during return journey with considerable individual variation and with more stopover sites used during post‐breeding migration. The diversity of migration routes followed suggests a relatively variable or flexible initial migration strategy, high individual route consistency as well as high fidelity for non‐breeding grounds.     

Birds on the move in the face of climate change: High species turnover in northern Europe

Species richness is predicted to increase in the northern latitudes in the warming climate due to ranges of many southern species expanding northwards. We studied changes in the composition of the whole avifauna and in bird species richness in a period of already warming climate in Finland (in northern Europe) covering 1,100 km in south–north gradient across the boreal zone (over 300,000 km²). We compared bird species richness and species‐specific changes (for all 235 bird species that occur in Finland) in range size (number of squares occupied) and range shifts (measured as median of area of occupancy) based on bird atlas studies between 1974–1989 and 2006–2010. In addition, we tested how the habitat preference and migration strategy of species explain species‐specific variation in the change of the range size. The study was carried out in 10 km squares with similar research intensity in both time periods. The species richness did not change significantly between the two time periods. The composition of the bird fauna, however, changed considerably with 37.0% of species showing an increase and 34.9% a decrease in the numbers of occupied squares, that is, about equal number of species gained and lost their range. Altogether 95.7% of all species (225/235) showed changes either in the numbers of occupied squares or they experienced a range shift (or both). The range size of archipelago birds increased and long‐distance migrants declined significantly. Range loss observed in long‐distance migrants is in line with the observed population declines of long‐distance migrants in the whole Europe. The results show that there is an ongoing considerable species turnover due to climate change and due to land use and other direct human influence. High bird species turnover observed in northern Europe may also affect the functional diversity of species communities.     

Independent trophic behavior and breeding success of a resident flycatcher and a coexisting migratory congener

Historical biases towards the study of Palearctic and Nearctic bird migration systems has resulted in a dearth of information on the ecology of intratropical migrants, which likely also play important ecological roles within their communities. For instance, there is little information on the foraging ecology and breeding biology of co‐occurring migrants and residents within the intratropical migratory system. Thus, we used two congeneric flycatchers – the resident Plain‐crested Elaenia Elaenia cristata (Aves: Tyrannidae), and the migrant Lesser Elaenia E. chiriquensis (Tyrannidae) – to check whether either of them showed foraging niche plasticity mediating their coexistence and the effects of such syntopy on their breeding behavior and fitness. To do so, we used an ensemble of methods that included the evaluation of food resource phenology, foraging behavior observations, stable isotope ratio analysis in birds’ tissues, and nest monitoring. Our results confirmed that residents have foraging behavior plasticity but reveals its trophic behavior and breeding success is independent of the coexistence with migrants. Since such results depart from the predictions based on the current (and often Northern‐biased) theories of bird migration, we discuss alternative hypotheses explaining our findings, including the influence of physiology on diet and the behavioral responses of nest predators. Once there is a clear knowledge gap about the interspecific interactions between residents and intratropical migrants, our study represents a simple, yet important, step towards understanding the mechanisms underlying this system.     

Mechanisms of population dynamics in trans-Saharan migrant birds: A review

Modern views on the mechanisms of population dynamics in Palearctic birds wintering in tropical Africa are discussed with regard to the distribution of migrants on the African continent and the influence of external factors on conditions of bird wintering. The data on long-term bird trapping at ten ornithological stations in Europe are analyzed. Among 142 long-term trends in the population dynamics of 18 bird species, 34% are negative, 11% are positive, and the remaining trends lack statistical significance. The proportion of negative trends in each species negatively correlates with the number of birds: the more abundant the species, the lower the probability of its long-term population decline. Population dynamics may largely be determined by environmental conditions in wintering regions. Population studies on ten Palearctic species showed that drastic population declines and even local extinction of birds in recent decades resulted from the dramatic decrease in their survival on African wintering grounds because of insufficient precipitation and, in particular, severe droughts in the Sahel.     

Breeding phenological response to spring weather conditions in common Finnish birds: resident species respond stronger than migratory species

National bird‐nest record schemes provide a valuable data source to study large‐scale changes in basic breeding biology and effects of climate change on birds. Using nest‐record scheme data from 26 common Finnish breeding bird species from whole Finland, we estimated the laydate of the first egg for 129 063 nesting attempts. We then investigated the relationship of mean spring temperature and spring precipitation sum to changes in the onset of laying over the period 1961–2012. In addition, we examine differences in response to these climatic variables for species grouped for different life history strategies; migration, diet and habitat. Finally, we test whether body size is related to the strength of phenological response. We show that 26 common Finnish breeding bird species have advanced their laying dates over time and to an increase in the mean spring temperature over the study period. When species are grouped according life history strategies, we find that breeding phenological change is negatively associated with changes in the mean spring temperature where residents respond strongest to changes in mean spring temperature, but also short‐ and long‐distance migrants advance laydates with increasing spring temperatures. Breeding phenological change is also associated with spring precipitation, where resident species delay and short‐distance migrants advance the onset of breeding. In addition we find that omnivorous species respond stronger than insectivorous species to changes in spring temperature. In contrast to results from an earlier study, we do not find evidence that small‐sized species respond stronger to spring temperature than large‐sized species. As climate warming is predicted to continue in the future, long‐term citizen science schemes, such as the Finnish nest‐card scheme, prove to be a valuable cost‐effective way to monitor the environment and allow investigation into how species are responding to changes in their environment.     

Climatic connectivity between Africa and Europe may serve as a basis for phenotypic adjustment of migration schedules of trans-Saharan migratory birds

Most European migratory birds wintering in sub‐Saharan Africa have anticipated arrival to the breeding areas over the past decades. This phenological change may be ultimately caused by warming of the Northern Hemisphere via evolutionary changes or phenotypic plasticity in migration behavior. First arrival dates are negatively predicted by temperatures upon arrival to the breeding grounds. This seems puzzling, because migrants should be unable to predict weather conditions at long range. Migrants can enjoy diverse fitness benefits from early arriving. However, if weather conditions at destination cannot be predicted, early arrival can also entail severe costs. If meteorological conditions in Europe during breeding covary with those in sub‐Saharan Africa during late winter, long‐distance migrants may have a clue to predict meteorological conditions in their breeding areas while they are still in Africa and adjust their migration schedule consequently, an idea that has never been tested. We analyzed the correlation between March–April temperature anomalies (T_an) in Europe and February T_an in the Sahel and sub‐Sahel, where long‐distance migrants winter or stop‐over. T_an in Africa negatively predicted T_an in Europe, the association being particularly strong (unsigned effect size, z_r>0.35) for eastern Sahel and northern and eastern Europe, where the risks of early arrival may be larger. However, the strength of the correlations between T_an in the two continents has declined during the last 25 years; thus, possibly, partly compromising adaptive mechanisms of adjustment of migration. The existence of such climatic connectivity leads to several predictions, including that positive T_an in Africa should delay arrival. Consistent with this prediction, we found that first arrival dates of seven long‐distance migratory species positively covaried with February T_an in Africa. Thus, while wintering, migrants might be able to predict meteorological conditions at the beginning of the breeding season, and phenotypically adjust migration schedules to optimally tune arrival date.     

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.     

Long-term population declines in Danish trans-Saharan migrant birds

Capsule Long-distance migrant birds show less favourable trends than sedentary/short-distance species.

Aims To use breeding bird surveys to contrast population trends amongst common species according to their migration pattern.

Methods Changes in abundance of 62 Danish breeding sedentary, short-distance (Europe/North Africa) or long-distance (trans-Saharan) migrants were described by fitting log linear regression models to point-count census data gathered during 1976–2005.

Results Trans-Saharan migrants declined by 1.3% per annum during this period, while short-distance migrants and sedentary species increased by 1.4% and 1.0% per annum, respectively. There were no significant decadal declines amongst species using different summer breeding habitats, except for wetlands, and there was no consistent variation in trends associated with wintering regions or habitats or diet.

Conclusions More information is urgently needed on diet, feeding ecology, habitat requirements, winter distribution and intra-African movements of the commoner European summer visitors to identify causes of the declines and highlight when in the annual cycle detrimental effects occur. Studies linking individuals on their breeding, staging and wintering grounds are especially needed. Danish trends resemble those from elsewhere in Europe, confirming that restoration to favourable conservation status requires inter-continental action to meet European and global targets to reduce or halt biodiversity loss.     

Process in the evolution of bird migration and pattern in avian ecogeography

Current ideas about the evolution of bird migration equate its origin with the first appearance of fully migratory populations, and attribute its evolution to a selective advantage generated by increased breeding success, gained through temporary emigration from resident populations to breed in under-exploited seasonal areas. I propose an alternative hypothesis in which migration first appears as a temporary directional shift away from the breeding site outside the reproductive period, in response to seasonal variation in the direction and/or severity of environmental gradients. Fully migratory populations then appear through either extinction of sedentary phenotypes, or colonisation of vacant seasonal areas by migrants. Where colonisation occurs, resident ancestral populations can be driven to extinction by competition from migrants which invade their range outside the breeding season, resulting in fully migratory species. An analogous process drives the evolution of migration between high latitudes and the tropics, since extension of breeding range into higher latitudes may drive low latitude populations to extinction, resulting in an overall shift of breeding range. This process can explain reverse latitudinal gradients in avian diversity in the temperate zone, since the breeding ranges of migratory species concentrate in latitudes where they enjoy the highest breeding success. Near absence of forest-dwelling species among Palaearctic-African migrants is attributable to the lack of forest in northern Africa for much of the Tertiary, which has precluded selection both for southward extension of migration by west Palaearctic forest species, and northward breeding colonisation by African forest species.     

Repeatability of individual migration routes,wintering sites,and timing in a long‐distance migrant bird

Migratory birds are often faithful to wintering (nonbreeding) sites, and also migration timing is usually remarkably consistent, that is, highly repeatable. Spatiotemporal repeatability can be of advantage for multiple reasons, including familiarity with local resources and predators as well as avoiding the costs of finding a new place, for example, nesting grounds. However, when the environment is variable in space and time, variable site selection and timing might be more rewarding. To date, studies on spatial and temporal repeatability in short‐lived long‐distance migrants are scarce, most notably of first‐time and subsequent migrations. Here, we investigated repeatability in autumn migration directions, wintering sites, and annual migration timing in Hoopoes (Upupa epops), a long‐distance migrant, using repeated tracks of adult and first‐time migrants. Even though autumn migration directions were mostly the same, individual wintering sites often changed from year to year with distances between wintering sites exceeding 1,000 km. The timing of migration was repeatable within an individual during autumn, but not during spring migration. We suggest that Hoopoes respond to variable environmental conditions such as north–south shifts in rainfall during winter and differing onset of the food availability during spring migration.