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.     

Changes in the timing of spring and autumn migration in North American migrant passerines during a period of global warming

Butler (2003) used first arrival dates (FADs) of 103 migrant birds in northeastern USA and found that both long-distance migrants (LDMs; wintering south of the USA) and short-distance migrants (SDMs; wintering in the southern USA) arrived earlier in the second half of the 20th century than they had in the first, consistent with scenarios of global warming; the trend was stronger in SDMs. Using FADs to characterize migration systems can be problematic because they are data from one tail of a distribution, they comprise a mostly male population and they may not correlate well with the balance of the migration period. FADs also provide no information about autumn migration. This paper uses a banding dataset from Long Point Bird Observatory, Ontario, for 14 passerines for a period of global warming (1975–2000), taking these issues into account. The data were filtered to minimize effects of unequal netting effort (147 491 resulting records), and the passage dates then calculated in each season of each year for the 1st, 2nd and 3rd quartiles for regression analysis. Only two of 13 species analysed in the spring showed significantly earlier passage times, although the overall trend was towards earlier spring migration, especially among SDMs. Autumn responses were more prevalent, however, and in some cases more dramatic with six of 13 species showing delayed migration (four SDMs, two LDMs). Two LDMs exhibited earlier autumn migration. Where earlier spring migration occurred, both sexes appeared to contribute to the change. Where delayed migration occurred in autumn, both sexes and both adults and hatch-year birds appeared to contribute in at least some cases. The spring FAD results are consistent with those of Butler, but when the whole migration is considered, change is far from universal in spring and is in fact more substantial and complex in autumn.     

Protandry, sexual selection and climate change

Protandry refers to the earlier appearance of males before females at sites of reproduction. Sexual selection has been hypothesized to give rise to sex differences in benefits and costs of early arrival, thereby selecting for earlier appearance by the sex subject to more intense sexual selection. If sexual selection is more intense, there is a greater premium on early arrival among individuals of the chosen sex because of direct selection for earlier arrival. This hypothesis leads to the prediction that changes in the costs and benefits of early arrival related to changes in environmental conditions should particularly affect the sex that arrives first and hence the degree of protandry. I tested this hypothesis using the Barn Swallow Hirundo rustica. During 1971–2003, the degree of protandry increased significantly in a Danish population because males advanced arrival date while females did not. This earlier arrival by males compared with females was correlated with a significant increase by over 1.2 standard deviations in the length of the outermost tail feathers of males, a secondary sexual character, suggesting direct selection on both protandry and the secondary sexual character. Environmental conditions during spring migration in Northern Africa, as reflected by the normalized difference vegetation index, have deteriorated since 1984, resulting in increased mortality among males during spring migration, but not among females, and this deterioration of climatic conditions was positively correlated with an increasing degree of protandry. Likewise, an increase in April temperatures at the breeding grounds during recent decades is positively correlated with increased protandry, apparently because males can arrive earlier without increasing the fitness cost of early arrival. Local population size did not predict changes in arrival date. These findings suggest that rapid changes in climate can cause a change in degree of protandry and secondary sexual characters.     

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.     

Climatic effects on arrival and laying dates in a long-distance migrant, the Collared Flycatcher Ficedula albicollis

Long‐distance migrants may respond to climate change in breeding, wintering or staging area by changing their phenology. The geographical variation in such responses (e.g. coastal vs. continental Europe) and the relative importance of climate at different spatial scales remain unclear. Here we analysed variation in first arrival dates (FADs) and laying dates of the Collared Flycatcher Ficedula albicollis in a central European population, from 1973 to 2002. The North Atlantic Oscillation (NAO) index correlated weakly with local temperature during the laying period. Decreasing spring temperatures until 1980 were associated with a trend towards later laying. The rate of warming (0.2 °C per year) and laying advancement (0.4 days per year) since 1980 are amongst the highest values reported elsewhere. This long‐term trend in laying date was largely explained by the change in climatic factors. The negative effect of local spring temperature on laying was relatively stronger than that of NAO. The number of clutches initiated on a particular day was marginally affected by the temperature 3 days prior to laying and the response of females to daily variation in temperature did not change over years. Correspondence between the average population‐level and the individual‐level responses of laying date to climate variation suggests that the advancement of laying was due to phenotypic plasticity. Despite warmer springs and advanced laying, FADs did not change over years and were not correlated with local spring temperature. Marginal evidence suggests later departure from wintering grounds and faster migration across staging areas in warmer conditions. Advancement of arrival was probably constrained by low local temperatures in early spring just before arrival that have not changed over years. The interval between first arrival and laying has declined since 1980 (0.5 days per year), but the increasing temperature during that period may have kept the food supply approximately unchanged.     

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.     

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.     

Implications of climate change for North American wood warblers (Parulidae)

Since 1912, Neotropical–Nearctic migrant birds may now have up to 20 fewer days to travel between Southern Illinois, where spring is arriving later, and Northern Minnesota, where spring is arriving earlier, to exploit optimal habitat conditions (expanding leaves and caterpillar activity) for refueling and breeding. As case studies of the effect of climate change on bird migration, I analyzed two long‐term data sets of arrival times for eight species of northern breeding migratory wood warblers (Parulidae) gathered over a 100 year period in east‐central Illinois (IL, USA) and a 40 year period in western Minnesota (MN, USA). Six (IL) and seven (MN) of the wood warbler species showed no significant tendency to migrate earlier in response to earlier springs in their breeding range. These results suggest that climate change may force many species of long‐distance migratory songbirds to become uncoupled in the spring from their food resources that are driven by temperature.     

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.     

Avian population consequences of climate change are most severe for long-distance migrants in seasonal habitats

One consequence of climate change is an increasing mismatch between timing of food requirements and food availability. Such a mismatch is primarily expected in avian long-distance migrants because of their complex annual cycle, and in habitats with a seasonal food peak. Here we show that insectivorous long-distance migrant species in The Netherlands declined strongly (1984–2004) in forests, a habitat characterized by a short spring food peak, but that they did not decline in less seasonal marshes. Also, within generalist long-distance migrant species, populations declined more strongly in forests than in marshes. Forest-inhabiting migrant species arriving latest in spring declined most sharply, probably because their mismatch with the peak in food supply is greatest. Residents and short-distance migrants had non-declining populations in both habitats, suggesting that habitat quality did not deteriorate. Habitat-related differences in trends were most probably caused by climate change because at a European scale, long-distance migrants in forests declined more severely in western Europe, where springs have become considerably warmer, when compared with northern Europe, where temperatures during spring arrival and breeding have increased less. Our results suggest that trophic mismatches may have become a major cause for population declines in long-distance migrants in highly seasonal habitats.     

Spring migration timing of Sylvia warblers in Tatarstan (Russia) 1957–2008

The timing of when migrant birds return to breed is a key component of studies of the impact of climate change upon bird populations. However, such data are not distributed evenly across the World, and in the Northern Hemisphere are underrepresented in Asia and the east of Europe. Therefore, to help rectify this bias, we analysed first arrival dates (FAD) of four species of Sylvia warblers (Blackcap Sylvia atricapilla, Whitethroat S.communis, Lesser Whitethroat S.curruca and Garden Warbler S.borin) collected in the Tatarstan Republic of Russia between 1957 and 2008. Over the whole period the species returned to their breeding sites between three and six days earlier; these trends were significant except for Whitethroat. Advances in arrival were especially apparent in the two earlier species, Blackcap and Lesser Whitethroat, mainly because local temperatures for March had risen substantially. Except for Whitethroat, FADs were significantly related to temperatures in the African wintering ground and/or in Tatarstan. Whilst significant correlations occurred between FADs of some of the species, there was considerable variability in these relationships indicating a species-specific response to rising temperatures. Changes in FADs in this eastern extremity of Europe were smaller than in Central and Western Europe.     

The relative importance of conditions in wintering and passage areas on spring arrival dates: the case of long-distance Iberian migrants

Remote sensing data have been used in previous studies to assess the effects of winter ecological conditions in Africa on biological parameters recorded in bird populations during the following breeding season in Europe. Based on the results of these studies, we hypothesized that a high productivity of vegetation during the winter and, thus, high resource availability, should advance the arrival of long-distance migrants to the European breeding areas due to enhanced ecological conditions. To test this hypothesis, between 1982 and 2000 we examined the first arrival date to the Iberian Peninsula of five species (White Stork, Cuckoo, Common Swift, Barn Swallow and Nightingale) in relation to several explanatory variables: ecological conditions in their African wintering grounds and passage areas, as reflected by the normalized difference vegetation index (NDVI), temperature and precipitation in their passage areas and the winter North Atlantic Oscillation (NAO). Ecological conditions in the wintering areas were important for White Stork, Cuckoo and Barn Swallow phenology, while both NDVI in passage areas and NAO did not have an effect on any species. Migratory birds arrived earlier after winters with high vegetation productivity in Africa. Temperature in passage areas was important for the later species (i.e. Cuckoo, Common Swift and Nightingale), although in all cases the true relevance of this factor was scarce due to the poor explanatory capacity of the models. These species were recorded in the Iberian Peninsula earlier in the spring of those years with warmer temperatures in passage areas. The nexus between African NDVI and arrival phenology is hypothesized through increases in wintering survival rates and/or the faster acquisition of pre-migratory body condition and progression through sub-Saharan areas. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Temperature‐related increases in grass growth and greater competition for food drive earlier migrational departure of wintering Whooper Swans

Understanding the departure decisions of migratory birds is critical for determining how changing climatic conditions will influence subsequent arrival times on the breeding grounds. A long‐term dataset (1972–2008) of Whooper Swan Cygnus cygnus departure dates from a wintering site in Ireland was used to assess the factors determining the timing of migration. Early and late migrating swans showed different departure patterns. Earlier wintering ground departure was more pronounced for the first 50% of the population than the last 10% of departing individuals. Earlier departure was associated with an increase in February temperatures at the wintering site for all departure phases except the date when the last individual departed. The date by which the first 50% of Swans had departed was earlier with increasing numbers of wintering Swans, suggesting that competition on the wintering grounds may further influence the timing of departure. The results also suggested that departure is mediated by the influence of spring temperature on food resources, with increased February grass growth in warmer years enabling earlier departure of migrating Swans. To determine why arrival dates in the breeding ground have altered, environmental conditions in the wintering grounds must be taken into account.     

Multi‐decadal trends in spring arrival of avian migrants to the central Arctic coast of Alaska: effects of environmental and ecological factors

Warming in the Arctic has caused the transition from winter to summer to occur weeks earlier over the last half century, yet little is known about whether avian migrants have altered their timing of arrival on breeding areas to match this earlier seasonal transition. Over a 50‐yr period, we examined trends in the timing of the first arrival for 16 avian migrant species at the terminus of their northward migration along the central Arctic coast of Alaska and compared these trends to factors potentially influencing migration phenology. Date of first arrival occurred an average of 0.12 d yr^?1 or 6 d (range = 3–10 d) earlier across all species and did not differ significantly among species between 1964 and 2013. Local climatic variables, particularly temperature, had a greater effect on a species first arrival date than did large‐scale climatic predictors. First arrival date was 1.03 d earlier for every 1°C annual change in temperature, but there was nearly a 2‐fold difference in the range of responses across species (0.69–1.33 d °C^?1), implying that some species did better than others at timing their arrival with changing temperature. There was weak support for an influence of foraging strategy, migration distance, and flight path on timing of first arrival. Our findings, like others from temperate latitudes, indicate that avian migrants are responsive to changing environmental conditions, though some species appear to be more adaptive than others.     

Haemosporidian parasitism in the blackcap Sylvia atricapilla in relation to spring arrival and body condition

Parasites can exert strong selection on hosts. Throughout the year migrants are exposed to different sets of parasites, which may affect life history traits such as migratory schedules. Here, we studied the relationship between parasite infection and arrival date of blackcaps Sylvia atricapilla to their breeding grounds in Germany throughout a period of six years (2007–2012). We used two data sets, one that included all blackcaps and one that included only recaptured birds. We assesed whether parasites influence spring arrival to breeding grounds, and for the recaptured data set, we analysed temporal variation in parasitism (i.e. infection status and parasitaemia) throughout the breeding season. We used both microscopy and PCR (a fragment of ? 479 bp of the mtDNA cyt b) to determine haemosporidian infection. Blackcaps were mostly infected with Haemoproteus parabelopolskyi (lineages SYAT01 and SYAT02). Infection status, but not parasitaemia, was constant through time for individual birds; meaning that once a bird is infected, it most likely will retain the infection for life. We found that infection by haemosporidian parasites has no relationship to arrival date in this blackcap population; however, infection by H. parabelopolskyi has a marginally significant effect on arrival date of recaptured blackcaps, somewhat delaying their arrival to breeding grounds. Birds captured later in the season were more likely to be infected than those from early spring, and parasitaemia was frequently lower in birds captured earlier in the season compared to those captured later (summer).     

Variation in climate warming along the migration route uncouples arrival and breeding dates

Migratory species are of special concern in the face of global climate change, since they may be affected by changes in the wintering area, along the migration route and at the breeding grounds. Here we show that migration and breeding times of a trans‐Saharan migrant, the pied flycatcher Ficedula hypoleuca, closely follow local temperatures along the migration route and at the breeding grounds. Because of differences in long‐term temperature trends of short within‐spring periods, the migration period and the time interval between migration and breeding dates of this species have extended in SW Finland. Temperatures in northern parts of Central Europe have risen at the time when the first migrants arrive there, facilitating their migration northward. Temperatures later in the spring have not changed, and the last individuals arrive at the same time as before. The timing of breeding has not advanced because temperatures at the breeding site after arrival have not changed. These results show that the pied flycatchers can speed up their migration in response to rising temperatures along the migration route. Our results strongly indicate that the effects of climate change have to be studied at the appropriate time and geographical scales for each species and population concerned.     

Predation can select for later and more synchronous arrival times in migrating species

For migratory species, the timing of arrival at breeding grounds is an important determinant of fitness. Too early arrival at the breeding ground is associated with various costs, and we focus on one understudied cost: that migrants can experience a higher risk of predation if arriving earlier than the bulk of the breeding population. We show, using both a semi‐analytic and simulation model, that predation can select for later arrival. This is because of safety in numbers: predation risk becomes diluted if many other individuals, either con‐ or heterospecific, are already residing in the area. Predation risk dilution can also select for more synchronous arrival because deviating from the current population‐wide norm to earlier or later dates leads to higher predation risk or to failures in territory acquisition, respectively. The fact that selection for high arrival synchrony can in some cases be more important than selection for a specific date (early or late) within the season is an example of an ‘evolutionary priority effect’: whichever strategy – in this case a particular arrival time – becomes established in a population can remain stable over long periods of time; there are many possible equilibria (multiple stable states) which the population can remain at. Mixed arrival strategies are also possible under some circumstances.     

Protandrous migration and variation in morphological characters in Emberiza buntings at an East Asian stopover site

The effect of the timing of spring migration on reproductive success differs between the sexes. As a consequence, various sex‐specific tactics relating to the timing of migration have evolved in migratory avian groups. Various hypotheses have been proposed to explain differential migration to breeding or wintering grounds, and inter‐ and intrasexual size differences are often considered one of the proximate mechanisms. We investigated arrival patterns in the spring by individuals of each sex, sexual size dimorphism and related morphological variables, and the relationship between size variation and arrival date in five bunting species that passed through an East Asian migratory flyway stopover site in 2006–08. Males of all the study species arrived before females, and significant sexual dimorphism was observed. Several morphological characters, including total length, wing‐length and tail‐length, contributed to the size variation. Although larger males arrived earlier, there was no relationship between arrival date and size in females. Our study confirmed that East Asian buntings display a discriminated protandrous migration pattern at the stopover site as well as at the breeding grounds. This is consistent with the view that larger body size in males is favoured due to its association with early arrival to help ensure access to the best resources and hence enhanced mating success.     

Overtaking on migration: does longer distance migration always incur a penalty?

For many migratory bird species, the latitudinal range of the winter distribution spans thousands of kilometres, thus encompassing considerable variation in individual migration distances. Pressure to winter near breeding areas is thought to be a strong driver of the evolution of migration patterns, as individuals undertaking a shorter migration are generally considered to benefit from earlier arrival on the breeding grounds. However, the influence of migration distance on timing of arrival is difficult to quantify because of the large scales over which individuals must be tracked. Using a unique dataset of individually‐marked Icelandic black‐tailed godwits Limosa limosa islandica tracked throughout the migratory range by a network of hundreds of volunteer observers, we quantify the consequences of migrating different distances for the use of stop‐over sites and timing of arrival in Iceland. Modelling of potential flight distances and tracking of individuals from across the winter range shows that individuals wintering further from the breeding grounds must undertake a stop‐over during spring migration. However, despite travelling twice the distance and undertaking a stop‐over, individuals wintering furthest from the breeding grounds are able to overtake their conspecifics on spring migration and arrive earlier in Iceland. Wintering further from the breeding grounds can therefore be advantageous in migratory species, even when this requires the use of stop‐over sites which lengthen the migratory journey. As early arrival on breeding sites confers advantages for breeding success, the capacity of longer distance migrants to overtake conspecifics is likely to influence the fitness consequences of individual migration strategies. Variation in the quality of wintering and stopover sites throughout the range can therefore outweigh the benefits of wintering close to the breeding grounds, and may be a primary driver of the evolution of specific migration routes and patterns.