Use of ring recoveries to predict habitat suitability in small passerines

Aim Abundance of small passerines may be distributed over large wintering areas according to geographical patterns that are difficult to detect. This may prevent the identification and conservation of the most suitable sectors, or the detection of local and regional features affecting the species during winter. In this paper, we explore the usefulness of ring recoveries to predict bird distribution in wintering grounds by using Maxent, one of the presence‐only techniques available for modelling species distributions. We test whether suitability indices obtained in 2.5 × 2.5 km UTM squares from ring recoveries were positively correlated with the actual abundance of robins (Erithacus rubecula) and blackcaps (Sylvia atricapilla) inside the squares. Location Spain. Methods We used 686 and 1139 localities across the country in which blackcap and robin ring recoveries were recorded by the Spanish Office of Migratory Species to generate the suitability maps. In addition, we sampled bird abundance inside 70 control squares independent of ring recovery localities along a belt crossing mountains, highlands and lowlands of the Iberian Peninsula during January from 2006 to 2011. Results Suitability indices predicted by Maxent were positively correlated with robin (r = 0.42, n = 70, P < 0.001) and blackcap (r = 0.52, n = 70, P < 0.001) abundances in the 70 control squares along the belt crossing the Iberian Peninsula. Main conclusions These results suggest that the use of distribution models with ring recoveries may be used to describe the habitat suitability of the winter ranges of small, common passerines. This also means that the huge number of ring recoveries stored in national and transnational data banks could be used to explore the factors shaping bird ranges and to forecast the geographical distribution of suitable wintering areas of migratory birds in large, poorly known regions. This may be useful in biogeography and conservation.     

Spring arrival along a migratory divide of sympatric blackcaps (Sylvia atricapilla)

The recent formation of a migratory divide in the blackcap (Sylvia atricapilla) involves sympatrically breeding birds migrating to different overwintering quarters. Within the last 50 years, a novel migratory strategy has evolved resulting in an increasing proportion of birds now wintering in Britain instead of migrating to the traditional sites in the Mediterranean area. This rapid microevolution has been attributed to allochronic spring arrival of migrants from the respective wintering quarters leading to assortative mating. Moreover, blackcaps wintering in Britain may experience fitness advantages owing to improved local wintering conditions. We used stable hydrogen isotope signatures (δD) to scrutinize the degree of temporal segregation of blackcaps upon spring arrival and to test for carry-over effects in body condition associated with the disparate wintering environments. Although we found that migrants from Britain arrive significantly earlier on German breeding grounds than migrants from the Mediterranean region, we also found a considerable overlap in arrival times. In a resampling model, the mean probability of assortative mating of birds wintering in Britain is ≤28% in both years. These results suggest that allochrony alone is not a strong isolating barrier between the two subpopulations. Migrants from both wintering locations did not differ in terms of body mass, mass-tarsus residuals or mass-tarsus ratio and arrived in a similar reproductive disposition. Thus, blackcaps wintering in Britain do not gain an apparent fitness advantage on spring migration due to carry-over effects in body condition. Future studies should explore additional factors such as differences in song quality and habitat that might contribute to the rapid microevolution of the blackcap.     

Spatial Isolation and Temporal Variation in Fitness and Condition Facilitate Divergence in a Migratory Divide

A novel migratory polymorphism evolved within the last 60 years in blackcaps (Sylvia atricapilla) breeding sympatrically in southwestern Germany. While most individuals winter in the traditional areas in the Mediterranean, a growing number of blackcaps started migrating to Britain instead. The rapid microevolution of this new strategy has been attributed to assortative mating and better physical condition of birds wintering in Britain. However, the isolating barriers as well as the physical condition of birds are not well known. In our study, we examined whether spatial isolation occurred among individuals with distinct migratory behaviour and birds with different arrival dates also differed in physical and genetic condition. We caught blackcaps in six consecutive years upon arrival on the breeding grounds and assigned them via stable isotope analysis to their wintering areas. Analysis of the vegetation structure within blackcap territories revealed different microhabitat preferences of birds migrating to distinct wintering areas. Blackcaps arriving early on the breeding grounds had higher survival rates, better body condition and higher multilocus heterozygosities than later arriving birds. We did however not find an effect of parasite infection status on arrival time. Our results suggest that early arriving birds have disproportionate effects on population dynamics. Allochrony and habitat isolation may thus act together to facilitate ongoing divergence in hybrid zones, and migratory divides in particular.     

Sex and age-specific differences in wing pointedness and wing length in blackcaps Sylvia atricapilla migrating through the southern Baltic coast

The blackcap Sylvia atricapilla shows a complex migratory pattern and is a suitable species for the studies of morphological migratory syndrome, including adaptations of wing shape to different migratory performance. Obligate migrants of this species that breed in northern, central, and Eastern Europe differ by migration distance and some cover shorter distance to the wintering grounds in the southern part of Europe/North Africa or the British Isles, although others migrate to sub-Saharan Africa. Based on ˃40 years of ringing data on blackcaps captured during autumn migration in the Southern Baltic region, we studied age- and sex-related correlations in wing pointedness and wing length of obligate blackcap migrants to understand the differences in migratory behavior of this species. Even though the recoveries of blackcaps were scarce, we reported some evidence that individuals which differ in migration distance differed also in wing length. We found that wing pointedness significantly increased with an increasing wing length of migrating birds, and adults had longer and more pointed wings than juvenile birds. This indicates stronger antipredator adaptation in juvenile blackcaps than selection on flight efficiency, which is particularly important during migration. Moreover, we documented more pronounced differences in wing length between adult and juvenile males and females. Such differences in wing length may enhance a faster speed of adult male blackcaps along the spring migration route and may be adaptive when taking into account climatic effects, which favor earlier arrival from migration to the breeding grounds.     

Habitat distribution of migratory and sedentary blackcaps Sylvia atricapilla wintering in southern Iberia: a morphological and biogeochemical approach

In migratory species, the way in which conspecifics from different breeding populations are distributed during the non‐breeding period is important from and ecological, evolutionary and conservation perspective, but such knowledge is still limited for most species. Migratory and sedentary blackcaps Sylvia atricapilla wintering in southern Spain can occupy two habitat types: forests and shrublands. According to earlier studies, blackcaps prefer forests over shrublands, and residents remain nearly restricted to forests. However, whether migrants with different breeding origin occupy the two habitats differently is unknown. We used morphological and biogeochemical data (hydrogen isotope ratios measured on feathers: δ²H_f), which show variation along the breeding range of the species, to answer this question. Isotope analyses supported the reliability of morphology as a method for distinguishing between migratory and sedentary blackcaps in sympatry, showing that sedentary individuals are rare in shrublands while migratory ones are abundant in both habitat types. However, migratory blackcaps scored similar δ²H_f values in forests and shrublands, and neither did vary in structural size or flight morphology between habitats. Our study suggests that migrants from a wide range of breeding origins end up mixing between forests and shrublands, which may explain the patterns of variation in space and time in the abundance of blackcaps in this area, and supports the view that inequalities may arise among migrants with the same origin but wintering in different habitats. Such inequalities might carry over into other stages of blackcaps’ life cycle contributing to the regulation of its migratory populations.     

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.     

Effects of habitat heterogeneity and local adaptation on the body condition of a forest passerine at the edge of its distributional range

We studied fluctuating asymmetry and feather growth rates as indicators of fitness of blackcaps ( Sylvia atricapilla ) breeding at the border of their distribution range in the Iberian Peninsula. Iberian blackcaps increase their abundance with increasing rainfall and ground cover of brambles. In border habitats, they are sedentary and morphologically different from migrants, suggesting that they could be adapted to peripheral conditions. We tested whether juvenile body condition depends on (1) distance from the centre of the range, (2) mean precipitation or (3) bramble cover. Controlling for environmental variation, we tested for differences between migratory and sedentary populations. Body condition varied across the Iberian gradient in parallel with changes in precipitation. Controlling for this effect, sedentary populations were in better body condition than migratory populations. Our results support the idea that environmental heterogeneity causes fitness to fluctuate across species' ranges, and also that local adaptation may mean that peripheral populations are more than a 'tail end' of the species.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 479–488.     

Regional variation in seasonality affects migratory behaviour and life-history traits of two Mediterranean passerines

Migratory birds may improve fecundity by moving to seasonal breeding areas, but may also suffer higher mortality rates as a cost of movement. However, the covariation among seasonality, migratoriness and life histories should change between species if their ecological features affect site-tenacity, survival or fecundity. In frugivorous birds, for instance, wandering in search of fruits may trigger broader migrations than territorial defence, and also may improve nonbreeding survival by preventing food shortages that eventually happen in discrete territories. We studied the variation in spatio-temporal distribution, life expectancy and fecundity in robins (Erithacus rubecula) and blackcaps (Sylvia atricapilla) distributed in three Iberian regions with a low, mid or high degree of environmental seasonality. In the Iberian Peninsula, robins and blackcaps are the two most intensive frugivores in winter; however, robins are territorial while blackcaps track fruit abundance among habitat patches. In the most seasonal area, robins and blackcaps decreased abundance in winter and showed a lower breeding-site tenacity, a shorter life expectancy and a larger clutch size. However, blackcaps tended to be more migratory than robins in all regions. Despite their stronger migratory behaviour, blackcaps showed a longer life expectancy and a smaller clutch size than robins in all regions. In robins, winter territoriality could decrease nonbreeding survival but also improve fecundity, because survivors are dominant and hence more efficient breeders. In blackcaps, however, the use of the most profitable habitat patches could improve nonbreeding survival, thereby allowing a similar recruitment than in robins with a lower reproductive investment. These results support that regional-scale changes in seasonality may affect migratory behaviour and hence the tradeoff between reproduction and survival in birds, doing so differently depending on the idiosyncrasy of each species.     

Is supplementary feeding in gardens a driver of evolutionary change in a migratory bird species?

Human activities are causing rapid environmental change at a global scale. Urbanization is responsible for some of the most extreme human‐altered habitats and is a known driver of evolutionary change, but evidence and understanding of these processes is limited. Here, we investigate the potential underlying mechanisms contributing to the contemporary evolution of migration behaviour in the Eurasian blackcap (Sylvia atricapilla). Blackcaps from central Europe have been wintering in urban areas of Britain with increasing frequency over the past 60 years, rather than migrating south to the Mediterranean. It has been hypothesized that the popularization of providing supplementary foods for wild birds within Britain may have influenced this marked migratory change, but quantifying the selective forces shaping evolutionary changes remains challenging. Using a long‐term national scale data set, we examine both the spatial distribution and interannual variation in blackcap wintering behaviour in Britain in relation to supplementary food availability and local climate. Over a 12‐year period, we show that blackcaps are becoming increasingly associated with the provision of supplementary foods in British gardens, and that the reliability of bird food supplies is influencing their winter distribution at a national scale. In addition, local climatic temperatures and broader scale weather variation are also important determinants of blackcap wintering patterns once they arrive in Britain. Based on our findings, we conclude that a synergistic effect of increased availability of feeding resources, in the form of garden bird food, coupled with climatic amelioration, has enabled a successful new wintering population to become established in Britain. As global biodiversity is threatened by human‐induced environmental change, this study presents new and timely evidence of the role human activities can play in shaping evolutionary trajectories.     

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.     

Habitat effects on resource tracking ability: do wintering Blackcaps Sylvia atricapilla track fruit availability?

If resource availability shapes population distribution, changes in resource abundance should cause parallel changes in population numbers. However, tracking ability may be disrupted by different environmental and behavioural factors that act at different spatial and temporal scales. Here we analyse the ability of wintering Blackcap Sylvia atricapilla populations to track spatio-temporal variation in fruit availability in southern Spain in two habitats (forests and shrublands) with different population structure. Former studies had shown that forests are equally used by both adult migrant and local Blackcaps, whereas shrublands are nearly monopolized by juvenile migrants. These differences might affect resource tracking: it should be disrupted in forests, as local birds remain over winter in their breeding territories, but not in shrublands where similarly competitive juvenile migrants can freely track the spatial distribution of fruits. We analysed the fruit-tracking ability of Blackcap populations among sites and years in both habitat types using a habitat-matching model, which predicts spatio-temporal changes in population abundance proportional to changes in resource availability. We counted Blackcaps and fruiting shrubs (dominated by Lentiscs Pistacia lentiscus and Wild Olives Olea europaea sylvestris ) during four winters in forest and shrubland patches. The abundance of fruits was always higher in shrublands than in forests. In shrublands, Blackcaps seemed to move freely across fruit-rich habitat patches, tracking changes in fruiting-shrub abundance among sites and years. However, such tracking was not observed in forests. This supports the view that fruit-tracking ability may be constrained by local factors, such as the social structure of populations occurring in different habitat types, which introduces spatio-temporal variation in the way fruit availability shapes the abundance distribution of these birds in their Mediterranean wintering grounds.     

Fruit tracking between sites and years by birds in Mediterranean wintering grounds

Knowledge of the ability of birds to track spatiotemporal variation in fruit distribution is essential for understanding plant-frugivore interactions. Arguably, although total fruit availability sets an upper limit to the number of birds that can exploit a habitat patch, not all species can equally distribute abundance according to variation in fruit resources. To explore this, we studied bird and fruit abundance in 1999–2005 in Mediterranean scrublands and woodlands of southern Spain. We analysed whether changes of fruit abundance in eight different sites during six winters could predict numerical changes of a set of frugivorous passerines of the area (blackcap Sylvia atricapilla , Sardinian warbler S. melanocephala , robin Erithacus rubecula , song thrush Turdus philomelos and blackbird T. merula ). We also investigated if all frugivores together tracked fruits better than individual species, thereby supporting a shared use of resources. Results showed strong inter-specific differences. Only the most abundant species (blackcaps and robins) tracked the spatial patterning of food despite strong differences in the use of space (vagrant and territorial, respectively). This suggests plastic behaviour of territorial robins, with individuals changing from strictly territorial to wandering, a flexibility that would favour between-site numerical arrangements according to food resources. Annual changes in bird numbers were independent of the availability of fruits, except for blackcaps, an abundant vagrant bird that tracked inter-winter changes in fruit abundance. The abundance of blackcaps fitted the spatiotemporal patterning of fruit resources better than the whole guild of frugivorous birds, inconsistent with the idea that these species track together the changing availability of fruit resources.     

Ecophysiological aspects of rapid population growth in a novel migratory blackcap (Sylvia atricapilla) population: an experimental approach

Summary Blackcaps (Sylvia atricapilla) that breed in central Europe have usually migrated to Mediterranean or African wintering grounds. In the past several decades, a portion of this breeding population has started migrating to the British Isles to overwinter and this population has increased dramatically. Several factors, including higher annual survivorship (due to supplemental feeding and reduced migratiry distance), assortative mating, and enhanced reproductive success may be involved in this rapid population growth. As part of an intensive, long-term study of this population, we tested the hypothesis that the differences in photoperiod experienced by British-wintering versus Mediterranean-wintering blackcaps might lead to relatively early vernal (i.e., migratory and/or reproductive) physiological condition in members of the former group. We found that birds exposed to photoperiodic conditions that simulated migration to Britain to overwinter generally initiated vernal migratory activity earlier than birds held under conditions simulating migration to traditional wintering areas in central Spain. This difference, coupled with the shorter migratory distance to the British Isles, leads to significantly earlier estimated arrival dates for blackcaps that winter in Britain compared to central Spain. Bimodality in arrival times suggests that assortative mating on central European breeding grounds might occur between members of the different wintering populations. Males exposed to British-winter photoperiods showed significantly earlier testicular development than males kept under Spanish-winter photoperiods. Early arrival on the breeding grounds, coupled with accelerated reproductive condition, should lead to a relatively early reproductive effort, perhaps increasing average reproductive success. In general, these results support the hypothesis that differences in photoperiod on the wintering grounds may play an important role in the dynamic state of this population.     

Wintering space use and site fidelity in a nomadic species,the snowy owl

Migratory species can exploit many habitats over vast geographic areas and adopt various patterns of space and habitat use throughout their annual cycle. In nomadic species, determinants of habitat use during the non‐breeding season are poorly known due to the unpredictability of their movement patterns. Here, we analysed variability in wintering space and habitat use by a highly nomadic species, the snowy owl, in eastern North America. Using 21 females tracked by satellite telemetry between 2007 and 2016, we 1) assessed how space use patterns in winter varied according to the type of environment (marine vs terrestrial), latitudinal zone (Arctic vs temperate), local snow conditions and lemming densities and 2) investigated winter habitat and site fidelity. Our results confirmed a high inter‐individual variation in patterns of habitat use by wintering snowy owls. Highly‐used areas were concentrated in the Arctic and in the marine and coastal environments. Owls wintering in the marine environment travelled over longer distances during the winter, had larger home ranges and these were divided in more smaller zones than individuals in terrestrial environments. Wintering home range sizes decreased with high winter lemming densities, use of the marine environment increased following high summer lemming densities, and a thick snow cover in autumn led to later settlement on the wintering ground. Contrary to expectations, snowy owls tended to make greater use of the marine environment when snow cover was thin. Snowy owls were highly consistent in their use of a given wintering environment and a specific latitudinal zone between years, but demonstrated flexibility in their space use and a modest site fidelity. The snowy owls’ consistency in wintering habitat use may provide them with advantages in terms of experience but their mobility and flexibility may help them to cope with changing environmental conditions at fine spatial scale.     

Individual variability and versatility in an eco-evolutionary model of avian migration

Seasonal migration is a complex and variable behaviour with the potential to promote reproductive isolation. In Eurasian blackcaps (Sylvia atricapilla), a migratory divide in central Europe separating populations with southwest (SW) and southeast (SE) autumn routes may facilitate isolation, and individuals using new wintering areas in Britain show divergence from Mediterranean winterers. We tracked 100 blackcaps in the wild to characterize these strategies. Blackcaps to the west and east of the divide used predominantly SW and SE directions, respectively, but close to the contact zone many individuals took intermediate (S) routes. At 14.0° E, we documented a sharp transition from SW to SE migratory directions across only 27 (10–86) km, implying a strong selection gradient across the divide. Blackcaps wintering in Britain took northwesterly migration routes from continental European breeding grounds. They originated from a surprisingly extensive area, spanning 2000 km of the breeding range. British winterers bred in sympatry with SW-bound migrants but arrived 9.8 days earlier on the breeding grounds, suggesting some potential for assortative mating by timing. Overall, our data reveal complex variation in songbird migration and suggest that selection can maintain variation in migration direction across short distances while enabling the spread of a novel strategy across a wide range.     

Seasonal matching of habitat quality and fitness in a migratory bird

When species occupy habitats that vary in quality, choice of habitat can be critical in determining individual fitness. In most migratory species, juveniles migrate independently of their parents and must therefore choose both breeding and winter habitats. Using a unique dataset of marked black-tailed godwits (Limosa limosa islandica) tracked throughout their migratory range, combined with analyses of stable carbon isotope ratios, we show that those individuals that occupy higher quality breeding sites also use higher quality winter sites. This seasonal matching can severely inflate inequalities in individual fitness. This population has expanded over the last century into poorer quality breeding and winter habitats and, across the whole population; individual birds tend to occupy either novel or traditional sites in both seasons. Winter and breeding season habitat selection are thus strongly linked throughout this population; these links have profound implications for a wide range of population and evolutionary processes. As adult godwits are highly philopatric, the initial choice of winter habitat by juveniles will be critical in determining future survival, timing of migration and breeding success.     

Tropical winter habitat limits reproductive success on the temperate breeding grounds in a migratory bird

Identifying the factors that control population dynamics in migratory animals has been constrained by our inability to track individuals throughout the annual cycle. Using stable carbon isotopes, we show that the reproductive success of a long-distance migratory bird is influenced by the quality of habitat located thousands of kilometres away on tropical wintering grounds. For male American redstarts (Setophaga ruticilla), winter habitat quality influenced arrival date on the breeding grounds, which in turn affected key variables associated with reproduction, including the number of young fledged. Based on a winter-habitat model, females occupying high-quality winter habitat were predicted to produce more than two additional young and to fledge offspring up to a month earlier compared with females wintering in poor-quality habitat. Differences of this magnitude are highly important considering redstarts are single brooded, lay clutches of only three to five eggs and spend only two-and-a-half months on the breeding grounds. Results from this study indicate the importance of understanding how periods of the annual cycle interact for migratory animals. Continued loss of tropical wintering habitat could have negative effects on migratory populations in the following breeding season, minimizing density-dependent effects on the breeding grounds and leading to further population declines. If conservation efforts are to be successful, strategies must incorporate measures to protect all the habitats used during the entire annual cycle of migratory animals.     

The strength of migratory connectivity for birds en route to breeding through the Gulf of Mexico

The strength of migratory connectivity is a measure of the cohesion of populations among phases of the annual cycle, including breeding, migration, and wintering. Many Nearctic‐Neotropical species have strong migratory connectivity between breeding and wintering phases of the annual cycle. It is less clear if this strength persists during migration when multiple endogenous and exogenous factors may decrease the cohesion of populations among routes or through time along the same routes. We sampled three bird species, American redstart Setophaga ruticilla, ovenbird Seiurus aurocapilla, and wood thrush Hylocichla mustelina, during spring migration through the Gulf of Mexico region to test if breeding populations differentiate spatially among migration routes or temporally along the same migration routes and the extent to which within‐population timing is a function of sex, age, and carry‐over from winter habitat, as measured by stable carbon isotope values in claws (δ¹³C). To make quantitative comparisons of migratory connectivity possible, we developed and used new methodology to estimate the strength of migratory connectivity (MC) from probabilistic origin assignments identified using stable hydrogen isotopes in feathers (δ²H). We found support for spatial differentiation among routes by American redstarts and ovenbirds and temporal differentiation along routes by American redstarts. After controlling for breeding origin, the timing of American redstart migration differed among ages and sexes and ovenbird migration timing was influenced by carry‐over from winter habitat. The strength of migratory connectivity did not differ among the three species, with each showing weak breeding‐to‐spring migration MC relative to prior assessments of breeding‐wintering connectivity. Our work begins to fill an essential gap in methodology and understanding of the extent to which populations remain together during migration, information critical for a full annual cycle perspective on the population dynamics and conservation of migratory animals.     

Range‐wide patterns of migratory connectivity in the western sandpiper Calidris mauri

Understanding the population dynamics of migratory animals and predicting the consequences of environmental change requires knowing how populations are spatially connected between different periods of the annual cycle. We used stable isotopes to examine patterns of migratory connectivity across the range of the western sandpiper Calidris mauri. First, we developed a winter isotope basemap from stable‐hydrogen (δD), ‐carbon (δ¹³C), and ‐nitrogen (δ¹⁵N) isotopes of feathers grown in wintering areas. δD and δ¹⁵N values from wintering individuals varied with the latitude and longitude of capture location, while δ¹³C varied with longitude only. We then tested the ability of the basemap to assign known‐origin individuals. Sixty percent of wintering individuals were correctly assigned to their region of origin out of seven possible regions. Finally, we estimated the winter origins of breeding and migrant individuals and compared the resulting empirical distribution against the distribution that would be expected based on patterns of winter relative abundance. For breeding birds, the distribution of winter origins differed from expected only among males in the Yukon‐Kuskokwim (Y‐K) Delta and Nome, Alaska. Males in the Y‐K Delta originated overwhelmingly from western Mexico, while in Nome, there were fewer males from western North America and more from the Baja Peninsula than expected. An unexpectedly high proportion of migrants captured at a stopover site in the interior United States originated from eastern and southern wintering areas, while none originated from western North America. In general, we document substantial mixing between the breeding and wintering populations of both sexes, which will buffer the global population of western sandpipers from the effects of local habitat loss on both breeding and wintering grounds.