Connecting breeding and wintering grounds of Neotropical migrant songbirds using stable hydrogen isotopes: a call for an isotopic atlas of migratory connectivity

There is an overdue and urgent need to establish patterns of migratory connectivity linking breeding grounds, stopover sites, and wintering grounds of migratory birds. Such information allows more effective application of conservation efforts by applying focused actions along movement trajectories at the population level. Stable isotope methods, especially those using stable hydrogen isotope abundance in feathers (δ²H_f) combined with Bayesian assignment techniques incorporating prior information such as relative abundance of breeding birds, now provide a fast and reliable means of establishing migratory connectivity, especially for Neotropical migrants that breed in North America and molt prior to fall migration. Here we demonstrate how opportunistic sampling of feathers of 30 species of wintering birds in Cuba, Venezuela, Guatemala, Puerto Rico, and Mexico, regions that have typically been poorly sampled for estimating migratory connectivity, can be assigned to breeding areas in North America through both advanced spatial assignment to probability surfaces and through simpler map lookup approaches. Incorporating relative abundance information from the North American Breeding Bird Survey in our Bayesian assignment models generally resulted in a reduction in potential assignment areas on breeding grounds. However, additional tools to constrain longitude such as DNA markers or other isotopes would be desirable for establishing breeding or molt origins of species with broad longitudinal distributions. The isotope approach could act as a rapid means of establishing basic patterns of migratory connectivity across numerous species and populations. We propose a large‐scale coordinated sampling effort on the wintering grounds to establish an isotopic atlas of migratory connectivity for North American Neotropical migrants and suggest that isotopic variance be considered as a valuable metric to quantify migratory connectivity. This initiative could then act as a strategic template to guide further efforts involving stable isotopes, light‐sensitive geolocators, and other technologies.     

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.     

Temperate migrants and resident bird species in Afro‐tropical savannahs show similar levels of ecological generalism

The specificity of an animal's habitat requirements will determine its ability to deal with anthropogenic climate and habitat change. Migratory birds are thought to be particularly vulnerable to such change, but theory predicts that they should be largely generalists. This prediction was tested with the aim of assessing whether migratory Palaearctic‐breeding birds wintering in the savannah biome of Africa are more or less generalist in their habitat use compared with taxonomically and ecologically similar Afro‐tropical resident species. The degree of specialization of these species groups to certain habitat characteristics was assessed and compared by calculating the relative occurrence of the species along habitat gradients, where wide occurrence indicates generalism and narrow occurrence indicates specialism. Palaearctic migrants as a group could not clearly be distinguished as generalists relative to Afro‐tropical residents with respect to habitat attributes. The only indication of greater flexibility in Palaearctic migrants was a significant tendency to use habitats over a wider latitudinal range. The results suggest that migrants are generalists, but not necessarily more generalist than taxonomically similar resident species that also occur over a wide range of habitat types within the savannah biome. The availability of specific habitat requirements on the wintering grounds in Africa is therefore unlikely to be a primary limiting factor for many Afro‐Palaearctic migratory bird species.     

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.     

Stable isotope analysis provides new information on winter habitat use of declining avian migrants that is relevant to their conservation

Winter habitat use and the magnitude of migratory connectivity are important parameters when assessing drivers of the marked declines in avian migrants. Such information is unavailable for most species. We use a stable isotope approach to assess these factors for three declining African-Eurasian migrants whose winter ecology is poorly known: wood warbler Phylloscopus sibilatrix, house martin Delichon urbicum and common swift Apus apus. Spatially segregated breeding wood warbler populations (sampled across a 800 km transect), house martins and common swifts (sampled across a 3,500 km transect) exhibited statistically identical intra-specific carbon and nitrogen isotope ratios in winter grown feathers. Such patterns are compatible with a high degree of migratory connectivity, but could arise if species use isotopically similar resources at different locations. Wood warbler carbon isotope ratios are more depleted than typical for African-Eurasian migrants and are compatible with use of moist lowland forest. The very limited variance in these ratios indicates specialisation on isotopically restricted resources, which may drive the similarity in wood warbler populations' stable isotope ratios and increase susceptibility to environmental change within its wintering grounds. House martins were previously considered to primarily use moist montane forest during the winter, but this seems unlikely given the enriched nature of their carbon isotope ratios. House martins use a narrower isotopic range of resources than the common swift, indicative of increased specialisation or a relatively limited wintering range; both factors could increase house martins' vulnerability to environmental change. The marked variance in isotope ratios within each common swift population contributes to the lack of population specific signatures and indicates that the species is less vulnerable to environmental change in sub-Saharan Africa than our other focal species. Our findings demonstrate how stable isotope research can contribute to understanding avian migrants' winter ecology and conservation status.     

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

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

Isotopic variation across the Audubon's–myrtle warbler hybrid zone

Differences in seasonal migratory behaviours are thought to be an important component of reproductive isolation in many organisms. Stable isotopes have been used with success in estimating the location and qualities of disjunct breeding and wintering areas. However, few studies have used isotopic data to estimate the movements of hybrid offspring in species that form hybrid zones. Here, we use stable hydrogen to estimate the wintering locations and migratory patterns of two common and widespread migratory birds, Audubon's (Setophaga auduboni) and myrtle (S. coronata) warblers, as well as their hybrids. These two species form a narrow hybrid zone with extensive interbreeding in the Rocky Mountains of British Columbia and Alberta, Canada, which has been studied for over four decades. Isotopes in feathers grown on the wintering grounds or early on migration reveal three important patterns: (1) Audubon's and myrtle warblers from allopatric breeding populations winter in isotopically different environments, consistent with band recovery data and suggesting that there is a narrow migratory transition between the two species, (2) most hybrids appear to overwinter in the south‐eastern USA, similar to where myrtle warblers are known to winter, and (3) some hybrid individuals, particularly those along the western edge of the hybrid zone, show Audubon's‐like isotopic patterns. These data suggest there is a migratory divide between these two species, but that it is not directly coincident with the centre of the hybrid zone in the breeding range. We interpret these findings and discuss them within the context of previous research on hybrid zones, speciation and migratory divides.     

High variation reduces the value of feather stable isotope ratios in identifying new wintering areas for aquatic warblers Acrocephalus paludicola in West Africa

Stable isotope analysis of feathers can be useful in the study of seasonal interactions and migratory connectivity in birds. For the Palaearctic–African migration system, however, the lack of isotope data from feathers of known origin in Africa renders the geographic assignment of birds captured on European breeding grounds to potential wintering areas problematic. Rectrices of the threatened aquatic warbler Acrocephalus paludicola grown in Africa were sampled across six European countries to assess whether birds in different breeding populations shared similar isotopic signatures and so were likely to have wintered in the same region in Africa. Freshly grown feathers of aquatic warblers collected at the only known wintering site in Senegal showed high variation in carbon, nitrogen, and hydrogen isotope ratios. Due to similarly high variation in isotope ratios of African‐grown feathers within all breeding populations, it was not possible to determine whether different populations wintered in different regions. However, isotope signatures of 20% of birds captured on European breeding grounds fell outside the range of those captured in Senegal, suggesting a wider wintering distribution than is currently known. We therefore assessed whether the origin of these feathers could be estimated by trying to establish isotopic gradients across sub‐Saharan West Africa. Feathers of three ecologically similar surrogate species were sampled from wetlands across a 3000 km east‐west and a 2000 km north–south transect. Within‐site variation in feather isotope ratios was frequently larger than the difference predicted by gradients across West Africa. Thus, predicting the origin of individual feathers using single‐isotope gradients was not reliable. The large within‐site variability of feather isotope ratios of a habitat specialist species like the aquatic warbler indicates that using feather isotope ratios will require large sample sizes from many locations, and may thus not be an efficient tool in identifying wintering areas of Palaearctic–African migrants.     

Effects of flyway‐wide weather conditions and breeding habitat on the breeding abundance of migratory boreal waterbirds

Anthropogenic habitat loss and climate change are among the major threats to biodiversity. Bioclimatic zones such as the boreal and arctic regions are undergoing rapid environmental change, which will likely trigger changes in wildlife communities. Disentangling the effects of different drivers of environmental change on species is fundamental to better understand population dynamics under changing conditions. Therefore, in this study we investigate the synergistic effect of winter and summer weather conditions and habitat type on the abundance of 17 migratory boreal waterbird species breeding in Finland using three decades (1986–2015) of count data. We found that above‐average temperatures and precipitations across the western and northern range of the wintering grounds have a positive impact on breeding numbers in the following season, particularly for waterbirds breeding in eutrophic wetlands. Conversely, summer temperatures did not seem to affect waterbird abundance. Moreover, waterbird abundance was higher in eutrophic than in oligotrophic wetlands, but long term trends indicated that populations are decreasing faster in eutrophic than in oligotrophic wetlands. Our results suggest that global warming may apparently benefit waterbirds, e.g. by increased winter survival due to more favourable winter weather conditions. However, the observed population declines, particularly in eutrophic wetlands, may also indicate that the quality of breeding habitat is rapidly deteriorating through increased eutrophication in Finland which override the climatic effects. The findings of this study highlight the importance of embracing a holistic approach, from the level of a single catchment up to the whole flyway, in order to effectively address the threats that waterbirds face on their breeding as well as wintering grounds.