Patterns and determinants of shorebird species richness in the circumpolar Arctic

Aim The intention with this study was first to investigate and describe the broad‐scale geographical patterns of species richness of breeding shorebirds (Charadriiformes; families: Charadriidae, Scolopacidae and Haematopodidae) throughout the arctic tundra biome. Secondly, after compensating for the positive relationship between net primary productivity (NPP) and species richness, the relative importance of additional ecological and historical variables for species richness was investigated. The main variables considered are NPP, length of snow‐ and ice‐free season, accessibility of regions depending on migratory flyway systems, tundra area at Pleistocene (120 and 20–18 ka bp ) and Holocene (8 ka bp ) times, and tundra area at present. Methods Information on shorebird species breeding distributions was compiled from distribution atlases and species accounts. The breeding distributions of shorebirds with ranges partly or completely in the Arctic (a total of 50 species) were mapped in ArcView 3.2 to create a raster grid layer of shorebird species richness at a 1° latitude × longitude resolution. The total and mean species richness value was calculated per each 10° of longitude sector of the Arctic. The relationships between species richness and the different climatic and environmental variables were analysed on the basis of this sector‐wise division of the arctic tundra. The influence of each variable on species richness was investigated using univariate and multivariate analyses (multivariate linear regression and general linear model). Results We found that patterns of breeding shorebird species richness in the Arctic tundra biome are to a large degree determined by the NPP, the length of the snow‐ or ice‐free season, the diversity of migratory flyways, as well as the historical extent of tundra habitat area during the maximum cooling of the last glacial period. Essentially, two main regions are distinguishable in the circumpolar Arctic regarding shorebird community richness. These are a species‐rich Beringia‐centred region and a species‐poor Atlantic‐centred region. Main conclusions The underlying explanations to these major trends may primarily be attributed to factors that operate at present through accessibility of areas from contemporary migration flyways, as well as processes that operated in the past during and after the last glacial cycle. The most prominent influence on the shorebird diversity was found for NPP in combination with the diversity of migratory flyways. These flyways provide the links between breeding and wintering resources, often separated by huge distances, and the geographical and ecological conditions associated with the shorebirds’ migration seem to be of particular importance for their breeding diversity in different sectors of circumpolar tundra.     

A polar system of intercontinental bird migration

Studies of bird migration in the Beringia region of Alaska and eastern Siberia are of special interest for revealing the importance of bird migration between Eurasia and North America, for evaluating orientation principles used by the birds at polar latitudes and for understanding the evolutionary implications of intercontinental migratory connectivity among birds as well as their parasites. We used tracking radar placed onboard the ice-breaker Oden to register bird migratory flights from 30 July to 19 August 2005 and we encountered extensive bird migration in the whole Beringia range from latitude 64 degrees N in Bering Strait up to latitude 75 degrees N far north of Wrangel Island, with eastward flights making up 79% of all track directions.The results from Beringia were used in combination with radar studies from the Arctic Ocean north of Siberia and in the Beaufort Sea to make a reconstruction of a major Siberian-American bird migration system in a wide Arctic sector between longitudes 110 degrees E and 130 degrees W, spanning one-third of the entire circumpolar circle. This system was estimated to involve more than 2 million birds, mainly shorebirds, terns and skuas, flying across the Arctic Ocean at mean altitudes exceeding 1 km (maximum altitudes 3-5 km). Great circle orientation provided a significantly better fit with observed flight directions at 20 different sites and areas than constant geographical compass orientation. The long flights over the sea spanned 40-80 degrees of longitude, corresponding to distances and durations of 1400-2600 km and 26-48 hours, respectively. The birds continued from this eastward migration system over the Arctic Ocean into several different flyway systems at the American continents and the Pacific Ocean. Minimization of distances between tundra breeding sectors and northerly stopover sites, in combination with the Beringia glacial refugium and colonization history, seemed to be important for the evolution of this major polar bird migration system.     

Does migration promote or restrict circumpolar breeding ranges of arctic birds?

Aim Migration has been suggested to promote large breeding ranges among birds because of the greater mobility of migratory compared with non‐migratory species, but migration has also been suggested to restrict breeding ranges because of evolutionary constraints imposed by the genetically based migration control programme. We aim to investigate the association between migration and the breeding ranges of both land birds and pelagic birds breeding in the Arctic region. Location The Arctic region. Methods Information on breeding and wintering ranges and migratory status of bird species breeding in the arctic tundra biome was compiled from the literature. The association between breeding range, migration distance and primary winter habitat was tested using multivariate generalized linear models and pair‐wise Mann–Whitney U‐tests. Phylogenetic effects were tested for using Mantel’s permutation tests. Results We found different relationships depending on the species’ major winter habitat. Among birds that are pelagic during winter, long‐distance migrants have the largest breeding ranges, while among terrestrial birds, residents and short‐distance migrants have the largest breeding ranges. Breeding ranges of coastal birds of all migratory distance classes are comparatively restricted. Main conclusions As a new explanation for this pattern we suggest that the possibility of colonizing large winter ranges is a key factor for the subsequent expansion of breeding ranges in arctic bird communities and possibly also in bird communities of other regions of the world. Because of the reversal in the relative extent of continents and oceans between the hemispheres, longitudinally wide winter ranges are more likely for long‐distance than short‐distance migrants among pelagic birds, while the reverse holds true for birds that use terrestrial winter habitats. For coastal birds both continents and oceans form barriers restricting colonization of extensive winter quarters and consequently also of extensive breeding ranges, regardless of the distance to the winter quarters.     

The role of migration for spatial turnover of arctic bird species in a circumpolar perspective

Several different factors may determine where species range limits are located within regions of otherwise continuously available habitat and suitable climate. Within the Arctic tundra biome many bird species are migratory and their breeding distributions are affected by migration routes that are in turn limited by factors such as suitable winter habitat, migratory stopover sites, geographical barriers and historical routes of colonization. We identified longitudinal zones in the circumpolar Arctic of pronounced changes in the avian species composition (high species spatial turnover; ‘species divides’). We tested for the association between migratory status and the geographical location and numbers of such species divides for species with non‐breeding habitats mainly within terrestrial, pelagic and coastal ecosystems. Our results demonstrate that migration is of profound importance for both the number and locations of species divides in the Arctic. Long‐distance migration is associated with a large number of divides among terrestrial and coastal arctic birds but with a reduced number of divides among pelagic birds. We suggest that long‐distance migration permits pelagic but not terrestrial and coastal birds to colonize large winter ranges, which in turn causes expansion of breeding ranges, with more homogenous communities and reduction of species divides as consequences, among the long‐distance migrants of pelagic but not of terrestrial and coastal birds. Furthermore, the divides among long‐distance migrants are situated in two main regions, the Beringia and Greenland zones, while divides among short‐distance migrants are more evenly spaced throughout the circumpolar Arctic. The Beringia and Greenland divides result largely from inter‐continental colonization of new breeding ranges but retainment of original winter quarters in a process of evolution through extension of migration routes, leading to aggregated divides in the meeting zones of major global flyways.     

Migratory routes of red‐necked phalaropes Phalaropus lobatus breeding in southern Chukotka revealed by geolocators

The migration routes of red‐necked phalaropes breeding around the Bering Sea are poorly known, despite the fact that the Bering Sea could mark the boundary between the East Palearctic populations that winter in the Pacific Ocean around the East Indies and the West Nearctic populations that winter in the Pacific Ocean off the coast of South America. Geolocator data retrieved from two male phalaropes tagged in southern Chukotka, Far Eastern Russia, confirm that birds breeding in this region belong to the East Palearctic population and winter in the East Indies, suggesting that the division line with the West Nearctic population is farther to the east. The routes taken by the two phalaropes were almost entirely pelagic, totaling around 18 000–20 000 km round‐trip, with the birds continuously on the move during migration, rather than resident in any particular stopover site, contrary to most other migratory shorebirds.     

Irruptive movements and breeding dispersal of snowy owls: a specialized predator exploiting a pulsed resource

Mobility and irruptive movements have been proposed as mechanisms that could allow some diet specialists to inhabit and breed in environments with highly unpredictable resources, like the arctic tundra. The snowy owl, one of the main avian predators of the tundra, is known to specialize on lemmings during the breeding season. These small mammals are also well known for their tremendous spatial and temporal variations in abundance. We examined the spring (pre‐breeding, from March to June) movements of snowy owls by tracking 9 breeding females in the Canadian Arctic for up to 3 yr with satellite transmitters. We used state‐space modeling to assess searching behavior and measure breeding dispersal distances. We also ascertain lemming abundance at some of the sites used by the marked owls. Tracked owls displayed searching movements for extended periods (up to 108 d) and traveled over large distances (up to 4093 km) each spring. The distance between furthest apart searching areas in a given year averaged 828 km (range 220 to 2433 km). Settlement date, distance between searching areas, traveled distance and the duration of prospecting movements were longer in the year where density of lemmings recorded in the eastern High‐Arctic (Bylot Island) was lowest. Nonetheless, snowy owls settled in areas where local lemming abundance was relatively high. Individual breeding dispersal distance between consecutive years averaged 725 km (range 18 to 2224). Overall, the high mobility of female snowy owls allowed these diet specialists to behave as irruptive migrants and to sustain their reproductive activities during consecutive years even under highly fluctuating resources.     

An avian terrestrial predator of the Arctic relies on the marine ecosystem during winter

Top predators of the arctic tundra are facing a long period of very low prey availability during winter and subsidies from other ecosystems such as the marine environment may help to support their populations. Satellite tracking of snowy owls, a top predator of the tundra, revealed that most adult females breeding in the Canadian Arctic overwinter at high latitudes in the eastern Arctic and spend several weeks (up to 101 d) on the sea‐ice between December and April. Analysis of high‐resolution satellite images of sea‐ice indicated that owls were primarily gathering around open water patches in the ice, which are commonly used by wintering seabirds, a potential prey. Such extensive use of sea‐ice by a tundra predator considered a small mammal specialist was unexpected, and suggests that marine resources subsidize snowy owl populations in winter. As sea‐ice regimes in winter are expected to change over the next decades due to climate warming, this may affect the wintering strategy of this top predator and ultimately the functioning of the tundra ecosystem.     

Geographical segregation in Dunlin Calidris alpina populations wintering along the East Atlantic migratory flyway – evidence from mitochondrial DNA analysis

Dunlin Calidris alpina is one of the most abundant shorebirds using coastal habitats in the East Atlantic migratory flyway, that links arctic breeding locations (Greenland to Siberia) with wintering grounds (West Europe to West Africa). Differential migration and winter segregation between populations have been indicated by morphometrics and ringing recoveries. Here, we analyse the potential of genetic markers (mitochondrial DNA – mtDNA) to validate and enhance such findings. We compared mtDNA haplotypes frequencies at different wintering sites (from north-west Europe to West Africa). All birds from West Africa had western (European) haplotypes, while the eastern (Siberian) haplotypes were only present in European winter samples, reaching higher frequencies further north in Europe. Compilation of published results from migrating birds also confirmed these differences, with the sole presence of European haplotypes in Iberia and West Africa and increasingly higher frequencies of Siberian haplotypes from south-west to north-west Europe. Comparison with published haplotype frequencies of breeding populations shows that birds from Greenland, Iceland, and North Europe were predominant in wintering grounds in West Africa, while populations wintering in West Europe originated from more eastern breeding grounds (e.g. North Russia). These results show that genetic markers can be used to enhance the integrative monitoring of wintering and breeding populations, by providing biogeographical evidence that validate the winter segregation of breeding populations.     

A long winter for the Red Queen: rethinking the evolution of seasonal migration

This paper advances an hypothesis that the primary adaptive driver of seasonal migration is maintenance of site fidelity to familiar breeding locations. We argue that seasonal migration is therefore principally an adaptation for geographic persistence when confronted with seasonality – analogous to hibernation, freeze tolerance, or other organismal adaptations to cyclically fluctuating environments. These ideas stand in contrast to traditional views that bird migration evolved as an adaptive dispersal strategy for exploiting new breeding areas and avoiding competitors. Our synthesis is supported by a large body of research on avian breeding biology that demonstrates the reproductive benefits of breeding‐site fidelity. Conceptualizing migration as an adaptation for persistence places new emphasis on understanding the evolutionary trade‐offs between migratory behaviour and other adaptations to fluctuating environments both within and across species. Seasonality‐induced departures from breeding areas, coupled with the reproductive benefits of maintaining breeding‐site fidelity, also provide a mechanism for explaining the evolution of migration that is agnostic to the geographic origin of migratory lineages (i.e. temperate or tropical). Thus, our framework reconciles much of the conflict in previous research on the historical biogeography of migratory species. Although migratory behaviour and geographic range change fluidly and rapidly in many populations, we argue that the loss of plasticity for migration via canalization is an overlooked aspect of the evolutionary dynamics of migration and helps explain the idiosyncratic distributions and migratory routes of long‐distance migrants. Our synthesis, which revolves around the insight that migratory organisms travel long distances simply to stay in the same place, provides a necessary evolutionary context for understanding historical biogeographic patterns in migratory lineages as well as the ecological dynamics of migratory connectivity between breeding and non‐breeding locations.     

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.     

Dichotomous strategies? The migration of Whimbrels breeding in the eastern Canadian sub‐Arctic

The conservation of migratory birds requires internationally coordinated efforts that, in turn, demand an understanding of population dynamics and connectivity throughout a species' range. Whimbrels (Numenius phaeopus) are a widespread long‐distance migratory shorebird with two disparate North American breeding populations. Monitoring efforts suggest that at least one of these populations is declining, but the level of migratory connectivity linking the two populations to specific non‐breeding sites or identifiable conservation threats remains unclear. We deployed light‐level geolocators in 2012 to track the migration of Whimbrels breeding near Churchill, Manitoba, Canada. In 2013, we recovered 11 of these geolocators, yielding complete migration tracks for nine individuals. During southbound migration, six of the nine Whimbrels stopped at two staging sites on the mid‐Atlantic seaboard of the United States for an average of 22 days, whereas three individuals made nonstop flights of ~8000 km from Churchill to South America. All individuals subsequently spent the entire non‐breeding season along the northern coasts of Brazil and Suriname. On their way north, all birds stopped at the same two staging sites used during southbound migration. Individuals staged at these sites for an average of 34 days, significantly longer than during southbound migration, and all departed within a 5‐day period to undertake nonstop flights ranging from 2600 to 3100 km to the breeding grounds. These extended spring stopovers suggest that female Whimbrels likely employ a mixed breeding strategy, drawing on both endogenous and exogenous reserves to produce their eggs. Our results also demonstrate that this breeding population exhibits a high degree of connectivity among breeding, staging, and wintering sites. As with other long‐distance migratory shorebirds, conservation efforts for this population of Whimbrels must therefore focus on a small, but widely spaced, suite of sites that support a large proportion of the population.     

Cross‐continental migratory connectivity and spatiotemporal migratory patterns in the great reed warbler

Migratory connectivity describes to which degree different breeding populations have distinct (non‐overlapping) non‐breeding sites. Uncovering the level of migratory connectivity is crucial for effective conservation actions and for understanding of the evolution of local adaptations and migratory routes. Here we investigate migration patterns in a passerine bird, the great reed warbler Acrocephalus arundinaceus, over its wide Western Palearctic breeding range using geolocators from Spain, Sweden, Czech Republic, Bulgaria and Turkey. We found moderate migratory connectivity: a highly significant spatial structure in the connections between breeding and sub‐Saharan non‐breeding grounds, but at the same time a partial overlap between individual populations, particularly along the Gulf of Guinea where the majority of birds from the Spanish, Swedish and Czech populations spent their non‐breeding period. The post‐breeding migration routes were similar in direction and rather parallel for the five populations. Birds from Turkey showed the most distinctive migratory routes and sub‐Saharan non‐breeding range, with a post‐breeding migration to east Africa and, together with birds from Bulgaria, a previously unknown pre‐breeding migration over the Arabian Peninsula indicating counter‐clockwise loop migration. The distances between breeding and sub‐Saharan non‐breeding sites, as well as between first and final sub‐Saharan non‐breeding sites, differed among populations. However, the total speed of migration did not differ significantly between populations; neither during post‐breeding migration in autumn, nor pre‐breeding migration in spring. There was also no significant relationship between the total speed of migration and distance between breeding and non‐breeding sites (neither post‐ nor pre‐breeding) and, surprisingly, the total speed of migration generally did not differ significantly between post‐breeding and pre‐breeding migration. Future challenges include understanding whether non‐breeding environmental conditions may have influenced the differences in migratory patterns that we observed between populations, and to which extent non‐breeding habitat fluctuations and loss may affect population sizes of migrants.     

Differentiation of tundra/taiga and boreal coniferous forest wolves: genetics, coat colour and association with migratory caribou

The grey wolf has one of the largest historic distributions of any terrestrial mammal and can disperse over great distances across imposing topographic barriers. As a result, geographical distance and physical obstacles to dispersal may not be consequential factors in the evolutionary divergence of wolf populations. However, recent studies suggest ecological features can constrain gene flow. We tested whether wolf-prey associations in uninterrupted tundra and forested regions of Canada explained differences in migratory behaviour, genetics, and coat colour of wolves. Satellite-telemetry data demonstrated that tundra wolves (n = 19) migrate annually with caribou (n = 19) from denning areas in the tundra to wintering areas south of the treeline. In contrast, nearby boreal coniferous forest wolves are territorial and associated year round with resident prey. Spatially explicit analysis of 14 autosomal microsatellite loci (n = 404 individuals) found two genetic clusters corresponding to tundra vs. boreal coniferous forest wolves. A sex bias in gene flow was inferred based on higher levels of mtDNA divergence (F(ST) = 0.282, 0.028 and 0.033; P < 0.0001 for mitochondrial, nuclear autosomal and Y-chromosome markers, respectively). Phenotypic differentiation was substantial as 93% of wolves from tundra populations exhibited light colouration whereas only 38% of boreal coniferous forest wolves did (chi(2) = 64.52, P < 0.0001). The sharp boundary representing this discontinuity was the southern limit of the caribou migration. These findings show that substantial genetic and phenotypic differentiation in highly mobile mammals can be caused by prey-habitat specialization rather than distance or topographic barriers. The presence of a distinct wolf ecotype in the tundra of North America highlights the need to preserve migratory populations.     

Red fox takeover of arctic fox breeding den: an observation from Yamal Peninsula, Russia

Here, we report from the first direct observation of a red fox (Vulpes vulpes) intrusion on an arctic fox (Vulpes lagopus) breeding den from the southern Arctic tundra of Yamal Peninsula, Russia in 2007. At the same time, as a current range retraction of the original inhabitant of the circumpolar tundra zone the arctic fox is going on, the red fox is expanding their range from the south into arctic habitats. Thus, within large parts of the northern tundra areas the two species are sympatric which gives opportunities for direct interactions including interference competition. However, direct first-hand observations of such interactions are rare, especially in the Russian Arctic. In the present study, we observed one red fox taking over an arctic fox breeding den which resulted in den abandonment by the arctic fox. On July 19, eight arctic fox pups were observed on the den before the red fox was observed on the same den July 22. The pups were never seen at the den or elsewhere after the red fox was observed on the den for as long as we stayed in the area (until August 10). Our observation supports the view that direct interference with red fox on breeding dens may contribute to the range retraction of arctic foxes from the southern limits of the Arctic tundra in Russia.     

Migratory connectivity in a declining bird species: using feather isotopes to inform demographic modelling

Aim Conservation programmes for endangered migratory species or populations require locating and evaluating breeding, stopover and wintering areas. We used multiple stable isotopes in two endangered European populations of wrynecks, Jynx torquilla L., to locate wintering regions and assess the degree of migratory connectivity between breeding and wintering populations. Location Switzerland and Germany. Methods We analysed stable nitrogen (δ¹⁵N), carbon (δ¹³C) and hydrogen (δD) isotopes from wing feathers from two populations of wrynecks to infer their wintering origins and to assess the strength of migratory connectivity. We tested whether variation in feather isotopic values within the Swiss population was affected by bird age and collection year and then considered differences in isotopic values between the two breeding populations. We used isotopic values of summer‐ and winter‐grown feathers to estimate seasonal distributions. Finally, we calculated a species‐specific δD discrimination factor between feathers and mean annual δD values to assign winter‐grown feathers to origin. Results Bird age and collection year caused substantial isotopic variation in winter‐grown feathers, which may be because of annually variable weather conditions, movements of birds among wintering sites and/or reflect asynchronous moulting or selection pressure. The large isotopic variance in winter‐grown feathers nevertheless suggested low migratory connectivity for each breeding population, with partially overlapping wintering regions for the two populations. Main conclusions Isotopic variance in winter‐grown feathers of two breeding populations of wrynecks and their geographical assignment point to defined, albeit overlapping, wintering areas, suggesting both leapfrog migration and low migratory connectivity. On this basis, integrative demographic models can be built looking at seasonal survival patterns with links to local environmental conditions on both breeding and wintering grounds, which may elucidate causes of declines in migratory bird species.     

Light‐level geolocators reveal migratory connectivity in European populations of pied flycatchers Ficedula hypoleuca

Understanding what drives or prevents long‐distance migrants to respond to environmental change requires basic knowledge about the wintering and breeding grounds, and the timing of movements between them. Both strong and weak migratory connectivity have been reported for Palearctic passerines wintering in Africa, but this remains unknown for most species. We investigated whether pied flycatchers Ficedula hypoleuca from different breeding populations also differ in wintering locations in west‐Africa. Light‐level geolocator data revealed that flycatchers from different breeding populations travelled to different wintering sites, despite similarity in routes during most of the autumn migration. We found support for strong migratory connectivity showing an unexpected pattern: individuals breeding in Fennoscandia (S‐Finland and S‐Norway) wintered further west compared to individuals breeding at more southern latitudes in the Netherlands and SW‐United Kingdom. The same pattern was found in ring recovery data from sub‐Saharan Africa of individuals with confirmed breeding origin. Furthermore, population‐specific migratory connectivity was associated with geographical variation in breeding and migration phenology: birds from populations which breed and migrate earlier wintered further east than birds from ‘late’ populations. There was no indication that wintering locations were affected by geolocation deployment, as we found high repeatability and consistency in δ¹³C and δ¹⁵N stable isotope ratios of winter grown feathers of individuals with and without a geolocator. We discuss the potential ecological factors causing such an unexpected pattern of migratory connectivity. We hypothesise that population differences in wintering longitudes of pied flycatchers result from geographical variation in breeding phenology and the timing of fuelling for spring migration at the wintering grounds. Future research should aim at describing how temporal dynamics in food availability across the wintering range affects migration, wintering distribution and populations’ capacity to respond to environmental changes.     

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.     

Wing shape and migration in shorebirds: a comparative study

Migration is an energetically expensive and hazardous stage of the annual cycle of non‐resident avian species, and requires certain morphological adaptations. Wing shape is one of the morphological traits that is expected to be evolutionarily shaped by migration. Aerodynamic theory predicts that long‐distance migrants should have more pointed wings with distal primaries relatively longer than proximal primaries, an arrangement that minimizes induced drag and wing inertia, but this prediction has mostly been tested in passerine species. We applied the comparative method of phylogenetically independent contrasts to assess convergent evolution between wing shape and migration within shorebirds. We confirmed the assumption that long‐distance migrants have less rounded wings than species migrating shorter distances. Furthermore, wing roundedness negatively correlates with fat load and mean distance of migratory flights, the basic components of migration strategies. After controlling for interspecific differences in body size, we found no support for a link between wing length and migration, indicating that wing shape is a more important predictor of shorebird migratory behaviour than wing length. The results suggest that total migration distance and migratory strategy may simultaneously act on the evolution of wing shape in shorebirds, and possibly in other avian species.     

Migratory connectivity and population-specific migration routes in a long-distance migratory bird

Knowledge about migratory connectivity, the degree to which individuals from the same breeding site migrate to the same wintering site, is essential to understand processes affecting populations of migrants throughout the annual cycle. Here, we study the migration system of a long-distance migratory bird, the Montagu''s harrier Circus pygargus, by tracking individuals from different breeding populations throughout northern Europe. We identified three main migration routes towards wintering areas in sub-Saharan Africa. Wintering areas and migration routes of different breeding populations overlapped, a pattern best described by ‘weak (diffuse) connectivity’. Migratory performance, i.e. timing, duration, distance and speed of migration, was surprisingly similar for the three routes despite differences in habitat characteristics. This study provides, to our knowledge, a first comprehensive overview of the migration system of a Palaearctic-African long-distance migrant. We emphasize the importance of spatial scale (e.g. distances between breeding populations) in defining patterns of connectivity and suggest that knowledge about fundamental aspects determining distribution patterns, such as the among-individual variation in mean migration directions, is required to ultimately understand migratory connectivity. Furthermore, we stress that for conservation purposes it is pivotal to consider wintering areas as well as migration routes and in particular stopover sites.     

Molecular Detection of Hematozoa Infections in Tundra Swans Relative to Migration Patterns and Ecological Conditions at Breeding Grounds

Tundra swans (Cygnus columbianus) are broadly distributed in North America, use a wide variety of habitats, and exhibit diverse migration strategies. We investigated patterns of hematozoa infection in three populations of tundra swans that breed in Alaska using satellite tracking to infer host movement and molecular techniques to assess the prevalence and genetic diversity of parasites. We evaluated whether migratory patterns and environmental conditions at breeding areas explain the prevalence of blood parasites in migratory birds by contrasting the fit of competing models formulated in an occupancy modeling framework and calculating the detection probability of the top model using Akaike Information Criterion (AIC). We described genetic diversity of blood parasites in each population of swans by calculating the number of unique parasite haplotypes observed. Blood parasite infection was significantly different between populations of Alaska tundra swans, with the highest estimated prevalence occurring among birds occupying breeding areas with lower mean daily wind speeds and higher daily summer temperatures. Models including covariates of wind speed and temperature during summer months at breeding grounds better predicted hematozoa prevalence than those that included annual migration distance or duration. Genetic diversity of blood parasites in populations of tundra swans appeared to be relative to hematozoa prevalence. Our results suggest ecological conditions at breeding grounds may explain differences of hematozoa infection among populations of tundra swans that breed in Alaska.