Winter-ground microhabitat use by differently coloured phenotypes affects return rate in a long-distance migratory bird.

Migratory bird populations are declining globally at alarming rates. Non-breeding site conditions affect breeding populations, but generalising non-breeding habitat conditions over large spatial regions cannot address potential fine-scale differences across landscapes or local populations. Plumage characteristics can mediate the effects of environmental conditions on individual fitness. However, whether different phenotypes use distinctive non-breeding sites, and whether they respond to non-breeding site conditions differently remains largely unknown. Stable isotopes (δ13C, δ15N, δ2H) of inert tissues are useful to infer habitat characteristics and geographic origins where those tissues were grown. We collected winter-grown feathers from pied flycatchers (Ficedula hypoleuca) on their breeding grounds over several years from males whose dorsal plumage colouration ranged continuously from brown to black and assessed their stable isotope values as proxies of local habitat conditions. Based on feather δ2H profiles we found that browner males spent their non-breeding season in drier habitats than black males. Assignment to origin analysis shows potential regional non-breeding ground separation between differently coloured males. High within-individual repeatability of both δ13C and δ15N indicate the pied flycatcher males return yearly to similar areas. Blacker males were more likely to return to the breeding grounds after dry years compared with brown males. The opposite was found in wet years. Our study demonstrates that different phenotypes are exposed to different non-breeding site conditions which can differentially affect individual survivorship. This has important ramifications for population dynamics under predicted climate change scenarios where especially brown phenotype pied flycatcher males may be under a risk of decreasing.

Multi-isotope (δ2H, δ13C, δ15N) feather profiles and morphometrics inform patterns of migratory connectivity in three species of North American swallows.

Aerial insectivorous birds have suffered steep population declines in North America over the last 60 years. A lack of information on migratory connectivity between breeding and non-breeding grounds for these species limits our ability to interpret factors affecting their population-specific trends. We determined likely Latin American non-breeding regions of Bank (Riparia riparia), Barn (Hirundo rustica) and Cliff (Petrochelidon pyrrhonota) swallow from populations across their breeding ranges. We used predicted feather hydrogen (δ2Hf) and carbon (δ13Cf) isoscapes for winter-grown feathers to indicate areas of highest probability of moult origin and incorporated these results into a cluster analysis to determine likely broad non-breeding regions. We also assessed variation in wing length among populations to determine the potential for this metric to differentiate population moult origins. We then investigated patterns of multi-isotopic (δ2Hf, δ13Cf, δ15Nf) and wing-length niche occupancy by quantifying niche size and overlap among populations under the assumption that broad niches were consistent with low within-species migratory connectivity and narrow and non-overlapping niches with higher connectivity. Multivariate assignment identified different non-breeding regions and potential clusters of moult origin generally corresponding to Central America and northern South America, eastern and south-central South America, and the western and southern part of that continent, with variation within and among populations and species. Separate niche space indicated different wintering habitat or areas used by species or populations whereas niche overlap indicated only potential spatial similarity. Wing length varied significantly among populations by species, being longer in the west and north for Bank and Cliff Swallow and longer in eastern Canadian Barn Swallow populations. Barn Swallow occupied consistently larger isotopic and wing length niche space than the other species. Comparisons among populations across species showed variable isotopic and wing-length niche overlap generally being greater within breeding regions and lower between western and eastern breeding populations supporting a general North American continental divide for all species with generally low migratory connectivity for all species. We present a novel approach to assessing connectivity using inexpensive and broad isotopic approaches that provides the basis for hypothesis testing using more spatially explicit expensive techniques.

Inferring spatial patterns of mercury exposure in migratory boreal songbirds: Combining feather mercury and stable isotope (δ2H) measurements.

Migratory songbirds breeding in the Canadian Boreal forest are exposed to mercury (Hg), a potent neurotoxin that impairs avian health, however, the degree of exposure depends on many factors. As breeding grounds are geographically remote and vast, the measurement of Hg in individual birds is impractical particularly at large spatial scales. Here, we present a Canada-wide dataset of nearly 2000 migratory songbirds that were used to assess summer Hg exposure of 15 songbird species sampled during fall migration. We measured Hg concentrations in tail feathers and related those to dietary guild, geographic capture location, age, sex and probable breeding ground locations using feather δ2H. Overall mean (±SE) feather Hg concentration was 1.49 ± 0.03 µg/g (N = 1946): however, a clear geographic gradient in feather Hg concentrations emerged being highest in East and lowest in West. Dietary guild was the next strongest predictor of feather Hg with insectivorous songbirds in Eastern Canada at particular risk due to Hg exposure on summer breeding grounds. This broad-scale assessment of Hg exposure in migratory songbirds in Canada can be used to guide future studies on finer-scale determinants of Hg exposure in birds.

Mercury exposure to swallows breeding in Canada inferred from feathers grown on breeding and non-breeding grounds.

Aerial insectivorous birds such as swallows have been the steepest declining groups of birds in North America over the last 50 years but whether such declines are linked to contaminants has not been examined. We sampled feathers from five species of swallow at multiple locations to assess total mercury [THg] exposure for adults during the non-breeding season, and for juveniles on the breeding grounds. We assessed Hg exposure to juvenile birds in crop- and grass-dominated landscapes to determine if land-use practices influenced feather [THg]. We assayed feathers for stable isotopes (δ2H, δ13C, δ15N) as proxies for relative habitat use and diet to determine their potential influence on feather [THg]. Feather [THg] was highest in adult bank swallows (Riparia riparia) and purple martins (Progne subis) from Saskatchewan and adult cliff swallows (Petrochelidon pyrrhonota) from western regions, indicating differential exposure to Hg on the non-breeding grounds. Juvenile bank, barn (Hirundo rustica) and tree (Tachycineta bicolor) swallows had lower feather [THg] in crop-dominated landscapes than grass-dominated landscapes in Saskatchewan, potentially resulting from lower use of wetland-derived insects due to wetland drainage and intensive agriculture. Feather [THg] was related to juvenile feather stable isotopes for several species, suggesting complex interactions with diet and environmental factors. Many individuals had feather [THg] values >2 µg/g, a threshold at which deleterious effects may occur. Our findings indicate differential Hg exposure among species of swallow, regions and land-uses and highlight the need for additional research to determine dietary and finer-scale land-use impacts on individual species and populations.

Migration distance does not predict blood parasitism in a migratory songbird.

Migration can influence host-parasite dynamics in animals by increasing exposure to parasites, by reducing the energy available for immune defense, or by culling of infected individuals. These mechanisms have been demonstrated in several comparative analyses; however, few studies have investigated whether conspecific variation in migration distance may also be related to infection risk. Here, we ask whether autumn migration distance, inferred from stable hydrogen isotope analysis of summer-grown feathers (δ 2Hf) in Europe, correlates with blood parasite prevalence and intensity of infection for willow warblers (Phylloscopus trochilus) wintering in Zambia. We also investigated whether infection was correlated with individual condition (assessed via corticosterone, scaled mass index, and feather quality). We found that 43% of birds were infected with Haemoproteus palloris (lineage WW1). Using generalized linear models, we found no relationship between migration distance and either Haemoproteus infection prevalence or intensity. There was spatial variation in breeding ground origins of infected versus noninfected birds, with infected birds originating from more northern sites than noninfected birds, but this difference translated into only slightly longer estimated migration distances (~214 km) for infected birds. We found no relationship between body condition indices and Haemoproteus infection prevalence or intensity. Our results do not support any of the proposed mechanisms for migration effects on host-parasite dynamics and cautiously suggest that other factors may be more important for determining individual susceptibility to disease in migratory bird species.

Stable isotopes reveal captive vs wild origin of illegally captured songbirds in France.

Naturally occurring stable isotopes of several elements are important tracers in the study the wildlife ecology including the identification of origins for migration research, investigations of trophic relationships, and in the forensic determination of illegally harvested and captive fauna. Extensive illegal trapping for use as decoys and in cuisine has contributed to drastic declines of Ortolan Bunting Emberiza hortulana populations breeding and migrating through France. We contrasted feather hydrogen isotope (δ2Hf) values in illegally captured and subsequently confiscated Ortolan Buntings (n=234), including feathers known to have grown in captivity (n=34) or of unknown growing environment (wild or cage), with δ2Hf values in birds legally caught (ringed and released) in the wild (n=40). We sought to determine if feathers from these bird groups could be differentiated based on this single isotope. Feathers grown in captivity had considerably lower δ2Hf relative to feathers of wild birds, which is potentially indicative of tap water consumption and the use of different diets in captive birds. Further, applying mixing models to δ2Hf values revealed similar proportions of captive vs. wild origins for birds illegally captured in 2012, 2014 and 2015, and a larger proportion of individuals with feathers grown in the wild in 2013. This potentially mirrors the confiscation of birds at poaching sites only in the former years, but also of recently caught wild buntings kept captive in 2013. Our results show that even a single stable isotope (δ2H) with good association with origins where feather keratins are produced is potentially useful in understanding origins of captive birds and may advance the monitoring of illegally captured birds.

Geographical assignment of polar bears using multi-element isoscapes.

Wide-ranging apex predators are among the most challenging of all fauna to conserve and manage. This is especially true of the polar bear (Ursus maritimus), an iconic predator that is hunted in Canada and threatened by global climate change. We used combinations of stable isotopes (13C,15N,2H,18O) in polar bear hair from > 1000 individuals, sampled from across much of the Canadian Arctic and sub-Arctic, to test the ability of stable isotopic profiles to 'assign' bears to (1) predefined managed subpopulations, (2) subpopulations defined by similarities in stable isotope values using quadratic discriminant analysis, and (3) spatially explicit, isotopically distinct clusters derived from interpolated (i.e. 'kriged') isotopic landscapes, or 'isoscapes', using the partitioning around medoids algorithm. A four-isotope solution provided the highest overall assignment accuracies (~80%) to pre-existing management subpopulations with accuracy rates ranging from ~30-99% (median = 64%). Assignment accuracies of bears to hierarchically clustered ecological groups based on isotopes ranged from ~64-99%. Multivariate assignment to isotopic clusters resulted in highest assignment accuracies of 68% (33-77%), 84% (47-96%) and 74% (53-85%) using two, three and four stable isotope groups, respectively. The resulting spatial structure inherent in the multiple stable isotopic compositions of polar bear tissues is a powerful forensic tool that will, in this case, contribute to the conservation and management of this species. Currently, it is unclear what is driving these robust isotopic patterns and future research is needed to evaluate the processes behind the pattern. Nonetheless, our isotopic approach can be further applied to other apex mammalian predators under threat, such as the large felids, providing that isotopic structure occurs throughout their range.

Unravelling migration connectivity reveals unsustainable hunting of the declining ortolan bunting.

In France, illegal hunting of the endangered ortolan bunting Emberiza hortulana has been defended for the sake of tradition and gastronomy. Hunters argued that ortolan buntings trapped in southwest France originate from large and stable populations across the whole of Europe. Yet, the European Commission referred France to the Court of Justice of the European Union (EU) in December 2016 for infringements to legislation (IP/16/4213). To better assess the impact of hunting in France, we combined Pan-European data from archival light loggers, stable isotopes, and genetics to determine the migration strategy of the species across continents. Ortolan buntings migrating through France come from northern and western populations, which are small, fragmented and declining. Population viability modeling further revealed that harvesting in southwest France is far from sustainable and increases extinction risk. These results provide the sufficient scientific evidence for justifying the ban on ortolan harvesting in France.

Long-Distance Range Expansion and Rapid Adjustment of Migration in a Newly Established Population of Barn Swallows Breeding in Argentina.

When bird populations spread, long-distance pioneering populations are often backfilled by a more slowly advancing front [1-3]. The Barn Swallow Hirundo rustica, a globally distributed passerine [4, 5], expanded its breeding range an exceptional 7,000 km when it began breeding 35 years ago in its regular wintering range in Argentina [6], subsequently expanding over 500 km from its starting point [7-11]. Trans-hemispheric breeding attempts have occurred previously in related swallows [12-14], but only this colonization has lasted. Comparative studies of birds show a remarkable diversity in patterns of change in migratory habits [15-21], and these Argentine-breeding swallows might retain ancestral patterns, breeding in Argentina but returning to North America for the austral winter. Feather isotopes from these birds are consistent with the alternative possibility that they migrate no farther than northern South America [22]. Because isotopic patterns cannot definitively distinguish these alternatives, we pursued a solar geolocator study [23, 24] to do so. Data from nine tagged birds show conclusively that Barn Swallows breeding in Argentina have rapidly changed their movements to migrate no farther north in austral winter than northern South America. The phenology of the annual cycles of molt, migration, and breeding for these Argentine-breeding swallows have all shifted by about 6 months, and we suggest that stimulatory day lengths and the proliferation of nesting substrates facilitated their colonization.

A Continent-Wide Migratory Divide in North American Breeding Barn Swallows (Hirundo rustica).

Populations of most North American aerial insectivores have undergone steep population declines over the past 40 years but the relative importance of factors operating on breeding, wintering, or stopover sites remains unknown. We used archival light-level geolocators to track the phenology, movements and winter locations of barn swallows (Hirdundo rustica; n = 27) from populations across North America to determine their migratory connectivity. We identified an east-west continental migratory divide for barn swallows with birds from western regions (Washington State, USA (n = 8) and Saskatchewan, Canada (n = 5)) traveling shorter distances to wintering areas ranging from Oregon to northern Colombia than eastern populations (Ontario (n = 3) and New Brunswick (n = 10), Canada) which wintered in South America south of the Amazon basin. A single swallow from a stable population in Alabama shared a similar migration route to eastern barn swallows but wintered farther north in northeast Brazil indicating a potential leap frog pattern migratory among eastern birds. Six of 9 (67%) birds from the two eastern populations and Alabama underwent a loop migration west of fall migration routes including around the Gulf of Mexico travelling a mean of 2,224 km and 722 km longer on spring migration, respectively. Longer migration distances, including the requirement to cross the Caribbean Sea and Gulf of Mexico and subsequent shorter sedentary wintering periods, may exacerbate declines for populations breeding in northeastern North America.