Natal origins and timing of migration of two passerine species through the southern Alps: inferences from multiple stable isotopes (δ2H, δ13C, δ15N, δ34S) and ringing data

Understanding spatial linkages between areas used by migratory animals during the annual cycle is fundamental to their conservation. Stable isotope measurements of animal tissues can be a valuable tool in understanding spatial connectivity and migration phenology of migratory wildlife. We inferred natal origins of two migratory passerines, European Pied Flycatcher Ficedula hypoleuca and European Robin Erithacus rubecula, captured during autumn migration in the Italian Alps, by combining feather δ²H (δ²H_f) and ring recovery data. We used a spatially explicit likelihood-based method to assign individuals to a precipitation δ²H surface calibrated to represent feather δ²H, together with the directional probability of origin from ring recoveries. The highest probabilities of origin for most individuals of both species were in central and north-eastern Europe. Seasonal trends in δ²H_f, which described the species’ migratory phenology through the Italian Alps, were correlated with feather δ¹³C, δ¹⁵N and δ³⁴S values, indicating strong spatial discrimination related to continental patterns for these isotopes. We demonstrate how this combined information can define catchment areas and migratory connectivity of birds intercepted in the Alps. We highlight the importance of ringing data in defining directional priors to define Bayesian-based probability surfaces using continental δ²H_f isoscapes, and how such information can be used to inform estimates of migratory connectivity.     

Space‐time tradeoffs in the development of precipitation‐based isoscape models for determining migratory origin

Precipitation stable isotope patterns over continental scales provide a fundamental tool for tracking origins of migratory species. Hydrogen isotopes from rain and environmental waters are assimilated into animal tissues and may thereby reveal the location where tissues were synthesized. Predictive isotopic maps (or isoscapes) of stable hydrogen isotope values in precipitation (δ²H_p) are typically generated by time‐averaging observations from a global network of stations that have been sampled irregularly in space and time. We previously demonstrated that restricting the temporal range in δ²H_p isoscapes to biologically relevant time frames did not improve predictions of geographic origin for two migratory species in North America and Europe; rather, it decreased the accuracy of assignment. Here, we examined whether the reduction in assignment accuracy stemmed from a decrease in the number of sampling stations available to support isoscape development for shorter time periods. Multiple regression models were used to predict the hydrogen isotope composition in precipitation using isotopic measurements from each station along with a suite of independent variables. The reduction in the number of stations with δ²H_p measurements used to estimate isoscape model parameters did not alter the accuracy and precision of assignments consistently. We also examined accuracy across a range of reduced station numbers and found that mean accuracy was affected only at very low numbers of stations, indicating that the spatial isotopic patterns in precipitation that are useful for assignment applications can be characterized with data from relatively limited data stations. The number and spatial distribution of stations may have more influence when geostatistical models are used to generate isoscapes, as they incorporate spatial correlation in the dataset. The results can be used to guide future research in understanding how data availability and constraints in creating δ²H_p isoscapes may affect predictions of geographic origins.     

Conservation through connectivity: can isotopic gradients in Africa reveal winter quarters of a migratory bird?

Conservation of migratory wildlife requires knowledge of migratory connectivity between breeding and non-breeding locations. Stable isotopes in combination with geographical isotopic patterns (isoscapes) can provide inferences about migratory connectivity. This study examines whether such an approach can be used to infer wintering areas in sub-Saharan Africa, where we lack such knowledge for many species, but where this method has not been used widely. We measured δ ²H, δ ¹³C and δ ¹⁵N in winter-grown feathers of a breeding Swiss and Spanish population of European hoopoe Upupa epops—a typical Palaearctic-Afrotropical migrant. δ ²H values predicted that ~70 % of the hoopoes spent the non-breeding season in the western portion of their potential winter range. This was corroborated by a shallow east–west gradient in feather-δ ²H values of museum specimens from known African origin across the potential winter range and by the recovery of Swiss hoopoes marked with geolocators. Hoopoes categorized as from eastern versus western regions of the wintering range were further delineated spatially using feather δ ¹³C and δ ¹⁵N. δ ¹⁵N showed no trend, whereas adults were more enriched in ¹³C in the western portion of the range, with eastern adults being in addition more depleted in ¹³C than eastern juveniles. This suggests that eastern juveniles may have occupied more xeric habitats than sympatric adults. We demonstrated that stable isotopes, especially δ ²H, could only very roughly delineate the winter distribution of a trans-Saharan Palaearctic migrant restricted primarily to the Sahelian and savanna belt south of the Sahara. Further refinements of precipitation isoscapes for Africa as well the development of isoscapes for δ ¹³C and δ ¹⁵N may improve assignment of this and other migrants.     

Improving provenance studies in migratory birds when using feather hydrogen stable isotopes

Hydrogen stable isotopes (δ²H) in feathers are used to determine the origin and migration strategy of birds. To identify the geographic location of the site of feather synthesis, calibration curves for the relation between feather δ²H and amount‐weighted growing‐season δ²H in precipitation are used to generate feather δ²H isoscapes. Factors like species, age and year might generate isotopic variation in calibration curves, but the extent to which accounting for variation may improve calibration curves and hence provenance determination of birds, is unknown. We compared three European calibration curves: 1) an existing multi‐species curve, uncorrected for age and year variation, and two species‐specific calibration curves, based on mallard Anas platyrhynchos feathers, of 2) varying age and year, and 3) juvenile natal origin, corrected for year variation. Calibration curves using ordinary least square linear regression (OLS) as opposed to standard major axis regression showed least bias in estimation. As expected, we found that a single species (mallard) OLS calibration curve corrected for age and year yielded the highest coefficient of determination, but was still surprisingly similar to the other two calibration curves. Nevertheless, when using feathers of known‐origin to assess provenance accuracy, the calibration curve that accounted for species, age and year variation yielded the best prediction in as many as 59% of the cases. Our study is the first to demonstrate implications of isotopic variation on assessing the origin of individual birds, but also highlights the relatively small gain in precision that is achieved by generating species, age and year specific calibration curves rather than resorting to more general alternatives.     

Flyway population delineation in Taiga Bean Geese Anser fabalis fabalis revealed by multi‐element feather stable isotope analysis

Fundamental to effective management of migratory waterbird populations is an understanding of their flyway delineation. Taiga Bean Geese Anser fabalis fabalis wintering in NW Denmark, Scotland and England are considered to originate from northern and central Sweden, southern and central Norway (‘Western flyway’), those wintering in southern Sweden, NE and southern Denmark are considered to originate from northern Fennoscandia and western Russia (‘Central flyway’), and those wintering in eastern Germany and Poland (which show far less favourable conservation status) are thought to come from western Siberia (‘Eastern 1 flyway’), although evidence to demonstrate this has largely been lacking. Evidence for different natal and moult origins of Taiga Bean Geese was investigated using stable isotope analyses of feathers of four elements (δ²H, δ¹³C, δ¹⁵N and δ³⁴S). There were significant differences in isotopic composition of feathers from Swedish (Central) and German (Eastern 1) wintering populations and those moulting in Sweden in late summer (Western), which validated the three proposed major management flyway units above. The strong continental gradient in the stable hydrogen isotope ratios in precipitation (δ²H_p) across the region was used to assign wintering birds geospatially to natal and moulting origin, indicating separate natal and moulting areas for German (n = 37, from western Siberia) and Swedish (n = 20, Fennoscandia and more western Russia) wintering birds. These results confirm the largely discrete nature of these three flyways and contribute significantly to our ability to deliver effective targeted and appropriate research, monitoring and management actions throughout the ranges of these flyways.     

Tracking Cats: Problems with Placing Feline Carnivores on ��18O, ��D Isoscapes

Background

Several felids are endangered and threatened by the illegal wildlife trade. Establishing geographic origin of tissues of endangered species is thus crucial for wildlife crime investigations and effective conservation strategies. As shown in other species, stable isotope analysis of hydrogen and oxygen in hair (δD_h, δ¹⁸O_h) can be used as a tool for provenance determination. However, reliably predicting the spatial distribution of δD_h and δ¹⁸O_h requires confirmation from animal tissues of known origin and a detailed understanding of the isotopic routing of dietary nutrients into felid hair.

Methodology/Findings

We used coupled δD_h and δ¹⁸O_h measurements from the North American bobcat (Lynx rufus) and puma (Puma concolor) with precipitation-based assignment isoscapes to test the feasibility of isotopic geo-location of felidae. Hairs of felid and rabbit museum specimens from 75 sites across the United States and Canada were analyzed. Bobcat and puma lacked a significant correlation between H/O isotopes in hair and local waters, and also exhibited an isotopic decoupling of δ¹⁸O_h and δD_h. Conversely, strong δD and δ¹⁸O coupling was found for key prey, eastern cottontail rabbit (Sylvilagus floridanus; hair) and white-tailed deer (Odocoileus virginianus; collagen, bone phosphate).

Conclusions/Significance

Puma and bobcat hairs do not adhere to expected pattern of H and O isotopic variation predicted by precipitation isoscapes for North America. Thus, using bulk hair, felids cannot be placed on δ¹⁸O and δD isoscapes for use in forensic investigations. The effective application of isotopes to trace the provenance of feline carnivores is likely compromised by major controls of their diet, physiology and metabolism on hair δ¹⁸O and δD related to body water budgets. Controlled feeding experiments, combined with single amino acid isotope analysis of diets and hair, are needed to reveal mechanisms and physiological traits explaining why felid hair does not follow isotopic patterns demonstrated in many other taxa.     

Dentine oxygen isotopes (δ18O) as a proxy for odontocete distributions and movements

Spatial variation in marine oxygen isotope ratios (δ¹⁸O) resulting from differential evaporation rates and precipitation inputs is potentially useful for characterizing marine mammal distributions and tracking movements across δ¹⁸O gradients. Dentine hydroxyapatite contains carbonate and phosphate that precipitate in oxygen isotopic equilibrium with body water, which in odontocetes closely tracks the isotopic composition of ambient water. To test whether dentine oxygen isotope composition reliably records that of ambient water and can therefore serve as a proxy for odontocete distribution and movement patterns, we measured δ¹⁸O values of dentine structural carbonate (δ¹⁸O_SC) and phosphate (δ¹⁸O_P) of seven odontocete species (n = 55 individuals) from regional marine water bodies spanning a surface water δ¹⁸O range of several per mil. Mean dentine δ¹⁸O_SC (range +21.2 to +25.5‰ VSMOW) and δ¹⁸O_P (+16.7 to +20.3‰) values were strongly correlated with marine surface water δ¹⁸O values, with lower dentine δ¹⁸O_SC and δ¹⁸O_P values in high‐latitude regions (Arctic and Eastern North Pacific) and higher values in the Gulf of California, Gulf of Mexico, and Mediterranean Sea. Correlations between dentine δ¹⁸O_SC and δ¹⁸O_P values with marine surface water δ¹⁸O values indicate that sequential δ¹⁸O measurements along dentine, which grows incrementally and archives intra‐ and interannual isotopic composition over the lifetime of the animal, would be useful for characterizing residency within and movements among water bodies with strong δ¹⁸O gradients, particularly between polar and lower latitudes, or between oceans and marginal basins.     

Linking breeding and wintering grounds of neotropical migrant songbirds using stable hydrogen isotopic analysis of feathers

 Recent studies have shown that stable hydrogen isotope ratios (δD) in the tissues of animals often correlate with δD of local precipitation. Here we examined the relationship between δD in feathers and growing season precipitation for neotropical migrant songbirds breeding over a continent-wide isotopic gradient. δD values were determined on feathers of 140 individuals of 6 species of wild insectivorous forest songbirds (Setophaga ruticilla, Empidonax minimus, Vermivora peregrinus, Catharus ustulatus, Seiurus aurocapillus, Hylocichla mustelina) taken from 14 breeding locations across North America. The δD of feathers was strongly correlated with the δD of growing season precipitation at breeding sites across North America. As feather hydrogen is metabolically inert after growth, this relationship was then used to assess the breeding origins of wintering migrants. Deuterium values of feathers from 64 individuals representing 5 species of migrants (Helmitheros vermivorus, Wilsonia citrina, Hylocichla mustelina, Dumetella carolinensis, Seirus aurocapillus) at a wintering site in Guatemala were consistent with those predicted from the known breeding ranges of these species. This study demonstrates hydrogen isotopes may become a powerful tool for linking breeding and wintering grounds of neotropical migrant songbirds, as well as other migratory species moving between isotopically distinct regions. Received: 25 August 1995 / Accepted: 28 May 1996     

Using stable hydrogen isotopes (δ2H) and ring recoveries to trace natal origins in a Eurasian passerine with a migratory divide

Despite recent advances in technology, it remains difficult to connect breeding and non‐breeding areas of populations of migratory organisms due to the challenges of year‐round tracking. Here, we used the Eurasian reed warbler Acrocephalus scirpaceus, a passerine with a pronounced migratory divide to demonstrate the promise of integrating several sources of information within the Bayesian modelling framework for the study of migratory connectivity. To this end, we combined data from stable hydrogen isotope ratios (δ²H) of feathers, ring recoveries, and the geographic delineation of sub‐populations on either side of the migratory divide. Feather δ²H measurements from local juvenile birds sampled across the breeding range tightly correlated with amount‐weighted mean annual precipitation δ²H values predicted for the natal sites. Predicted natal origins of birds intercepted en route in the Mediterranean region largely differed among the five stopover sites. Thanks to the different migratory pathways used by different breeding populations and the existence of a migratory divide, we were able to effectively narrow the assigned regions of origin. Our results show that spatial resolution of likelihood‐based assignments of geographic origins based on δ²H measurements may improve significantly when prior probabilities derived from population‐specific migratory directions are included. Integrating information from stable isotopes, ring recoveries, geolocators and other sources within the Bayesian modelling framework will provide an extremely useful toolbox for the study of animal movements in the future.     

Origins of irruptive migrations by Common Crossbills Loxia curvirostra into northwestern Europe revealed by stable isotope analysis

We used analyses of stable hydrogen isotope (δ²H) measurements in Common Crossbill feathers (δ²H_f) to infer the region of origin of Crossbills collected from different irruptions into Britain, Iceland and the Faeroes, comparing these values with those from birds sampled in breeding areas in Britain and elsewhere in the western Palaearctic. No differences in δ²H_f values were found between different species or sexes of Crossbills that could be presumed to have grown their feathers in the same region, but juveniles had lower δ²H_f values than adults that had grown their feathers in the same region. On the basis mainly of museum skins, immigrant birds were sampled from 30 different irruption years, spanning the period 1866–2009, with annual samples varying from one to 29 individuals. The variation in δ²H_f values within irruptions was substantially less than the variation between irruptions, indicating that irruptions in different years originated in different parts of the western Palaearctic boreal zone. Birds with lower δ²H_f values tended to arrive later in the migration season, which was consistent with the idea that they had travelled further. In 17 of the irruption years, the birds had mean δ²H_f values more than ?120‰, suggesting that they had originated somewhere in the region extending from northern Scandinavia to northwestern Russia. In these years the birds arrived early, in June and July. In 10 of the irruption years, the mean δ²H_f values were between ?120 and ?130‰, suggesting origins further east, in northern Russia, east of Archangel (about 40°E). In only three of the 30 years (1898, 2002, 2009) the mean δ²H_f values were even lower (< 130‰), and these birds arrived in late July, August and September. Birds in these three irruptions had probably come from Siberia, east of the Ural Mountains. In at least three irruption years (1898, 1927, 1985) the observed range of δ²H_f values suggested that birds had come from more than one of these regions, including east of the Urals in 1898 and 1927.     

Stable isotope analysis of modern human hair collected from Asia (China,India, Mongolia,and Pakistan)

We report isotopic data (δ²H, δ¹⁸O n = 196; δ¹³C, δ¹⁵N n = 142; δ³⁴S n = 85) from human hair and drinking water (δ²H, δ¹⁸O n = 67) collected across China, India, Mongolia, and Pakistan. Hair isotope ratios reflected the large environmental isotopic gradients and dietary differences. Geographic information was recorded in H and O and to a lesser extent, S isotopes. H and O data were entered into a recently developed model describing the relationship between the H and O isotope composition of human hair and drinking water in modern USA and pre‐globalized populations. This has anthropological and forensic applications including reconstructing environment and diet in modern and ancient human hair. However, it has not been applied to a modern population outside of the USA, where we expect different diet. Relationships between H and O isotope ratios in drinking water and hair of modern human populations in Asia were different to both modern USA and pre‐globalized populations. However, the Asian dataset was closer to the modern USA than to pre‐globalized populations. Model parameters suggested slightly higher consumption of locally producedfoods in our sampled population than modern USA residents, but lower than pre‐globalized populations. The degree of in vivo amino acid synthesis was comparable to both the modern USA and pre‐globalized populations. C isotope ratios reflected the predominantly C₃‐based regional agriculture and C₄ consumption in northernChina. C, N, and S isotope ratios supported marine food consumption in some coastal locales. N isotope ratios suggested a relatively low consumption of animal‐derived products compared to western populations. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Evaporation and carbonic anhydrase activity recorded in oxygen isotope signatures of net CO2 fluxes from a Mediterranean soil

The oxygen stable isotope composition (δ¹⁸O) of CO₂ is a valuable tool for studying the gas exchange between terrestrial ecosystems and the atmosphere. In the soil, it records the isotopic signal of water pools subjected to precipitation and evaporation events. The δ¹⁸O of the surface soil net CO₂ flux is dominated by the physical processes of diffusion of CO₂ into and out of the soil and the chemical reactions during CO₂–H₂O equilibration. Catalytic reactions by the enzyme carbonic anhydrase, reducing CO₂ hydration times, have been proposed recently to explain field observations of the δ¹⁸O signatures of net soil CO₂ fluxes. How important these catalytic reactions are for accurately predicting large‐scale biosphere fluxes and partitioning net ecosystem fluxes is currently uncertain because of the lack of field data. In this study, we determined the δ¹⁸O signatures of net soil CO₂ fluxes from soil chamber measurements in a Mediterranean forest. Over the 3 days of measurements, the observed δ¹⁸O signatures of net soil CO₂ fluxes became progressively enriched with a well‐characterized diurnal cycle. Model simulations indicated that the δ¹⁸O signatures recorded the interplay of two effects: (1) progressive enrichment of water in the upper soil by evaporation, and (2) catalytic acceleration of the isotopic exchange between CO₂ and soil water, amplifying the contributions of ‘atmospheric invasion’ to net signatures. We conclude that there is a need for better understanding of the role of enzymatic reactions, and hence soil biology, in determining the contributions of soil fluxes to oxygen isotope signals in atmospheric CO₂.     

Contributions of evaporation, isotopic non-steady state transpiration and atmospheric mixing on the delta18O of water vapour in Pacific Northwest coniferous forests

Changes in the ²H and ¹⁸O of atmospheric water vapour provide information for integrating aspects of gas exchange within forest canopies. In this study, we show that diurnal fluctuations in the oxygen isotope ratio (δ¹⁸O) as high as 4‰ were observed for water vapour (δ¹⁸O_vp) above and within an old‐growth coniferous forest in the Pacific Northwest region of the United States. Values of δ¹⁸O_vp decreased in the morning, reached a minimum at midday, and recovered to early‐morning values in the late afternoon, creating a nearly symmetrical diurnal pattern for two consecutive summer days. A mass balance budget was derived and assessed for the ¹⁸O of canopy water vapour over a 2‐d period by considering the ¹⁸O‐isoflux of canopy transpiration, soil evaporation and the air entering the canopy column. The budget was used to address two questions: (1) do δ¹⁸O values of canopy water vapour reflect the biospheric influence, or are such signals swamped by atmospheric mixing? and (2) what mechanisms drive temporal variations of δ¹⁸O_vp? Model calculations show that the entry of air into the canopy column resulted in an isotopically depleted ¹⁸O‐isoflux in the morning of day 1, causing values of δ¹⁸O_vp to decrease. An isotopically enriched ¹⁸O‐isoflux resulting from transpiration then offset this decreased δ¹⁸O_vp later during the day. Contributions of ¹⁸O‐isoflux from soil evaporation were relatively small on day 1 but were more significant on day 2, despite the small H₂¹⁶O fluxes. From measurements of leaf water volume and sapflux, we determined the turnover time of leaf water in the needles of Douglas‐fir trees as ≈ 11 h at midday. Such an extended turnover time suggests that transpiration may not have occurred at the commonly assumed isotopic steady state. We tested a non‐steady state model for predicting δ¹⁸O of leaf water. Our model calculations show that assuming isotopic steady state increased isoflux of transpiration. The impact of this increase on the modelled δ ¹⁸O_vp was clearly detectable, suggesting the importance of considering isotopic non‐steady state of transpiration in studies of forest ¹⁸O water balance.     

18O composition of CO2 and H2O ecosystem pools and fluxes in a tallgrass prairie: Simulations and comparisons to measurements

In this paper we describe measurements and modeling of ¹⁸O in CO₂ and H₂O pools and fluxes at a tallgrass prairie site in Oklahoma. We present measurements of the δ¹⁸O value of leaf water, depth‐resolved soil water, atmospheric water vapor, and Keeling plot δ¹⁸O intercepts for net soil‐surface CO₂ and ecosystem CO₂ and H₂O fluxes during three periods of the 2000 growing season. Daytime discrimination against C¹⁸OO, as calculated from measured above‐canopy CO₂ and δ¹⁸O gradients, is also presented. To interpret the isotope measurements, we applied an integrated land‐surface and isotope model (ISOLSM) that simulates ecosystem H₂¹⁸O and C¹⁸OO stocks and fluxes. ISOLSM accurately predicted the measured isotopic composition of ecosystem water pools and the δ¹⁸O value of net ecosystem CO₂ and H₂O fluxes. Simulations indicate that incomplete equilibration between CO₂ and H₂O within C₄ plant leaves can have a substantial impact on ecosystem discrimination. Diurnal variations in the δ¹⁸O value of above‐canopy vapor had a small impact on the predicted δ¹⁸O value of ecosystem water pools, although sustained differences had a large impact. Diurnal variations in the δ¹⁸O value of above‐canopy CO₂ substantially affected the predicted ecosystem discrimination. Leaves dominate the ecosystem ¹⁸O‐isoflux in CO₂ during the growing season, while the soil contribution is relatively small and less variable. However, interpreting daytime measurements of ecosystem C¹⁸OO fluxes requires accurate predictions of both soil and leaf ¹⁸O‐isofluxes.     

Feather deuterium as an indicator of age‐class in the Pectoral Sandpiper Calidris melanotos

We measured deuterium isotope ratios (δD_f) in primary feathers to distinguish first‐year from older Pectoral Sandpipers Calidris melanotos captured in Barrow, Alaska, during the breeding season. δD_f showed a distinct bimodal distribution, and model‐based clustering placed the δD_f values into two non‐overlapping groups. More negative δD_f corresponded to Arctic areas, probably identifying first‐year birds with Arctic‐grown juvenile feathers retained from the previous year. The more positive values corresponded to lower latitudes, possibly identifying older birds that grew their feathers at non‐Arctic latitudes.     

Stable Isotope Ecology of Extant Tapirs from the Americas

Understanding the ecology of modern and ancient forests can help clarify the evolution of forest dwelling mammals. It is first necessary, however, to elucidate the source and extent of stable isotope variation in forest taxa. Tapirs are of particular interest because they are model organisms for identifying forest environments due to their highly conservative diet and habitat preferences. Here, stable carbon and oxygen isotopic compositions of extant tapirs are quantified to test hypotheses regarding ontogenetic diet shifts, stable isotope variation at the population level, and relationships between stable isotopes and climatic variables. A population of extant tapirs (Tapirus bairdii) demonstrates low δ¹³C variation (～2–3‰) and increased δ¹³C values in late erupting teeth, indicating that juveniles consume ¹³C deplete milk and/or browse in denser forests. Disparate δ¹⁸O values of late erupting teeth are instead reflecting seasonal variation. Extant tapir (T. bairdii, Tapirus pinchaque, Tapirus terrestris) δ¹⁸O values are constrained by climatic and geographic variables. Most notably, δ¹⁸O values of T. bairdii decrease with decreasing precipitation frequency. Tapirus terrestris is typically present in areas with greater precipitation than T. bairdii and δ¹⁸O values instead positively correlate with δ¹³C values. These data indicate that tapirs in wetter areas are getting a larger proportion of their water from leaves experiencing less evaporation in denser canopies, while T. bairdii is interpreted to increase its consumption of water via drinking when present in drier areas. An understanding of extant tapir stable isotope ecology improves ecological interpretations of these elusive mammals both today and in the past. Abstract in Spanish is available at http://www.blackwell‐synergy.com/loi/btp .     

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.     

Origins of harvested American black ducks: stable isotopes support the flyover hypothesis

Waterfowl management is more effective when based on detailed information on population connectivity between breeding, wintering, and stopover sites. For the American black duck (Anas rubripes), a species of conservation concern, estimates for the fall age ratio at harvest differed depending on whether harvest data were derived from Canada or the United States, suggesting regional differences. Within Canada, hunters in Atlantic Canada were more likely to harvest black ducks from nearby breeding locations compared to hunters in southern Ontario and Quebec, Canada, who were more likely to harvest individuals from the Boreal Softwood and Taiga Shield of eastern Canada. Black ducks harvested in the United States are thought to originate predominantly from northern portions of the breeding range, leading to the flyover hypothesis, which postulates that black ducks produced in the Boreal Softwood and Taiga Shield region are less susceptible to harvest by hunters in Atlantic Canada and northeastern United States. To test the flyover hypothesis, we examined regional and temporal differences in the origins of harvested black ducks using feathers from wings (n = 664) submitted by hunters to the species composition and parts collection surveys across 3 hunting seasons (2017–2018, 2018–2019, 2019–2020). We used a likelihood-based assignment method that relied on feather stable-hydrogen isotopes (δ²H) and stable-carbon isotopes (δ¹³C) to determine the natal or molt origin of individuals harvested within eastern Canada and the United States. We also used a spatial clustering technique to group harvested individuals by area of origin without a priori knowledge of such regions. Adult female black ducks originated farther south compared to males and juveniles. All sexes and ages of black ducks harvested in Atlantic Canada showed predominantly southern origins, while those harvested in the United States and other Canadian provinces primarily originated farther north within the boreal, supporting the flyover hypothesis. By contrast, we found no relationship between timing of harvest or peaks of migration and individual origin. After combining band returns and stable isotopes, we inferred 2 distinct stocks: the Mississippi flyway stock and the Atlantic flyway stock. We recommend that regional demographic parameters, particularly for Atlantic Canada, be directly measured to promote more effective conservation of black ducks and optimize harvest opportunities in the United States and Canada.     

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 (δ²H_f) 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.