A test of geographic assignment using isotope tracers in feathers of known origin

We used feathers of known origin collected from across the breeding range of a migratory shorebird to test the use of isotope tracers for assigning breeding origins. We analyzed δD, δ¹³C, and δ¹⁵N in feathers from 75 mountain plover (Charadrius montanus) chicks sampled in 2001 and from 119 chicks sampled in 2002. We estimated parameters for continuous-response inverse regression models and for discrete-response Bayesian probability models from data for each year independently. We evaluated model predictions with both the training data and by using the alternate year as an independent test dataset. Our results provide weak support for modeling latitude and isotope values as monotonic functions of one another, especially when data are pooled over known sources of variation such as sample year or location. We were unable to make even qualitative statements, such as north versus south, about the likely origin of birds using both δD and δ¹³C in inverse regression models; results were no better than random assignment. Probability models provided better results and a more natural framework for the problem. Correct assignment rates were highest when considering all three isotopes in the probability framework, but the use of even a single isotope was better than random assignment. The method appears relatively robust to temporal effects and is most sensitive to the isotope discrimination gradients over which samples are taken. We offer that the problem of using isotope tracers to infer geographic origin is best framed as one of assignment, rather than prediction.     

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     

The use of isotope tracers for identifying populations of migratory birds

 To determine whether stable isotopes can be used for identifying the geographic origins of migratory bird populations, we examined the isotopic composition of hydrogen (deuterium, δD), carbon (δ¹³C), and strontium (δ⁸⁷Sr) in tissues of a migratory passerine, the black-throated blue warbler (Dendroica caerulescens), throughout its breeding range in eastern North America. δD and δ¹³C values in feathers, which are grown in the breeding area, varied systematically along a latitudinal gradient, being highest in samples from the southern end of the species’ breeding range in Georgia and lowest in southern Canada. In addition, δD decreased from east to west across the northern part of the breeding range, from New Brunswick to Michigan. δ⁸⁷Sr ratios were highest in the Appalachian Mountains, and decreased towards the west. These patterns are consistent with geographical variation in the isotopic composition of the natural environment, i.e., with that of precipitation, plants, and soils for δD, δ¹³C, and δ⁸⁷Sr, respectively. Preliminary analyses of the δD and δ¹³C composition of feathers collected from warblers in their Caribbean winter grounds indicate that these individuals were mostly from northern breeding populations. Furthermore, variances in isotope ratios in samples from local areas in winter tended to be larger than those in summer, suggesting that individuals from different breeding localities may mix in winter habitats. These isotope markers, therefore, have the potential for locating the breeding origins of migratory species on their winter areas, for quantifying the degree of mixing of breeding populations on migratory and wintering sites, and for documenting other aspects of the population structure migratory animals – information needed for studies of year-round ecology of these species as well as for their conservation. Combining information from several stable isotopes will help to increase the resolution for determining the geographic origins of individuals in such highly vagile populations. Received: 24 April 1995 / Accepted: 2 June 1996     

Linking isotopic and migratory patterns in a pelagic seabird

The value of stable isotope analysis in tracking animal migrations in marine environments is poorly understood, mainly due to insufficient knowledge of isotopic integration into animal tissues within distinct water masses. We investigated isotopic and moult patterns in Cory’s shearwaters to assess the integration of different stable isotopes into feathers in relation to the birds’ transoceanic movements. Specimens of Mediterranean Cory’s shearwater Calonectris diomedea diomedea caught accidentally by Catalan longliners were collected and the signatures of stable isotopes of C (δ¹³C), N (δ¹⁵N) and S (δ³⁴S) were analysed in 11 wing and two tail feathers from 20 birds, and in some breast feathers. Based on isotopic signatures and moult patterns, the feathers segregated into two groups (breeding and wintering), corresponding to those grown in the Mediterranean or Atlantic regions, respectively. In addition, feathers grown during winter, i.e. moulted in Atlantic waters, were grouped into two isotopically distinct profiles, presumably corresponding to the two main wintering areas previously identified for Mediterranean Cory’s shearwater in tracking studies. N signatures mainly indicated the Mediterranean-to-Atlantic migration, whereas C and S signatures differed according to the Atlantic wintering area. Our results indicate that isotopic signatures from distant oceanic regions can integrate the feathers of a given bird and can indicate the region in which each feather was grown. This study thus underscores how stable isotope analysis can link marine animals to specific breeding and wintering areas, and thereby shed new light on studies involving assignment, migratory connectivity and carry-over effects in the marine environment. Xavier Ruiz deceased 27 April 2008.     

Comparison of trace element and stable isotope approaches to the study of migratory connectivity: an example using two hirundine species breeding in Europe and wintering in Africa

Analyses of stable isotopes and trace elements in feathers may provide important information about location and habitat use during molt, thereby enabling the investigation of migratory connectivity and its ecological consequences in bird species that breed and winter in different areas. We have compared the conclusions arrived at based on the use of these two methods on the same samples of feathers from two migratory birds, the Sand Martin Riparia riparia and the Barn Swallow Hirundo rustica. We investigated the effects of location, age and sex on stable isotope (δ¹³C, δ¹⁵N, δD) and trace element profiles (As, Cd, Mg, Mn, Mo, Se, Sr, Co, Fe, Zn, Li, P, Ti, V, Ag, Cr, Ba, Hg, Pb, S, Ni and Cu). The feathers of adults at the breeding grounds were removed, forcing in birds to grow new feathers at the breeding grounds; this enabled a comparison of composition of feathers grown in Europe and Africa by the same individual. Stable isotope and trace element profiles varied geographically, even at micro-geographic scales, and also among age classes. The results of both methods suggest that food composition and/or source differs between adults and nestlings in the breeding area and that food and/or molting location changes with the age of individuals in Africa. In an attempt to determine the usefulness of data obtained from composition of feathers, we performed discriminant function analyses on information obtained on stable isotopes and trace elements in order to assess the correctness of the classification of group membership. When feathers molted in Africa were compared to those molted in Europe, trace element profiles of the 22 elements generally had a much greater resolution than the stable isotope profiles based on three stable isotopes. The proportion of correctly classified samples was also greater for analyses based on trace elements than for those based on stable isotopes.     

On the use of stable oxygen isotope (δ18O) measurements for tracking avian movements in North America

Tracking migratory animals has benefitted using measurements of naturally occurring stable isotopes of hydrogen (δ²H) in keratinous tissues such as hair and feathers to link animal origins to continental patterns or isoscapes of δ²H in precipitation. However, for most taxa, much less information exists on the use of stable oxygen isotope ratios (δ¹⁸O) despite the fact that δ²H and δ¹⁸O are strongly linked in environmental waters through the meteoric relationship and the possibility of using both isotopes to infer greater information on origins and climatic conditions where tissues are grown. A fundamental requirement of using stable isotopes to assign individuals and populations to origins is the development of a rescaling function linking environmental food web signals to the tissue of interest and for birds, this has not been carried out. Here, we derived the relationship between H and O isotopes in known source feathers of 104 individuals representing 11 species of insectivorous passerines sampled across the strong precipitation isoscape of North America. We determined again a strong expected relationship between feather δ²H (δ²H_f) and long‐term amount‐weighted precipitation δ²H (δ²H_p; r² = 0.77), but the corresponding relationship between δ¹⁸O_f and δ¹⁸O_p was poor (r² = 0.32) for the same samples. This suggests that δ²H measurements are currently more useful for assignment of insectivorous songbirds to precipitation isoscapes but does not preclude other uses of the δ¹⁸O_f data. Currently, mechanisms responsible for the decoupling of H and O isotopes in food webs is poorly known, and we advocate a much broader sampling of both isotopes in the same keratinous tissues across precipitation isotope gradients and across taxa to resolve this issue and to increase the power of using water isotopes to track migratory animals.     

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

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

Fundamental limits to the accuracy of deuterium isotopes for identifying the spatial origin of migratory animals

Deuterium isotope analyses have revolutionized the study of migratory connectivity because global gradients of deuterium in precipitation (δD_P) are expressed on a continental scale. Several authors have constructed continental scale base maps of δD_P to provide a spatial reference for studying the movement patterns of migratory species and, although they are very useful, these maps present a static, 40-year average view of the landscape that ignores much underlying inter-annual variation. To more fully understand the consequences of this underlying variation, we analyzed the GNIP deuterium data, the source for all current δD_P maps, to estimate the minimum separation in δD_P (and latitude) necessary to conclude with a given level of confidence that distinct δD_P values represent different geographic sites. Extending analyses of δD_P successfully to deuterium in tissues of living organisms, e.g., feathers in migratory birds (δD_F), is dependent on the existence of geographic separation of δD_P, where every geographic location has a distribution of values associated with temporal variability in δD_P. Analyses were conducted for three distinct geographic regions: North America, eastern North America (east of longitude 100°W), and Argentina. At the 80% confidence level, the minimum separation values were 12, 7, and 14° of latitude (equivalent to 53, 31, and 32‰) for North America, eastern North America, and Argentina, respectively. Hence, in eastern North America, for example, one may not be able to accurately assign individual samples to sites separated by less than about 7° of latitude as the distributions of δD_P were not distinct at latitudes <7° apart. Moreover, two samples that differ by less than 31‰ cannot be confidently said to originate from different latitudes. These estimates of minimum separation for δD_P do not include other known sources of variation in feather deuterium (δD_F) and hence are a first order approximation that may be useful, in the absence of more specific information for the system of interest, for planning and interpreting the results of new stable isotope studies.     

Birds of a feather winter together: migratory connectivity in the Reed Warbler Acrocephalus scirpaceus

To investigate migratory connectivity in the Reed Warbler Acrocephalus scirpaceus, we analysed (1) all available sub-Saharan ringing recoveries and (2) stable isotopes in feathers grown in Africa sampled at 17 European breeding sites across a migratory divide. A cluster analysis of ringing recoveries showed remarkable connectivity between breeding and non-breeding grounds. Two main clusters represented populations taking the two main migratory routes [southwesterly (SW) and southeasterly (SE)]. Stable isotope analysis confirmed the separation of wintering areas of SW- and SE-migrating populations. Higher δ¹⁵N values in feathers of SE-migrating birds indicated that they occupied more xeric biome types. Values of δ¹³C that did not differ significantly among populations were higher than those from feathers of known European origin and indicated a C4 biome. Three populations with an unknown migratory direction were assigned to the SE-migrating populations on the basis of δ¹⁵N values.     

Stable isotopes (δD and δ13C) are geographic indicators of natal origins of monarch butterflies in eastern North America

Wing membranes of laboratory and field-reared monarch butterflies (Danaus plexippus) were analyzed for their stable-hydrogen (δD) and carbon (δ¹³C) isotope ratios to determine whether this technique could be used to identify their natal origins. We hypothesized that the hydrogen isotopic composition of monarch butterfly wing keratin would reflect the hydrogen isotope patterns of rainfall in areas of natal origin where wings were formed. Monarchs were reared in the laboratory on milkweed plants (Asclepias sp.) grown with water of known deuterium content, and, with the assistance of volunteers, on native milkweeds throughout eastern North America. The results show that the stable hydrogen isotopic composition of monarch butterflies is highly correlated with the isotopic composition of the milkweed host plants, which in turn corresponds closely with the long-term geographic patterns of deuterium in rainfall. Stable-carbon isotope values in milkweed host plants were similarly correlated with those values in monarch butterflies and showed a general pattern of enrichment along a southwest to northeast gradient bisecting the Great Lakes. These findings indicate that natal origins of migratory and wintering monarchs in Mexico can be inferred from the combined δD and δ¹³C isotopic signatures in their wings. This relationship establishes that analysis of hydrogen and carbon isotopes can be used to answer questions concerning the biology of migratory monarch butterflies and provides a new approach to tracking similar migratory movements of other organisms. Received: 1 July 1998 / Accepted: 11 November 1998     

Hydrogen isotopic variation in migratory bird tissues of known origin: implications for geographic assignment

Continent-wide variation in hydrogen isotopic composition of precipitation is incorporated into animal diets, providing an intrinsic marker of geographic location at the time of tissue growth. Feathers from migratory birds are now frequently analyzed for stable-hydrogen isotopes (δD) to estimate the location of individuals during a preceding molt. Using known-origin birds, we tested several assumptions associated with this emerging technique. We examined hydrogen isotopic variation as a function of age, sex, feather type and the timing of molt in a marked population of American redstarts (Setophaga ruticilla) breeding in southeastern Ontario. We measured δD in feathers and blood from individuals that bred or hatched at our study site during the year in which those tissues were grown. Juvenile tissues from 5- to 10-day-old birds had more negative δD values than those from adults, which most likely reflected age-related differences in diet. Within adults, primary feathers had more negative δD values than contour feathers. The mean δD value in adult primary feathers was relatively consistent among years and with the value expected for our study population. However, among-individual variation in δD corresponded to an estimated latitudinal range of 6–8° (650–900 km). We conclude that feathers sampled from recently hatched juveniles may not provide a reliable estimate of expected local isotopic signatures for comparison with adult feathers of unknown origin. Furthermore, we urge researchers to use caution when using δD values in feathers to infer geographic origin, and suggest that the best approach is to assign individuals to broad geographic zones within a species’ potential molting range.     

Stable isotopes identify the natal origins of a generalist brood parasite, the brown-headed cowbird Molothrus ater

Identifying the natal origins of brood parasites is a major challenge that usually requires labor-intensive searching for nests of host species. Stable isotope analysis of feathers and other body tissues of parasitic young could be a possible tool for determining natal origins if tissues reflect the isotopic composition of the diet fed to nestlings. We measured the carbon (¹³C) and nitrogen (¹⁵N) isotope compositions of feathers for two age-classes of brown-headed cowbirds Molothrus ater at the Konza Prairie Biological Station near Manhattan, Kansas: nestlings raised by five species of songbird hosts in two different habitats, and juveniles captured after independence. Isotope values from cowbird nestlings did not differ among host species and we were unable to assign juvenile cowbirds to their natal hosts. However, nestlings raised in grassland habitat had feathers that contained significantly higher δ¹³C values and lower δ¹⁵N values than nestlings raised in shrub habitats. In addition, independent juveniles had isotopic signatures that were similar to cowbird nestlings raised on shrub habitats. Although dickcissel Spiza americana comprised the majority of samples from shrub habitats, our conclusions reflect the natural pattern of parasitism at the site and should be representative of cowbirds raised at Konza. We conclude that stable isotope analysis of feathers is effective for determining the natal origins of parasitic young if isotope values from nestlings are isotopically distinct among habitats.     

Does avian malaria infection affect feather stable isotope signatures?

It is widely accepted that stable isotope ratios in inert tissues such as feather keratin reflect the dietary isotopic signature at the time of the tissue synthesis. However, some elements such as stable nitrogen isotopes can be affected by individual physiological state and nutritional stress. Using malaria infection experiment protocols, we estimated the possible effect of malaria parasite infections on feather carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope signatures in juvenile common crossbills Loxia curvirostra. The birds were experimentally infected with Plasmodium relictum (lineage SGS1) and P. ashfordi (GRW2), two widespread parasites of passerines. Experimental birds developed heavy parasitemia of both parasites and maintained high levels throughout the experiment (33 days). We found no significant difference between experimental and control birds in both δ¹³C and δ¹⁵N values of feathers re-grown. The study shows that even heavy primary infections of malaria parasites do not affect feather δ¹³C and δ¹⁵N isotopic signatures. The results of this experiment demonstrate that feather isotope values of wild-caught birds accurately reflect the dietary isotopic sources at the time of tissue synthesis even when the animal’s immune system might be challenged due to parasitic infection.     

Estimating the latitudinal origins of migratory birds using hydrogen and sulfur stable isotopes in feathers: influence of marine prey base

Hydrogen stable isotope analysis of feathers is an important tool for estimating the natal or breeding latitudes of nearctic-neotropical migratory birds. This method is based on the latitudinal variation of hydrogen stable isotope ratios in precipitation in North America (deltaD(p)) and the inheritance of this variation in newly formed feathers (deltaD(f)). We hypothesized that the typically strong relationship between deltaD(p) and deltaD(f) would be decoupled in birds that forage in marine food webs because marine waters have relatively high deltaD values compared to deltaD values for local precipitation. Birds that forage on marine prey bases should also have feathers with high delta(34)S values, since delta(34)S values for marine sulfate are generally higher than delta(34)S values in terrestrial systems. To examine this potential marine effect on feather stable isotope ratios, we measured deltaD and delta(34)S in the feathers of nine different species of raptors from both inland and coastal locations across North America. Feathers from coastal bird-eating raptors had consistently higher deltaD and delta(34)S values than feathers from inland birds. Birds that had high delta(34)S values also deviated strongly from the typical relationship between deltaD(p) and deltaD(f). We recommend measuring both sulfur and hydrogen stable isotope ratios in feathers when some members of a migrant population could potentially forage in marine habitats. We suggest using a practical cutoff of delta(34)S >10 per thousand to remove marine-foraging birds from a migrant sample when using stable isotopes of hydrogen to estimate the latitudinal origins of migrants because high deltaD(f) values for marine-foraging birds could potentially distort estimates of origins.     

Conservation genetics of snowy plovers (<Emphasis Type="Italic">Charadrius alexandrinus</Emphasis>) in the Western Hemisphere: population genetic structure and delineation of subspecies

We examined the genetic structure of snowy plovers (Charadrius alexandrinus) in North America, the Caribbean, and the west coast of South America to quantify variation within and among breeding areas and to test the validity of three previously recognized subspecies. Sequences (676 bp) from domains I and II of the mitochondrial control region were analyzed for 166 snowy plovers from 20 breeding areas. Variation was also examined at 10 microsatellite loci for 144 snowy plovers from 14 breeding areas. The mtDNA and microsatellite data provided strong evidence that the Puerto Rican breeding group is genetically divergent from sites in the continental U.S. (net sequence divergence = 0.38%; F _ST for microsatellites = 0.190). Our data also revealed high levels of differentiation between sites from South America and North America (net sequence divergence = 0.81%; F _ST for microsatellites = 0.253). In contrast, there was little genetic structure among breeding sites within the continental U.S. Our results suggest that snowy plovers in Florida should be considered part of C. a. nivosus (rather than part of C. a. tenuirostris, where they are currently placed), whereas snowy plovers from Puerto Rico should be considered part of C. a. tenuirostris. Snowy plovers in South America should remain a separate subspecies (C. a. occidentalis). Although U.S. Pacific and Gulf Coast breeding areas were not genetically distinct from other continental U.S. sites, demographic isolation, unique coastal habitats, and recent population declines suggest they warrant special concern.     

Global application of stable hydrogen and oxygen isotopes to wildlife forensics

Stable isotopes are being increasingly used in wildlife forensics as means of determining the origin and movement of animals. The heavy isotope content of precipitated water and snow (D_p, ¹⁸O_p) varies widely and systematically across the globe, providing a label that is incorporated through diet into animal tissue. As a result, these isotopes are potentially ideal tracers of geographic origin. The hydrogen and oxygen isotope tracer method has excellent potential where (1) spatial variation of precipitation isotopes exist, and (2) strong, mechanistic relationships link precipitation and isotope ratios in biological tissue. Here, we present a method for interpolation of precipitation isotope values and use it to create global basemaps of growing-season (GS) and mean annual (MA) D_p and ¹⁸O_p. The use of these maps for forensic application is demonstrated using previously published isotope data for bird feathers (D_f) in North America and Europe. The precipitation maps show that the greatest potential for applying hydrogen and oxygen isotope forensics exists in mid- to high-latitude continental regions, where strong spatial isotope gradients exist. We demonstrate that D_f/D_p relationships have significant predictive power both in North America and Europe, and show how zones of confidence for the assignment of origin can be described using these predictive relationships. Our analysis focuses on wildlife forensics, but the maps and approaches presented here will be equally applicable to criminal forensic studies involving biological materials. These maps are available in GIS format at .     

The influence of breeding colony and sex on mercury,selenium and lead levels and carbon and nitrogen stable isotope signatures in summer and winter feathers of <Emphasis Type="Italic">Calonectris</Emphasis> shearwaters

Contamination in marine foodwebs is nowadays of great environmental concern owing to the increasing levels of pollution in marine ecosystems from different anthropogenic sources. Seabirds can be used as indicators of regional contaminant patterns across large temporal and spatial scales. We analysed Hg, Se and Pb levels as well as stable isotope ratios of C (¹³C/¹²C, δ¹³C) and N (¹⁵N/¹⁴N, δ¹⁵N) in breeding- and winter-season feathers on males and females of two related shearwater species, providing information on spatiotemporal patterns of contaminants as well as the influence of the trophic ecology of these seabirds on contaminant levels. During the breeding season, Se and Pb concentrations were highest at the Cape Verde archipelago, showing no differences among the other colonies or between the sexes. However, Hg levels varied among colonies, being highest in the Mediterranean, probably resulting from the larger emissions and fallout of this pollutant in Europe. Feathers grown during breeding also showed sexual differences in Hg concentrations and δ¹³C. Differences in Hg concentration between sexes are mainly due to egg-laying decontamination in females. In contrast, differences in Hg among colonies are probably related to differences in trophic ecology, as indicated by δ¹³C and δ¹⁵N measurements. Contaminant concentrations in winter-grown feathers did not show any relationship with stable isotope values but were affected by contaminant loads associated with the breeding season. These findings suggest that the interpretation of contaminant levels of migratory species from feathers moulted out of the breeding season should be made with caution because those values could reflect exposures to contaminants acquired during the breeding season. We conclude that factors other than feeding ecology may play an important role in the interpretation of contaminant levels and their annual dynamics at several spatial scales. Consideration of the relevant temporal context provided by isotopic signatures and contaminant concentrations is important in deciphering contaminant information based on various tissues. Xavier Ruiz: deceased 27 April 2008.     

Trophic ecology of Pacific salmon (<Emphasis Type="Italic">Oncorhynchus</Emphasis> spp.) in the ocean: a synthesis of stable isotope research

Increasing interest in the marine trophic dynamics of Pacific salmon has been motivated by the recognition of their sensitivity to changing climate and to the competitive effects of hatchery fish on wild stocks. It has become more common to use stable isotopes to supplement traditional diet studies of salmon in the ocean; however, there have been no integrated syntheses of these data to determine whether stable isotope analyses support the existing conventional wisdom of feeding strategies of the Pacific salmon. We performed a meta-analysis of stable isotope data to examine the extent of trophic partitioning among five species of Pacific salmon during their marine lives. Pink, sockeye, and chum salmon showed very high overlap in resource use and there was no consistent evidence for chum relying on alternative food webs dominated by gelatinous zooplankton. δ¹⁵N showed that Chinook and coho salmon fed at trophic levels higher than the other three species. In addition, these two species were distinctly enriched in ¹³C, suggesting more extensive use of coastal food webs compared to the more depleted (pelagic) signatures of pink, sockeye, and chum salmon. This paper presents the first synthesis of stable isotope work on Pacific salmon and provides δ¹⁵N and δ¹³C values applicable to research on the fate of the marine derived nutrients these organisms transport to freshwater and riparian ecosystems.     

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.     

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

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