Incorporating site and year‐specific deuterium ratios (δ2H) from precipitation into geographic assignments of a migratory bird

The study of migratory connectivity is rapidly growing in ornithology, as is the technology used to measure it. While use of extrinsic markers, such as archival tags, is becoming more prevalent, for many small species the best tool available for tracking birds remains intrinsic markers, such as stable‐hydrogen isotope ratios (δ²H). Many researchers have raised concerns that spatial and temporal environmental variation introduces a large amount of error into isotope‐based assignments, limiting their utility. Here, using feathers, we sought to address these issues in developing δ²H base maps for assigning pied flycatchers Ficedula hypoleuca of known origin to 15 sites across the breeding range (approx. 4 020 800 km²). We evaluated the effects of including random site variation and year‐specific precipitation δ²H (δ²H_P) maps on assignments, compared to using mean annual growing season δ²H_p and no site effects. We found a positive correlation between feather δ²H (δ²H_F) and mean annual δ²H_P, resulting from large scale geographic variation. Repeatability of feather δ²H for individuals sampled in multiple years was strong overall, but variable among populations. Annual variation in δ²H_P explained 21% of within individual variation in δ²H_F. Neither year‐ nor site‐specific methods improved assignment precision or accuracy. All three methods assigned flycatchers of unknown origin captured at an African overwintering site to similar breeding areas. However, methods using long‐term means of δ²H_p assigned birds more precisely than year‐specific methods. Our results suggest that annual variation in this system is primarily a result of food web or individual level processes and that random site effects are not strong enough to drastically impact accuracy. We conclude that improvements in isotope based geographic assignments will rely on the addition of prior information, such as relative abundance in a Bayesian framework, or additional intrinsic markers.     

Feather isotope analysis discriminates age-classes of Western, Least, and Semipalmated sandpipers when plumage methods are unreliable

ABSTRACT Avian age‐class discrimination is typically based on the completeness of the first prebasic molt. In several calidrid sandpiper species, juvenal flight feathers grown on Arctic breeding grounds are retained through the first three migrations. Thereafter, flight feathers are grown annually at temperate migratory stopover sites during the fall or on the subtropical wintering grounds. Standard methods for distinguishing age classes of sandpipers rely on a combination of traits, including body plumage, coloration of protected inner median covert edges, and extent of flight feather wear. We tested the ability of stable hydrogen isotope ratios in flight feathers (δD_f) to distinguish young birds in their first winter through second fall from older adults in three calidrid sandpiper species, Western (Calidris mauri), Least (C. minutilla), and Semipalmated (C. pusilla) sandpipers. We compared the apparent reliability of the isotope approach to that of plumage‐based aging. The large expected differences in δD_f values of flight feathers grown at Arctic versus non‐Arctic latitudes enabled use of this technique to discriminate between age‐classes. We determined δD_f values of known Arctic‐grown feathers from juveniles that grew their flight feathers on the breeding grounds. Flight feather δD_f values of southward‐migrating adults showed bimodal distributions for all three species. Negative values overlapped with species‐specific juvenile values, identifying putative second fall birds with high‐latitude grown juvenal feathers retained from the previous year. The more positive values identified older adults who grew their feathers at mid‐ and low latitudes. Importantly, δD_f analysis successfully identified first‐winter and second‐fall birds not detected by plumage‐based aging. Flight feather wear alone was a poor basis for age classification because scores overlapped extensively between putative second fall birds and older adults. Flight feather hydrogen isotope analysis enables more definitive assignment of age classes when standard plumage methods are unreliable.     

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.     

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

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

Stable isotope ratios in winter‐grown feathers of Great Reed Warblers Acrocephalus arundinaceus,Clamorous Reed Warblers A. stentoreus and their hybrids in a sympatric breeding population in Kazakhstan

Analyses of the stable isotope composition of feathers can provide significant insight into the spatial structure of bird migration. We collected feathers from Great Reed Warblers Acrocephalus arundinaceus, Clamorous Reed Warblers A. stentoreus and a small sample of their hybrids in a sympatric breeding population in Kazakhstan to assess natural variation in stable isotope signatures and delineate wintering sites. The Great Reed Warbler is a long‐distance migrant that overwinters in sub‐Saharan Africa, whereas the Clamorous Reed Warbler performs a short‐distance migration to the Indian sub‐continent. Carbon (δ¹³C), nitrogen (δ¹⁵N) and deuterium (δD) isotope signatures were obtained from winter‐grown feathers of adult birds. There were highly significant differences in δD and less significant differences in δ¹³C between Great and Clamorous Reed Warblers. Thus, our results show that the stable isotope technique, and in particular the deuterium (δD) signal, resolves continental variation in winter distribution between these closely related Acrocephalus species with sympatric natal origin. The isotope signatures of hybrid Great × Clamorous Reed Warblers clustered with those of the Great Reed Warblers. Hence, a parsimonious suggestion is that the hybrids undergo moult in Afrotropical wintering grounds, as do the Great Reed Warblers. The observed δD values fell within the range of expected values based on available precipitation data collected at precipitation stations across the wintering continents of each species. However, the power to predict the winter origin of birds in our study system using these data was weak as the expected values ranged widely at this broad continental scale.     

Quantifying structural variation in contour feathers to address functional variation and life history trade‐offs

Body feathers are important to many interactions birds have with their physical and social environments, such as streamlining the body for flight, thermoregulation, and social signaling. Birds differ dramatically in the texture of their body plumage depending on species and age class, likely reflecting different functional demands and age‐related trade‐offs in feather production. Despite the important insights potentially offered by studying variation in the structure of body feathers, there is no clear system for quantifying this variation. We present methods for quantifying age and species differences in the structure of body feathers. Most variation in our measures is due to species and age‐class differences, with little variance attributable to individual birds or to differences among feathers sampled from the same bird. We use our measures to test the hypothesis that the loosely‐textured plumage characteristic of many juvenile passerines reflects a trade‐off between investment in feather quality and rapid body growth that promotes early fledging. The structure of juvenile feathers was negatively correlated with duration of the nestling period among ten species of New World warblers (Parulidae), suggesting a trade‐off between investment in feathers and investment in rapid somatic development promoting fledging. Systematic studies of variation in the structure of body feathers will likely offer numerous other insights into avian biology.     

Rainfall at African wintering grounds predicts age‐specific probability of haemosporidian infection in a migratory passerine bird

In migratory species breeding in temperate zones and wintering in tropical areas, the prevalence of blood parasites may be affected by migratory strategies and winter habitat choice. We explored whether African winter habitat was linked to the probability of haemosporidian infection in the House Martin Delichon urbicum breeding in Spain, and tested for potential differences between age‐classes. As a proxy for winter habitat features, we analysed stable isotope (δ²H, δ¹³C and δ¹⁵N) values of winter‐grown feathers moulted in tropical Africa. Rainfall at the African winter grounds was related to the probability of being infected with haemosporidians and this effect differed among age‐classes. We found that haemosporidian prevalence was similar for young and experienced birds wintering in habitats of higher rainfall (²H‐depleted), whereas there were great differences in winter habitats of lower rainfall (²H‐enriched), with young having a much higher prevalence compared with experienced birds. Likewise, experienced birds wintering in habitats of higher rainfall had a higher probability of haemosporidian infection compared with experienced birds wintering in habitats of lower rainfall. By contrast, young birds wintering in habitats of lower rainfall had a higher probability of haemosporidian infection compared with young birds wintering in habitats of higher rainfall. These outcomes highlight the interaction of age with haemosporidian infection in the migratory ecology of the House Martin, which may drive carry‐over effects in this long‐distance aerial insectivore.     

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.     

Variability in stable isotopes of snowy owl feathers and contribution of marine resources to their winter diet

The snowy owl is an elusive arctic predator known for its nomadic behaviour. Satellite tracking has revealed that some adult snowy owls could make an extensive use of the marine environment during the non‐breeding season. However, the relative contribution of marine resources to their diet is unknown. Stable isotope analyses can be useful to document the diet of mobile animals during periods of the year when individuals are less accessible. This study aimed to assess variation in isotopic values (δ¹³C and δ¹⁵N) of various feather types, and the usefulness of feathers to determine the contribution of the marine environment to the winter diet of snowy owls captured in summer. We sampled feathers coming from 6 body regions of 18 breeding females at two sites in the eastern Canadian Arctic in 2013 and 2014. Prior to analyses, diet‐tissue discrimination factors of snowy owl feathers were established in captivity. Variability in isotopic values among feather types was relatively low and pairwise correlations in isotopic values between feathers on the same individual were variable and often low, which suggests differences in the diet at the time when various feathers were synthesized. Diet reconstruction models detected a contribution of marine sources to snowy owl feathers ranging from 4 to 19% among feather types. However, the marine contribution was highly variable when single feathers were examined within individuals, ranging from 3 to 71%. This indicated that no single feather type could be used alone to reliably infer the contribution of marine resources to the winter diet of owls, possibly due to a high variability in the timing and sequence of molt. For asynchronous molters like snowy owls, we recommend sampling multiple feathers from various body regions, excluding wing feathers, to investigate winter diet or habitat use.     

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.     

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.     

Wing moult and mass change in free‐living mallard Anas platyrhynchos

Captured free‐living male mallard Anas platyrhynchos at Abberton in southern Britain showed peak mass gain immediately prior to simultaneous remex moult. Individuals of both sexes were heavier before shedding wing feathers than when flightless confirming literature accounts that show mallard accumulate fat stores in anticipation of moult to contribute to meeting energy needs during remex re‐growth. Over the course of four seasons, males lost 13 17% of initial body mass on average during re‐growth of flight feathers, females 13 23%. Based on energy expenditure of 1.3 times BMR, male mallard were estimated to be able to fulfil 42 60% and females 41 82% of their energy needs throughout moult from stores. Free‐flying male mallard fed ad libitum in a predator‐free environment did not differ in starting body mass or rate of mass loss during wing moult compared to free‐living Abberton birds, suggesting depletion of fat stores, irrespective of available sources of exogenous energy. Based on this evidence, we reject that the hypotheses that mass loss in moulting mallard is due to 1) simple energy stress and 2) restrictions on feeding and consider that 3) attaining the ability to fly at an earlier stage on incompletely grown flight feathers is not the primary factor shaping this trait. Rather, we consider the accumulation and subsequent depletion of fat stores, together with reductions in energy expenditure, enable mallard to re‐grow feathers as rapidly as possible by exploiting habitats that offer safety from predators, but do not necessarily enable them to balance energy budgets during the flightless period of remex feather re‐growth.     

Trophic level delineation and resource partitioning in a South African afromontane forest bird community using carbon and nitrogen stable isotopes

Southern African forests are naturally fragmented yet hold a disproportionately high number of bird species. Carbon and nitrogen stable isotopes were measured in feathers from birds captured at Woodbush (n = 27 species), a large afromontane forest in the eastern escarpment of Limpopo province, South Africa. The δ¹³C signatures of a range of forest plants were measured to categorise the food base. Most plants sampled, including two of five grass species, had δ¹³C signatures typical of a C₃ photosynthetic pathway (?29.5 ± 1.9‰). Three grass species had a C₄ signature (?12.0 ± 0.6‰). Most bird species had δ¹³C values representing a predominantly C₃‐based diet (?24.8‰ to ?20.7‰). δ¹⁵N values were as expected, with higher levels of enrichment associated with a greater proportion of dietary animal matter. The cohesive isotopic niche defining most species (n = 22), where the ranges for δ¹³C and δ¹⁵N were 2.4‰ and 3.4‰, respectively, highlight the difficulties in understanding diets of birds in a predominantly C₃‐based ecosystem using carbon and nitrogen stable isotopes. However, variation in isotopic values between and within species provides insight into possible niche width and the use of resources by different birds within a forest environment.     

Tracing site‐specific isotopic signatures along a Blue Tit Cyanistes caeruleus food chain

Food chains culminating with temperate insectivorous passerines are well described, yet whether trophic webs can be site‐specific remains a largely unexplored question. In the case of site‐ or habitat‐specificity of food webs, stable isotope signatures of bird feathers may enable assignment of unmarked individuals to a site or a habitat of origin. We address this question in landscapes that include contrasting forest habitat patches with either deciduous Downy Oak Quercus humilis or evergreen Holm Oak Quercus ilex as dominant tree species. First, we examine the spatial variation across habitats and sites in the stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) along the oak leaf–Tortrix moth Tortrix viridana caterpillar–Blue Tit Cyanistes caeruleus food chain. Secondly, we assess whether the isotopic signatures allow for correct assignment of individual birds to their site or habitat of origin. At the scale of the landscape, stable isotope values enabled identification of the different components of the Blue Tit food chain: from oak leaves to Blue Tit nestlings and yearling birds. However, isotopic signatures were site‐specific (i.e. geographical) more than habitat‐specific (i.e. deciduous vs. evergreen oaks). Discriminant analyses correctly assigned 85% of nestlings and 83% of resident yearling birds, indicating a pronounced effect of site on Blue Tit feather isotopic signatures. We thereby demonstrate that isotopes reflect a stronger association of locally born birds to the local features of their habitat than that of un‐ringed yearling birds, whose plumage may have grown while in a wider geographical area. This study provides evidence of site‐specific isotopic signatures from oak leaves to Blue Tit feathers at a fine spatial scale.     

A Quantitative Analysis of Flight Feather Replacement in the Moustached Tree Swift Hemiprocne mystacea,a Tropical Aerial Forager

The functional life span of feathers is always much less than the potential life span of birds, so feathers must be renewed regularly. But feather renewal entails important energetic, time and performance costs that must be integrated into the annual cycle. Across species the time required to replace flight feather increases disproportionately with body size, resulting in complex, multiple waves of feather replacement in the primaries of many large birds. We describe the rules of flight feather replacement for Hemiprocne mystacea, a small, 60g tree swift from the New Guinea region. This species breeds and molts in all months of the year, and flight feather molt occurs during breeding in some individuals. H. mystacea is one to be the smallest species for which stepwise replacement of the primaries and secondaries has been documented; yet, primary replacement is extremely slow in this aerial forager, requiring more than 300 days if molt is not interrupted. We used growth bands to show that primaries grow at an average rate of 2.86 mm/d. The 10 primaries are a single molt series, while the 11 secondaries and five rectrices are each broken into two molt series. In large birds stepwise replacement of the primaries serves to increase the rate of primary replacement while minimizing gaps in the wing. But stepwise replacement of the wing quills in H. mystacea proceeds so slowly that it may be a consequence of the ontogeny of stepwise molting, rather than an adaptation, because the average number of growing primaries is probably lower than 1.14 feathers per wing.     

Inferring the ecology of willow warblers during their winter moult by sequential stable isotope analyses of remiges

We present a comparison of feather stable isotope (δ¹³C, δ¹⁵N) patterns representing the habitat and diet conditions for two subspecies of willow warblers Phylloscopus trochilus that breed in parapatry, but winter in different regions of sub‐Saharan Africa. Previous analyses have shown that on average winter moulted innermost primaries (P1) show subspecific differences in δ¹⁵N values, although individuals show substantial variation for both δ¹³C and δ¹⁵N within the subspecies. We examined whether corresponding variation in the timing of the winter moult, as reflected by consistent intra‐wing correlations for individual's δ¹³C and δ¹⁵N values, could explain some of the previously observed isotopic variation. Further, differential subspecific adaptations to winter precipitation patterns across Africa might result in a variable degree of site fidelity or itinerancy during moult. We found no consistent trend in isotopic values from innermost to outermost primaries, thus inter‐individual variation in the timing of moult does not explain the subspecific isotopic variation for P1. Patterns in wing feather δ¹³C and δ¹⁵N values indicated that 41% of the individuals from both subspecies shifted their diet or habitats during winter moult. Importantly, despite well‐documented itinerancy in willow warblers during the winter, 59% of the individuals had feather isotope values consistent with stable use of habitats or diets during winter moult. Repeatability analyses suggest that individuals of both subspecies initiate moult in similar habitats from year‐to‐year while feeding on isotopically similar diets.     

Individual migratory patterns of two threatened seabirds revealed using stable isotope and geolocation analyses

Aim

Intraspecific variability in the migratory movements of seabirds is being revealed far more complex than hitherto recognized, and our lack of understanding undermines their effective protection. Our aim is to test whether the isotopic values of a single feather of two threatened seabirds, the Mediterranean (Puffinus yelkouan) and the Balearic (Puffinus mauretanicus) shearwaters allow the geographic assignment of birds to their non‐breeding areas.

Location

These two species are known to use three main non‐breeding areas: the NE Atlantic Ocean, the W Mediterranean and the Aegean‐Black seas.

Methods

We clustered in three groups the δ¹³C and δ¹⁵N values of the first primary feather (P1), inferred to be grown during the non‐breeding period, of 34 Mediterranean and 56 Balearic shearwaters accidentally caught by longliners. To link the isotopic values of P1 to its corresponding non‐breeding area, we performed a discriminant function analysis (DFA) based on the three clusters and applied this function to feathers of known origin: P1 from seven Mediterranean shearwaters from Hyères Archipelago (France) tracked with geolocators and body feathers from six chicks from Balearic shearwaters. To link the moulting patterns to the areas where the feathers were grown, we applied the DFA to a sequence of primary feathers of eight Balearic and five Mediterranean shearwaters (caught by longliners).

Results

Isotopic and tracking data indicate that none of the Mediterranean shearwaters migrated to the Atlantic. The cluster and discriminant function analyses revealed that 8% of Balearic and 54% of Mediterranean shearwater moulted P1 in the Mediterranean Sea. Migratory movements were reflected in the changing isotopic values of the primary sequences.

Main conclusions

Stable isotope analyses (SIA) are a powerful approach to reveal the intraspecific variability in the migratory patterns of seabirds that use distinct isotopic areas over their annual cycle. The assignment of birds to their non‐breeding areas by means of SIA is a simple and effective tool that can help to evaluate the impact of human activities in remote areas not only at population but also at individual level, which is an essential knowledge for the management and conservation of threatened species.

     

Using hydrogen isotopes of freshwater fish tissue as a tracer of provenance

Hydrogen isotope (δ²H) measurements of consumer tissues in aquatic food webs are useful tracers of diet and provenance and may be combined with δ¹³C and δ¹⁵N analyses to evaluate complex trophic relationships in aquatic systems. However, δ²H measurements of organic tissues are complicated by analytical issues (e.g., H exchangeability, lack of matrix‐equivalent calibration standards, and lipid effects) and physiological mechanisms, such as H isotopic exchange with ambient water during protein synthesis and the influence of metabolic water. In this study, δ²H (and δ¹⁵N) values were obtained from fish muscle samples from Lake Winnipeg, Canada, 2007–2010, and were assessed for the effects of species, feeding habits, and ambient water δ²H values. After lipid removal, we used comparative equilibration to calibrate muscle δ²H values to nonexchangeable δ²H equivalents and controlled for H isotopic exchange between sample and laboratory ambient water vapor. We then examined the data for evidence of trophic δ²H enrichment by comparing δ²H values with δ¹⁵N values. Our results showed a significant logarithmic correlation between fork length and δ²H values, and no strong relationships between δ¹⁵N and δ²H. This suggests the so‐called apparent trophic compounding effect and the influence of metabolic water into tissue H were the potential mechanisms for δ²H enrichment. We evaluated the importance of water in controlling δ²H values of fish tissues and, consequently, the potential of H isotopes as a tracer of provenance by taking account of confounding variables such as body size and trophic effects. The δ²H values of fish appear to be a good tracer for tracking provenance, and we present a protocol for the use of H isotopes in aquatic ecosystems, which should be applicable to a broad range of marine and freshwater fish species. We advise assessing size effects or working with fish of relatively similar mass when inferring fish movements using δ²H measurements.     

Wintering areas predict age‐related breeding phenology in a migratory passerine bird

Understanding connections between breeding, stopover and wintering grounds for long‐distance migratory birds can provide important insight into factors influencing demography and the strength of carry‐over effects among various periods of the annual cycle. Using previously described, multi‐isotope (δ¹³C, δ¹⁵N, δ²H) feather isoscapes for Africa, we identified the most probable wintering areas for house martins Delichon urbica breeding at Badajoz in southwestern Spain. We identified two most‐probable wintering areas differing in isotopic signature in west Africa. We found that the probability to winter in the isotopic cluster two was related to age and sex of individuals. Specifically, experienced males (i.e. two years or older) winter in the isotopic cluster two with a greater probability than experienced females, whereas first‐year females winter in the isotopic cluster two with a greater probability than first‐year males. In addition, wintering area was correlated with breeding phenology, with individuals wintering in the isotopic cluster two initiating their clutches earlier than those wintering in the isotopic cluster one. For birds wintering in the isotopic cluster two, there was no relationship between age and clutch initiation date. In contrast, young birds wintering in the isotopic cluster one initiated their clutches earlier than experienced birds wintering in this area. There was no significant correlation between wintering area and clutch size or the number of fledglings produced. We hypothesize that the relationship among social status, population density and winter habitat quality should be the most important driver of the carry‐over effect we found for this population.     

Large‐scale geographic variation in iridescent structural ornaments of a long‐distance migratory bird

Iridescent colours produced during moult likely play an important role in pair formation in birds. We sought to quantify geographic variation in such colouration in a duck species, Eurasian teal Anas crecca, in winter (when mating occurs) to evaluate whether this variation reflects birds’ breeding origins or differential individual migration strategies in both males and females. We combined information on feather production region and individual attributes (body size, sex and age) of Eurasian teal from 82 wintering sites in France. Feather production region (moult site or natal origin) was inferred using feather deuterium values (δD_f). We performed spectral measurements to evaluate speculum colour and brightness contrasts for 1052 teal collected over four years. Colouration differed strongly among wintering regions, with birds wintering in eastern France exhibiting higher colour contrast than those wintering in the west. Body size and colouration were positively related. There were no differences in cohort‐specific δD_f values between separate wintering regions in France, indicating that within a winter quarter teal originated from areas across the entire breeding range. Overall, patterns of spatial variation in feather colouration were related most closely to body size which was consistent with predictions of a differential migration hypothesis, with larger and more colour‐contrasting birds wintering closer to their breeding grounds. Because moult speed is also known to affect colour production, early breeders or individuals that skipped reproduction may have invested more or earlier in their feather quality to gain potential advantages in monopolizing future mates.