Philopatry of winter moult area in migratory Great Reed Warblers Acrocephalus arundinaceus demonstrated by stable isotope profiles

Stable carbon- (δ¹³C), nitrogen- (δ¹⁵N) and hydrogen (δD) isotope profiles in feathers of migratory Great Reed Warblers Acrocephalus arundinaceus recaptured for 2 or more years in 6 successive years were examined to test whether the isotope profiles of individual warblers appeared to be consistent between years. Similar isotopic signatures in successive years suggested that individual birds tended to return and grow their feathers in Afro-tropical wintering habitats that generate similar δ¹³C, δ¹⁵N and δD signatures. Previous studies have shown that Great Reed Warblers exhibit strong natal and breeding philopatry, with most of the surviving birds returning to the breeding site. The present study of feather δ¹³C, δ¹⁵N and δD isotopic values demonstrate the year-to-year fidelity might also include the African moulting sites in this migratory species.     

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.     

Winter habitat use does not influence spring arrival dates or the reproductive success of Yellow Warblers breeding in the arctic

Winter habitat use can influence the breeding success of migratory songbirds in temperate regions due to its impact on bird condition and breeding phenology. How such carry-over effects vary with latitude is unknown. To address this question, we examined how winter habitat use, inferred from δ¹³C and δ¹⁵N signatures in winter-grown feathers, influenced the breeding phenology and productivity of Yellow Warblers (Setophaga petechia) at the extreme north of their range in the Canadian arctic (68°N) and compared this population with midlatitude Yellow Warbler (51°N) and American Redstart (Setophaga ruticilla; 44°N) populations reported in previous studies. In the arctic, we examined male arrival dates, female clutch initiation dates and the relationship between these timing variables and the number and quality of offspring produced within the season. In contrast to warblers breeding at midlatitudes, we find no support for an impact of winter habitat use on breeding phenology or productivity. Male arrival dates and female clutch initiation dates in both young and older individuals were not correlated with isotopic signatures acquired on the wintering grounds. Males with enriched δ¹⁵N signatures paired more rapidly after arrival, indicating a possible relationship between winter habitat use and condition. This relationship did not enhance annual productivity for these individuals, however, as the negative relationship between breeding phenology and reproductive success in our arctic population was significantly weaker than among Yellow Warblers breeding further south. This reduction or absence of timing effects on productivity in the north effectively removes one pathway through which carry-over effects can act.     

Identifying the winter grounds of the recently described Barbary Reed Warbler (Acrocephalus baeticatus ambiguus)

The Iberian and North African populations of reed warblers have been described recently as a separate taxon, ambiguus, forming a sister clade to the Sahelian subspecies minor of African Reed Warbler Acrocephalus baeticatus. Although the breeding range of ambiguus has been identified, the migration strategy of its populations remained unknown. We deployed geolocators and sampled the innermost primary from breeding adults in Spain for stable hydrogen (δ²H) analyses and also analysed stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopes in feathers collected in two reed warbler taxa (Acrocephalus scirpaceus and Acrocephalus baeticatus ambiguus) in Morocco, to identify the moulting and wintering sites of these populations. Ring recoveries, geolocator tracks and probabilistic assignments to origin from δ²H values indicate that Spanish ambiguus are likely to moult south of the Sahara and winter in West Africa, probably from Mauretania to southern Mali and Ivory Coast. Moroccan ambiguus, however, undergo post-breeding moult north of the Sahara, and possibly then migrate to West Africa. With other populations of ambiguus breeding from Algeria to Libya and probably wintering further east in the Sahelian belt, the Barbary Reed Warbler can therefore be considered a trans-Saharan migrant, with a post-breeding moult strategy that varies between populations, and probably structured according to breeding latitude.     

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.     

Discriminating geographic origins of migratory waders at stopover sites: insights from stable isotope analysis of toenails

In this study we test the potential of stable isotope analysis to reveal wintering origins of waders mixing at stopover sites, using the dunlin Calidris alpina as a case study. We determined stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope signatures of toenails of dunlins captured during winter at reference sites along the East‐Atlantic Flyway, from Mauritania to the United Kingdom. Afterwards, during spring migration, dunlins were sampled at the Tagus estuary, Portugal, and assigned to their wintering grounds according to their stable isotope signatures. Toenails from wintering dunlins at different sites had significantly different δ¹³C and δ¹⁵N signatures, despite some overlap in isotopic carbon ratios of birds from Morocco, Portugal and the UK. Among birds sampled during migration in Portugal, we found a clear bimodal pattern in δ¹³C values, corresponding to passage migrants from Mauritania (enriched δ¹³C values) and wintering birds from the Tagus estuary (depleted δ¹³C values). The first passage migrants from Mauritania appeared at the Tagus estuary by the end of March, with peak numbers during late April and early May. Our study provides evidence that isotopic signatures of toenails can play a determinant role in tracing the wintering origins of migrant dunlins at their stopover areas. Toenails, instead of feathers, can be the powerful and innovating tissue to sample in wader studies, allowing to bridge the gap in the field of migratory connectivity between sites used in different phases of the life cycle of waders.     

Breeding phenology,reproductive traits and apparent survival of sympatric Marsh Warbler Acrocephalus palustris and Blyth’s Reed Warbler Acrocephalus dumetorum

Capsule: Sympatric Marsh Warblers Acrocephalus palustris and Blyth’s Reed Warblers Acrocephalus dumetorum differ significantly in their life history traits.

Aims: To provide a direct comparison of demographic parameters among two sympatric populations of the closely related Marsh Warbler and Blyth’s Reed Warbler.

Methods: We examined breeding phenology and reproductive traits at a 25?ha study plot. We use program MARK to estimate daily nest survival and adult apparent survival rates.

Results: On average, Marsh Warblers laid the first eggs 3 days later than Blyth’s Reed Warblers. Mean clutch size in the Marsh Warbler was significantly lower than in the Blyth’s Reed Warbler. There are no significant differences between the two species for nest daily survival, duration of incubation and nestling periods. Apparent survival of adults was slightly higher in Marsh Warblers than in Blyth’s Reed Warblers.

Conclusion: Our findings suggest that two ecologically similar sympatric species differ significantly in terms of life history traits. We assume that observed differences could be the result of adaptations to environmental factors in the central parts of the species’ ranges or due to differences in mortality on migratory pathways or wintering grounds.     

The use and function of snake skins in the nests of Great Reed Warblers Acrocephalus arundinaceus

We documented and experimentally tested the use of snake skins in construction of nests in a Great Reed Warbler Acrocephalus arundinaceus population in southwestern Slovakia. There was no difference in predation rates of artificial nests that did or did not contain sloughs. We suggest that snake skins in Great Reed Warbler nests may serve as a post‐pairing signal revealing female parental quality.     

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.     

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.     

Changes in migration phenology and biometrical traits of Reed,Marsh and Sedge Warblers

Global environmental processes like climate change could severely affect population level migratory behaviour of long range migrant birds. We analyzed changes in migration phenology and biometrics of three closely-related long-distance migrant Acrocephalus species. We used the records of 12 063 Sedge, 12 913 Reed, and 5 409 Marsh Warblers caught and ringed between 1989–2009, at a Hungarian stopover site. Quantile regressions were used to analyse the changes in spring and autumn migration phenology. Median spring arrival date of Sedge and Reed Warblers shifted 6.5 and 7.5 days earlier, respectively. Autumn arrival of all species shifted one (Reed and Marsh Warblers) or two (Sedge Warbler) weeks later. Mean body mass of adult Reed and Marsh Warblers decreased in spring (by 0.3 and 0.2 grams, respectively) and in autumn (by 0.8 and 0.2 grams, respectively) while body mass of adult Sedge Warblers decreased only in autumn (by 0.4 grams). Mean wing length of all species increased significantly (range of change: 0.6–1 mm). Despite the fact that the studied species are closely related, all three have remarkably different migration strategies. However, similar patterns can be observed in the studied parameters, indicating that global processes may have general effects on these species, albeit through markedly different mechanisms.     

Using stable isotopes to link breeding population trends to winter ecology in Willow Warblers,Phylloscopus trochilus

Capsule Stable isotope composition of feathers grown in Africa indicates that Willow Warblers, Phylloscopus trochilus, breeding in areas of Britain experiencing different population trends may vary in their distribution, timing of moult and/or use of prey resources during the non-breeding season.

Aims To compare stable isotope ratios of feathers of Willow Warblers breeding in parts of Britain with differing rates of population change.

Methods Feathers were collected from three regions with differing recent population trends; northern Scotland (population increased), western Wales (declined slightly) and eastern England (declined rapidly). Collection occurred at ringing sites run by British Trust for Ornithology (BTO) volunteers as part of the Constant Effort Sites (CES) scheme between April and July in 2008 and 2009.

Results Willow Warblers from northern Scotland had significantly higher δ ¹⁵N and δ ¹³C values than those from eastern England, although the regional mean values did not differ greatly. There was no significant difference in δ ¹⁵N and δ ¹³C values between the sexes, but δ ¹⁵N values were significantly greater in individuals caught later in the breeding season.

Conclusions During the premigratory moult period in Africa, Willow Warblers from different parts of Britain may vary in their distribution, timing of moult and/or use of prey resources.     

The identification and distribution of the enigmatic Large‐billed Reed Warbler Acrocephalus orinus

We re‐evaluate the characteristics of the little‐known Large‐billed Reed Warbler Acrocephalus orinus based on plumage and biometrical comparisons of 10 newly discovered specimens of A. orinus with a large sample of Blyth’s Reed Warbler Acrocephalus dumetorum, the species most closely resembling A. orinus. Using specimens whose identity was verified by DNA analyses, we found that the most important criteria for identifying orinus are its long and comparatively broad bill, with typically uniformly dark upper mandible, and on average longer tarsi and claws with a thin and pointed hind claw. In contrast, previously suggested differences in wing formula, plumage colour and shape of flight‐feathers are very minor, variable and seem of limited use for identification of orinus. Graphical analysis of hind claw length against bill to skull length should assist researchers in locating museum or live specimens of orinus that otherwise might be overlooked as dumetorum, and multivariate analysis unambiguously separates the two species based on our present dataset. A review of the scattered literature and specimen records suggests that the breeding distribution of orinus lies between southeastern Kazakhstan and northeastern Afghanistan, within the breeding range of dumetorum, but that orinus may well winter predominantly east of the main wintering range of dumetorum. Further evidence to support this assessment of the species' breeding range was obtained in June 2009, with the discovery of a population in the Pamir Mountains of Afghanistan.     

Erratum: Isotopic (δ2Hf) evidence of “loop migration” and use of the Gulf of Maine Flyway by both western and eastern breeding populations of Blackpoll Warblers

Our earlier report of breeding/natal ground assignment, through the use of feather deuterium samples (δ²H_f) of Blackpoll Warblers (Setophaga striata) during spring and fall migration at various sites in North America, contained an erroneous breeding origin map derived for a group of birds sampled during spring migration in Florida. Updated analyses revealed a broader breeding range assignment than previously reported, but this change does not affect our original conclusions that Blackpoll Warblers exhibit a “loop migration” pattern as they move between breeding and wintering areas.     

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.     

Experimental enlargement of nest size does not increase risk of predation or brood parasitism in the Great Reed Warbler Acrocephalus arundinaceus

We assessed whether nest size affects the probability of nest loss using dyads of large and small (large being twice the size of small) inactive Great Reed Warbler Acrocephalus arundinaceus nests placed at similar sites in Great Reed Warbler territories. Large nests were not predated significantly more frequently than small nests. Experimentally enlarged active Great Reed Warbler nests suffered non‐significantly higher predation compared with non‐manipulated control nests. Our experiments did not support the nest‐size hypothesis and suggested that nest size does not appear to be a factor affecting the risk of nest predation in this species. The probability of brood parasitism by the Common Cuckoo Cuculus canorus was also unaffected by experimental nest enlargement, supporting the commonly accepted hypothesis that the Common Cuckoo searches for suitable host nests by host activity during nest building rather than nest size.     

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.     

Resource use for reproduction depends on spring arrival time and wintering area in an arctic breeding shorebird

Resources for egg production may come from body reserves stored before breeding (“capital breeders”) or from food acquired at the breeding site (“income breeders”). Arctic migrants were long thought to be capital breeders, because they often arrive at a time when local food availability is still limited. However, later evidence suggested that arctic breeding shorebirds are primarily income breeders, or that they use a mixed strategy depending on laying date. We explored the relationship between laying date and resource use for reproduction in the pectoral sandpiper Calidris melanotos breeding in the Alaskan arctic by contrasting carbon isotope (δ¹³C) values of the local diet and of maternal plasma, cellular blood, feather and claw with those of the eggs produced. Our results revealed that early breeding females utilize resources for egg production that were acquired recently at staging areas, whereas later breeding females mostly relied on nutrients derived from local food sources. These findings suggest that the resource allocation strategy used for reproduction differs among females, and varies depending on the timing of arrival and the start of reproduction. The arrival date at the breeding ground and laying date may critically depend on non‐breeding season events such as winter habitat choice, staging areas or migration routes. By comparing maternal feather δ¹³C, claw δ¹³C and feather δD, we examined whether non‐breeding season events influenced the use of resources for egg production through variation in capture date or clutch initiation date. Female pectoral sandpipers originating from moulting areas characterized by higher (more positive) δD signatures were caught earlier and started laying earlier, and they used stored resources for reproduction. Using regional maps of δD values for precipitation in the wintering sites in South America, we compared the spatial variation in the observed feather δD signatures. This analysis indicated that female pectoral sandpipers with higher δD signatures, presumably coming from more north‐easterly wintering sites in southern America, started laying earlier and used mostly stored resources for egg production, compared to females that wintered (or at least moulted) further south. Our results thus show that winter moulting habitat is linked to breeding resource allocation strategy in this high‐arctic breeding shorebird.     

The Large-billed Reed Warbler Acrocephalus orinus revisited

The Large-billed Reed Warbler Acrocephalus orinus is known only from the type specimen, collected in Himachal Pradesh, India, in 1867. The specimen is poorly prepared, and it has been suggested that it could represent an isolated form of the Clamorous Reed Warbler A. stentoreus or an aberrant Blyth's Reed Warbler A. dumetorum . We tested the affinity of A. orinus by (1) re-examining the morphology of the type specimen and (2) amplifying and sequencing a portion of its mitochondrial cytochrome b gene. Both the morphological and the mitochondrial analyses showed the specimen to be similar to dumetorum , but distinct enough to qualify as a species of its own. Relative to dumetorum , it has a more rounded wing, longer bill, longer and more graduated tail with more pointed tail feathers, and larger claws. The divergence in mitochondrial DNA between orinus and dumetorum was 7.8%, well above the value expected between subspecies. A. orinus is smaller than any of the forms of A. stentoreus or the related Australian Reed Warbler A. australis . It has a somewhat longer first primary, more pointed tail feathers and paler, less robust feet and claws. DNA comparison places it in the clade of small unstreaked Acrocephalus warblers, and apart from the clade of large unstreaked warblers that contains stentoreus and australis .     

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.