Moult of overwintering Wood Warblers <i>Phylloscopus sibilatrix</i> in an annual-cycle perspective

Wood Warblers, an Afro-Palearctic migrant species, are declining steadily in Europe likely due to mortality outside their breeding grounds. However, little is known about their overwintering, and records about the sensitive life-cycle stage of moult in Africa are practically absent. To fill this gap, we report on moult of Wood Warblers captured over two winters (January–February) in 2019–2020 in Cameroon. We caught 14 individuals, of which 12 were monitored for flight feather moult. All inspected individuals showed advanced stages of flight feather renewal. Despite low sample sizes, Underhill-Zucchini moult models aptly explained variation in primary and secondary moult (R2 = 0.61). Estimated moult onset date was 26 December, completion date was 25 February, and moult duration was 61 days. These findings fit well with experimental data on the annual cycle and the timing of recently published migration tracks of Wood Warblers. Jointly, the data suggest that moult timing is set by an internal programme, which enables Wood Warblers to organise their multi-stage migration such that they reach suitable moulting habitat in time, and can depart in time with a fresh plumage for the breeding grounds. In our study, moult occurred during the peak of the dry season, which in Cameroon nonetheless shows high relative humidity. During our mist-netting on 28 cocoa plantations of varying shade cover, Wood Warblers were caught on 6 farms whose canopies were comparatively open. These data suggest that the birds encounter in Cameroon relatively stable climatic conditions for moult, and do not measurably prefer closed-canopy forests. Our findings are important, because successful moult increases survival prospects and because moult needs to be safely embedded in a migratory life cycle. Hence, information on moult timing and location is essential for identifying year-round vulnerabilities of Wood Warblers.<br/

Stable isotope analysis provides new information on winter habitat use of declining avian migrants that is relevant to their conservation

Winter habitat use and the magnitude of migratory connectivity are important parameters when assessing drivers of the marked declines in avian migrants. Such information is unavailable for most species. We use a stable isotope approach to assess these factors for three declining African-Eurasian migrants whose winter ecology is poorly known: wood warbler Phylloscopus sibilatrix, house martin Delichon urbicum and common swift Apus apus. Spatially segregated breeding wood warbler populations (sampled across a 800 km transect), house martins and common swifts (sampled across a 3,500 km transect) exhibited statistically identical intra-specific carbon and nitrogen isotope ratios in winter grown feathers. Such patterns are compatible with a high degree of migratory connectivity, but could arise if species use isotopically similar resources at different locations. Wood warbler carbon isotope ratios are more depleted than typical for African-Eurasian migrants and are compatible with use of moist lowland forest. The very limited variance in these ratios indicates specialisation on isotopically restricted resources, which may drive the similarity in wood warbler populations' stable isotope ratios and increase susceptibility to environmental change within its wintering grounds. House martins were previously considered to primarily use moist montane forest during the winter, but this seems unlikely given the enriched nature of their carbon isotope ratios. House martins use a narrower isotopic range of resources than the common swift, indicative of increased specialisation or a relatively limited wintering range; both factors could increase house martins' vulnerability to environmental change. The marked variance in isotope ratios within each common swift population contributes to the lack of population specific signatures and indicates that the species is less vulnerable to environmental change in sub-Saharan Africa than our other focal species. Our findings demonstrate how stable isotope research can contribute to understanding avian migrants' winter ecology and conservation status

Habitat characteristics of wintering Wood Warbler Phylloscopus sibilatrix in the Centre Region of Cameroon: conservation implications

Populations of many Afro-Palearctic birds have declined, with those wintering in sub-Saharan Africa, such as Wood Warbler Phylloscopus sibilatrix, particularly affected. In this study we investigated the relationship between habitat characteristics and Wood Warbler presence/absence in the Centre Region of Cameroon. A total of six transects were established in three habitat types (forest, forest&ndash;savanna transitional zone and savanna). Call playback surveys were conducted monthly from November 2015 to April 2016 to determine Wood Warbler presence/absence. Detailed habitat measurements were also recorded in each transect. A total of 86 responses were recorded: 33 (mean 6.6 &plusmn; 2.3) in forest habitat, 47 (mean 9.4 &plusmn; 3.36) in the forest&ndash;savanna transitional zone, and 6 (mean 2 &plusmn; 1.1) in savanna habitat. Wood Warbler presence increased significantly with the number of trees between 3 and 7 m in height, and decreased significantly with the number of shrubs between 0.5 and 3 m in height. Anthropogenic disturbance such as the agricultural cycle and burning were not found to have an effect on Wood Warblers presence/absence. We conclude that Wood Warblers overwinter in all three habitat types with probability of detection greatest in the forest&ndash;savanna transitional habitat with a relatively low canopy and an open understorey. Forest clearance in sub-Saharan Africa potentially threatens wintering habitat for Wood Warblers

An East-West Comparison of Migration in North American Wood Warblers

That western and eastern songbird migration routes are distinct ecological systems has been proposed for over 100 years. Nonetheless, this distinction has not been widely recognized nor have there been any comparative studies that quantitatively evaluate the differences and similarities between western and eastern songbird migration systems. We drew from previously published research on wood warblers to highlight patterns in stopover ecology that suggest fundamental differences between western and eastern migrants. In particular, we compared biogeography, evolutionary relationships, and stopover ecology of wood warblers from western and eastern North America and found: (1) multiple lines of evidence that indicate western wood warblers are geographically isolated from eastern con-specifics or congeners throughout the annual cycle, (2) eastern and western wood warbler taxa are distinct evolutionary units, (3) migrant wood warblers captured in the Southwest tended to carry lower fat loads and be comprised of more after-hatch-year birds than is typical of eastern migrants, (4) frugivory is unknown in wood warblers endemic to the Northwest or Southwest and (5) relative to other regions and seasons, riparian vegetation is heavily used by western wood warblers in the spring. We think that further examination and synthesis of these differences would yield a more mechanistic understanding of Nearctic-Neotropical avian migration. On this basis, we elaborate our view that (1) an improved understanding of western songbird migration ought to be a high priority for science, conservation, and education, and (2) large-scale coordinated research efforts would be the most effective strategy for advancing our knowledge of passerine migration in the West. /// Por más de 100 años se ha propuesto que las rutas de migración de aves paseriformes del este y del oeste son sistemas ecológicos diferentes. Sin embargo, esta distinción no ha sido reconocida ampliamente, ni se han realizado estudios comparativos que evalúen cuantitativamente las diferencias y similitudes entre los sistemas de migración del este y del oeste. Revisamos estudios previamente publicados sobre aves de bosque de la familia Parulidae (Reinitas) para destacar los patrones ecológicos de las paradas migratorias que sugieran diferencias fundamentals entre las aves paseriformes migratorias el este y del oeste. Particularmente, comparamos la biogeografía, las relaciones evolutivas y la ecología de las paradas migratorias de las Reinitas del este y del oeste de Norteamérica y encontramos: (1) múltiples líneas de evidencia que indican que las Reinitas del oeste se encuentran geográficamente aisladas de sus coespecíficos o cogenéricos del este durante todo el ciclo anual, (2) los taxa del este y del oeste son unidades evolutivas independientes, (3) las Reinitas migratorias capturadas en el suroeste tendieron a presentar menores cargas de grasa y una mayor proporción de aves eclosionadas durante el año, que lo es típicamente observado para las aves migratorias del este, (4) la frugivoría no se conoce en las Reinitas endémicas del noroeste o sudoeste y (5) en relación a otras regiones o estaciones, la vegetación riparia es usada intensamente durante la primavera por las Reinitas del oeste. Creemos que estudios adicionales y una síntesis de estas diferencias podrían producir un mejor entendimiento mecaní stico de la migración Neártico-Neotropical de aves. Basados en esto, sugerimos que (1) un mejor entendimiento de la migración de aves paserinas del oeste debería tener alta prioridad para la ciencia, conservación y educación y (2) que los esfuerzos de investigación coordinados a gran escala serían la estrategia más efectiva para progresar en nuestro conocimiento sobre la migración de las aves paserinas en el oeste

Spatial behavior and habitat use in widely separated breeding and wintering distributions across three species of long-distance migrant <i>Phylloscopus</i> warblers

Aim: To investigate the ecological relationship between breeding and wintering in specialist and generalist long-distance migratory species, and the links between densities and range sizes. Location: Denmark, Senegal and Ghana. Methods: We use radio tracking to study spatial behavior and habitat use in three morphologically and ecologically similar and closely related Phylloscopus species on their widely separated breeding and wintering distributions. During wintering and breeding, willow warblers P. trochilus (winter: n = 9, breeding: n = 13), chiffchaffs P. collybita (n = 11, n = 7), and wood warblers P. sibilatrix (n = 17, n = 14) were tracked. Results: Willow warblers P. trochilus increased home range sizes in winter, whereas it was similar in chiffchaffs P. collybita and wood warblers P. sibilatrix, in both seasons. Home ranges overlapped more in winter than in the breeding season. In winter, home range overlap was similar among species but larger overlap during breeding was indicated for willow warblers. Tree cover was unrelated to home range size but significantly higher in breeding than in winter in all species. However, whereas willow warblers and wood warblers maintained some degree of tall tree cover inside their home ranges in winter, chiffchaffs changed from more than 80% to <1% tree cover, indicating a niche shift. Main conclusions: Individuals of all three species showed changes between breeding and wintering areas in spatial behavior and habitat availability, with larger overlap in winter. The differences in patterns were potentially related to being generalist (willow warbler) or specialist (chiffchaff and wood warbler). These ecological relationships are important for the conservation of migrants and for understanding the link between breeding and wintering distributions and ecology

Geographic range size and evolutionary age in birds

Together with patterns of speciation and extinction, post-speciation transformations in the range sizes of individual species determine the form of contemporary species-range-size distributions. However, the methodological problems associated with tracking the dynamics of a species' range size over evolutionary time have precluded direct study of such range-size transformations, although indirect evidence has led to several models being proposed describing the form that they might take. Here, we use independently derived molecular data to estimate ages of species in six monophyletic groups of birds, and examine the relationship between species age and global geographic range size. We present strong evidence that avian range sizes are not static over evolutionary time. In addition, it seems that, with the regular exception of certain taxa (for example island endemics and some threatened species), range-size transformations are non-random in birds. In general, range sizes appear to expand relatively rapidly post speciation; subsequently, and perhaps more gradually, they then decline as species age. We discuss these results with reference to the various models of range-size dynamics that have been proposed

Migration Distance and Sexual Dichromatism: Cost of Migration is Correlated with Reduced Female Coloration in Wood Warblers

Avian sexual dichromatism has been observed to be greater at higher latitudes, and one of the long-standing explanations for the underlying evolutionary forces underlying this geographic pattern is the breeding latitude hypothesis, which focuses on changes in male ornamentation and posits that sexual selection for male ornamentation is stronger at higher latitudes. I offer an alterative hypothesis that focuses on how changes in female coloration can drive the evolution of sexual dichromatism. I propose that the costs of migration, including increased predation risk and high energetic costs, negatively select against elaborate plumage coloration, and that the distance a species migrates is positively related to these costs. Furthermore, because positive directional selection for ornamentation is generally greater on males than on females, I propose that the costs associated with longer migration have imposed stronger net negative selection on female ornamentation compared to male ornamentation. Specifically, I predict that migration distance is positively related to sexual dichromatism. To address both the breeding latitude and migration distance hypotheses, I tested how migration distance and breeding latitude predict sexual differences in plumage coloration among wood warblers (Parulidae) using independent contrasts to control for phylogeny. Both migration distance and breeding latitude independently predict sexual dichromatism, and based on model-selection analyses, my results indicate that migration distance has more predictive power than breeding latitude. Thus, these results provide evidence consistent with the mechanism that changes in female ornamentation have driven the evolution of sexual dichromatism. Furthermore, these results suggest that incremental increases in migration distances are associated with concurrent increases in the costs associated with ornamentation. Future studies of both migratory and non-migratory taxa should consider how costs associated with female ornamentation might contribute to the evolution of sexual dimorphism

Predator recognition and differential behavioural responses of adult Wood Warblers Phylloscopus sibilatrix

This is the final version of the article. Available from Springer Verlag via the DOI in this record.Birds often engage in nest defence against predators to improve breeding success, but defence efficiency requires the capability to assess the threat level posed by potential predators. For species with low breeding-site tenacity, which may encounter varying occurrence and density of predators in different areas, threat recognition could be compromised due to naivety, and so predator recognition may focus on broad key features to diminish the risk of misidentification. We experimentally tested this hypothesis by recording behavioural reactions of the nomadic wood warbler Phylloscopus sibilatrix to objects reflecting various levels of threat: least weasel and Eurasian jay taxidermy mounts, an inanimate object and an empty display mount. To assess actual nest predators, we used remote cameras to record predation events at wood warbler nests. As in other studies in Western Europe, Eurasian jay was found to be the main nest predator, with occasional predation by least weasel. The reaction of adult warblers to the models was generally to remain silent and on nests during the incubation stage presumably due to the need to maintain efficient nest camouflage and concealment. During the nestling stage, behavioural responses of adult warblers, calling and suspended feeding of young, showed the strongest effects from the jay taxidermy mount, moderate to the weasel and weakest to the inanimate object and empty mount. As the reaction of wood warblers reflected the degree of genuine threat posed by the predators depicted by the models, we conclude that predator recognition may be present in this species.We thank John Mallord, Chris Orsman and Andrew Cristinacce for monitoring nests in the wider study area as part of a project jointly funded by RSPB and Natural England within the Action for Birds in England partnership, Louise and Gary Clewey for assistance in the field and landowners Natural England, Woodland Trust, National Trust and Devon Wildlife Trust

Testosterone increase in free-living young Blackcaps <i>Sylvia atricapilla</i> and Wood Warblers <i>Phylloscopus sibilatrix</i> during post-juvenile molt with possible implications for juvenile dispersal

Testosterone may affect many aspects of passerine maturation including the expression of plumage coloration and spring migration in adult birds. However, how the testosterone level changes during bird development is not well known. We compared the testosterone profile during juvenile development of a sexually dimorphic species, the Blackcap Sylvia atricapilla, and a monomorphic species, the Wood Warbler Phylloscopus sibilatrix. We first tested whether testosterone influences plumage coloration and increases at the onset of molt in males of the dimorphic but not the monomorphic species. We found that a testosterone level increase occurred in both sexes and species during later stages of molt, and thus was not related to male plumage coloration. We also investigated whether the increase in testosterone level coincides with juvenile dispersal. If testosterone affects dispersal behavior in these species, both sexes should show an elevated testosterone level during dispersal, but this increase should occur earlier in the Wood Warbler, which disperses earlier than the Blackcap. In juvenile Blackcaps, the increase in testosterone level occurred on the 43-68th day after hatching, while in Wood Warblers it occurred on the 32-36th day (i.e. 11-32 days earlier). The increase in testosterone level coincided with the onset of the post-juvenile dispersal in both species. This study provides the first direct evidence of a testosterone level increase during juvenile development in two free-living migratory birds. While not correlated with species plumage coloration, this increase might affect other aspects of behavior, e.g. juvenile dispersal

The influence of wind direction on the capture of the wood warbler (Phylloscopus sibilatrix), an uncommon migratory species in the western Mediterranean

The wood warbler (Phylloscopus sibilatrix) is a migratory species in the western Mediterranean wintering in the Gulf of Guinea region, West Africa. In autumn and spring, this species, along with the populations breeding in Ireland and Britain, uses the Italian peninsula as its main axis of migration. From the data of captured birds at several ringing stations in the western Mediterranean (Balearic Islands and coastal Iberian Peninsula), we analyzed capture rates of the species during spring migration from 1993 to 2007. Based on the selection of days with a significant number of captures and those without captures, we analyzed the effect of wind direction over the western Mediterranean to determine a relationship between winds and the number of captures. From a total of 663 wood warblers captured between 1993 and 2007, a total of 31 days have been selected as significant days with a high number of captures, and 31 days have been selected as no-capture days. On days of maximum captures, winds coming from an easterly direction, i.e. Algeria and Tunisia, were dominant, indicating days with a clear eastern component. Contrary to expected results, captures were also made on days when the wind direction was predominantly from a northerly direction. Analysis of the origin of the winds in north eastern Spain (western Mediterranean) revealed that the majority of northerly winds originated from Africa and not from Europe as is usual for this region. Days or periods selected as no-capture days were characterized by winds coming from a northerly (European origin) or westerly direction