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

Declining numbers of Blackpoll Warblers (Setophaga striata) have been documented at long‐term migration monitoring sites as well as in breeding areas. However, the “loop migration” of Blackpoll Warblers makes it difficult to ascribe population change at migration monitoring sites to specific breeding populations. Individuals from all populations across the breeding range of Blackpoll Warblers concentrate in fall along the Atlantic coastline of eastern North America prior to initiating a transoceanic flight to wintering areas. In spring, Blackpoll Warblers return along a different route, moving north into the southeastern United States where birds from eastern and western breeding populations then diverge during migration to reach their respective breeding areas. To monitor breeding populations outside of breeding areas and identify factors potentially affecting those populations, we must be able to identify where birds captured during migration breed and map seasonal variation in population‐specific flyways. To “map” population‐specific migration movements of Blackpoll Warblers, we used feather deuterium (δ²H_f) values and a spatially explicit model to assign molt origins of 289 Blackpoll Warblers moving through sites in the Gulf of Maine (GOM) region and at three locations further west and south (northern Great Lakes area, Pennsylvania, and Florida). The assignment method was validated with feather samples from 35 birds captured during the breeding season at Churchill, Manitoba, Canada. As predicted, the spatial pattern of movement within and between seasons reflected “loop migration.” Blackpoll Warblers captured during fall migration in the GOM region included birds from across their breeding range, whereas birds captured during the spring were exclusively from northeastern populations. During fall migration, Blackpoll Warblers captured at two sites west of the GOM were from breeding areas further northwest than those from western Canada that were captured in the GOM. Blackpoll Warblers captured in eastern Florida during spring migration were assigned exclusively to breeding areas in the northeast, suggesting that eastern and western populations diverge soon after entering the United States. Finally, most Blackpoll Warblers sampled at Manomet Bird Observatory originated from breeding populations in Alaska and western Canada that have shown a similar (70–90%) decline over the same period. Our results, therefore, not only document the “loop migration” pattern of Blackpoll Warblers, but, by mapping patterns of connectivity between breeding and non‐breeding areas, may help target conservation efforts for breeding populations of Blackpoll Warblers where most needed.     

Effects of migration distance and sex on stopover timing and refueling by Wilson's Warblers

Migration distance and sex are integral to avian migration strategies, yet these intrinsic factors are understudied with respect to their effects on stopover ecology and behavior. We investigated how individual variation in migration distance and sex affected body condition, refueling performance, and stopover durations of Wilson's Warblers (Cardellina pusilla) during spring and fall 2006 at a stopover site in southern Arizona. Migration distance of individuals was inferred using the hydrogen stable isotope ratio of feathers (δ²H_f) as an index of breeding latitude, refueling rate was assessed using plasma triglyceride and β‐OH‐butyrate, and minimum length of stay was estimated by re‐sighting color‐banded birds. In the spring, migration distance and sex were strong determinants of the timing of migration by Wilson's Warblers, with males and shorter‐distance migrants passing through the site earlier than females and longer‐distance migrants. Later‐arriving migrants also had higher fat scores and refueling rates. However, neither migration distance nor sex independently affected body condition (fat score or size‐corrected mass), refueling rate, or minimum length of stay. In a smaller sample of fall migrants, we found that longer‐distance migrants and males had higher refueling rates than shorter‐distance migrants and females. Our results show that differences in migration distance can, under some circumstances, affect how birds use stopover sites. Stable isotope analysis and other methods can provide information on migration distance, and should be integrated with measurements of timing, fuel stores, refueling performance, and departure behavior to gain a deeper understanding of bird migration.     

Fall migration and breeding origins of Canada Warblers moving through northern Colombia

Canada Warblers (Cardellina canadensis) are long‐distance Neotropical migrants, but little is known about their migratory behavior and ecology. We examined the fall migration of Canada Warblers at two sites, Darién and the Sierra Nevada de Santa Marta, in northern Colombia from 2011 to 2015 using constant‐effort mist‐netting. Our objectives were to determine: 1) breeding origins and connectivity patterns, 2) migratory pathways, 3) the phenology of migration, 4) possible differences in movements between ages and sexes, 5) their body condition when arriving in Colombia, and 6) evidence of stopover and refueling. Stable hydrogen isotopes (δ²H_f) in flight feathers were analyzed to estimate breeding origins of captured Canada Warblers in North America. The δ²H_f values revealed that most Canada Warblers captured in the Darién likely originated from the central and northeastern regions of their breeding range. The capture of all but one of 162 Canada Warblers in the Darién also indicates a migration route through Central American rather than across the Caribbean Sea. Most captured birds were hatch‐year birds (91% vs. 9% after hatch‐year birds), and we captured more females (67%) than males (33%). Canada Warblers migrated through the Darién from 20 September to early November, with most arriving in mid‐October. Most (89%) individuals arrived with low fuel reserves. These results combined with estimated flight ranges revealed that 46% of the individuals captured in the Darién likely needed to refuel to continue migrating, whereas 31% could continue 50 to 200 km beyond our capture site. However, no individuals were recaptured so stopover duration could not be determined. Canada Warblers may adopt a strategy of 1‐d stopovers and short flights or, alternatively, the Darién may represent low‐quality habitat and birds quickly left our study site in search of suitable habitat. Further study is needed to determine the possible importance of other (montane) habitats for Canada Warblers in the Darién region to prioritize conservation actions.     

Crosswise migration by Yellow Warblers,Nearctic‐Neotropical passerine migrants

Yellow Warblers (Setophaga petechia) are abundant breeding birds in North America, but their migratory and non‐breeding biology remain poorly understood. Studies where genetic and isotopic techniques were used identified parallel migration systems and longitudinal segregation among eastern‐ and western‐breeding populations of Yellow Warblers in North America, but these techniques have low spatial resolution. During the 2015 breeding season, we tagged male Yellow Warblers breeding in Maine (N = 10) and Wisconsin (N = 10) with light‐level geolocators to elucidate fine‐scale migratory connectivity within the eastern haplotype of this species and determine fall migration timing, routes, and wintering locations. We recovered seven of 20 geolocators (35%), including four in Maine and three in Wisconsin. The mean duration of fall migration was 49 d with departure from breeding areas in late August and early September and arrival in wintering areas in mid‐October. Most individuals crossed the Gulf of Mexico to Central America before completing the final eastward leg of their migration to northern South America. Yellow Warblers breeding in Maine wintered in north‐central Colombia, west of those breeding in Wisconsin that wintered in Venezuela and the border region between Brazil, Colombia, and Venezuela. Our results provide an example of crosswise migration, where the more easterly breeding population wintered farther west than the more westerly breeding population (and vice versa), a seldom‐documented phenomenon in birds. Our results confirm earlier work demonstrating that the eastern haplotype of northern Yellow Warblers winters in northern South America, and provide novel information about migratory strategies, timing, and wintering locations of birds from two different populations.     

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.     

Timing of events on the breeding grounds for five species of sympatric warblers

ABSTRACT On the breeding grounds, migratory birds have limited time to breed and molt before autumn migration. However, few studies of long‐distance migrants have examined the phenology of these events to determine what life‐history trade‐offs might result if these activities overlap. From 2000 to 2007, I used banding data to determine the timing of migration, breeding, and primary molt for Yellow Warblers (Dendroica petechia), Yellow‐rumped Warblers (D. coronata coronata), American Redstarts (Setophaga ruticilla), Ovenbirds (Seiurus aurocapilla), and Canada Warblers (Wilsonia canadensis) at a study site in Alberta, Canada. Hatching date did not differ among species (P= 0.63), with means ranging from 27 June to 3 July. All species began primary molt between 12 July and 18 July, near the expected fledging date of offspring, and therefore all species exhibited overlap between postfledging parental care and molt. The duration of primary molt ranged from 28 d for Canada Warblers to 69 d for Yellow‐rumped Warblers. Yellow Warblers, Yellow‐rumped Warblers, and American Redstarts began autumn migration having completed about 50% of their primary molt. However, Ovenbirds departed when 21% of molt was complete, and Canada Warblers departed 2 d after completing molt. For all five species of warblers, molt did not overlap with nest‐bound breeding activities. However, molt did overlap with both postfledging care and migration. This suggests that initiating migration as soon as possible is important, possibly because earlier arrival on the wintering grounds may improve access to high quality winter habitat. Overall, warblers may maximize individual fitness by combining life‐history events that result in overlapping portions of the breeding cycle, molt, and migration.     

Fetal cannabinoid receptors and the “dis‐joint‐ed” brain

Microtubule turnover in the growing axons is required for directional axonal growth and synapse formation in the developing brain. In this issue of The EMBO Journal, Tortoriello et al ( 2014 ) show that the microtubule‐binding protein SCG10/stathmin‐2 is a specific molecular target for a CB₁ receptor‐mediated effect of Δ⁹‐tetrahydrocannabinol (THC), the psychoactive ingredient of smoked marijuana, in the fetal brain. Considering the role of CB₁ in modulating the specification and long‐distance migration of neurons in the perinatal brain, this study reveals an interesting mechanism potentially accounting for connectivity deficits during cortical development following exposure to CB₁ agonists or THC during pregnancy.     

Extreme genetic similarity does not predict non‐breeding distribution of two closely related warblers

Detailed knowledge of migratory connectivity can facilitate effective conservation of Neotropical migrants by helping biologists understand where and when populations may be most limited. We studied the migratory behavior and non‐breeding distribution of two closely related species of conservation concern, the Golden‐winged Warbler (Vermivora chrysoptera) and Blue‐winged Warbler (Vermivora cyanoptera). Although both species have undergone dynamic range shifts and population changes attributed to habitat loss and social interactions promoting competition and hybridization, full life‐cycle conservation planning has been limited by a lack of information about their non‐breeding ecology. Because recent work has demonstrated that the two species are nearly identical genetically, we predicted that individuals from a single breeding population would have similar migratory timing and overwintering locations. In 2015, we placed light‐level geolocators on 25 males of both species and hybrids in an area of breeding sympatry at the Fort Drum Military Installation in Jefferson and Lewis counties, New York. Despite extreme genetic similarity, non‐breeding locations and duration of migration differed among genotypes. Golden‐winged Warblers (N = 2) overwintered > 1900 km southeast of the nearest Blue‐winged Warbler (N = 3) and spent nearly twice as many days in migration; hybrids (N = 2) had intermediate wintering distributions and migratory timing. Spring migration departure dates were staggered based on distance from the breeding area, and all birds arrived at the breeding site within 8 days of each other. Our results show that Golden‐winged Warblers and Blue‐winged Warblers in our study area retain species‐specific non‐breeding locations despite extreme genetic similarity, and suggest that non‐breeding locations and migratory timing vary along a genetic gradient. If the migratory period is limiting for these species, our results also suggest that Golden‐winged Warblers in our study population may be more vulnerable to population decline than Blue‐winged Warblers because they spend almost twice as many days migrating.     

Geolocators reveal migratory connectivity between wintering and breeding areas of Golden‐winged Warblers

The conservation of migratory songbirds is often impeded by a lack of understanding of how populations in breeding and wintering areas are geographically linked (migratory connectivity). In recent years, light‐level geolocators have improved our understanding of migratory connectivity. Such information is valuable for evaluating how conservation efforts align between the breeding and non‐breeding areas of at‐risk species, and help to more effectively prioritize the allocation of conservation funding. Golden‐winged Warblers (Vermivora chrysoptera) are imperiled migratory songbirds, but the extent to which conservation efforts in their breeding and non‐breeding areas coincide with patterns of migratory connectivity are not well known. We used light‐level geolocators to evaluate the extent to which conservation actions targeting Golden‐winged Warblers in Nicaragua and in their breeding range in North America align with patterns of migratory connectivity. We recovered six of 22 geolocators that had been deployed on male Golden‐winged Warblers at the El Jaguar Reserve during the winter of 2015–2016. All six males migrated to breeding areas in the western Great Lakes region that includes eastern Minnesota, northern Wisconsin, southwestern Ontario, and Michigan's Upper Peninsula. All six males also had similar migration routes, with spring stopovers in southern Mexico, Guatemala, and Belize, a trans‐Gulf flight, and a stopover in the region of Louisiana, Arkansas, eastern Oklahoma, and Texas. Our results, in combination with those of previous studies, demonstrate strong migratory connectivity between portions of the breeding and winter distributions of Golden‐winged Warblers currently targeted for conservation. However, additional studies are needed to improve our understanding of the stopover ecology of Golden‐winged Warblers, especially in areas where they remain for extended periods of time. Finally, patterns of migratory connectivity revealed in our study should be used in combination with existing demographic parameters for Golden‐winged Warblers in the western Great Lakes and Nicaragua to help inform full life cycle population models for this imperiled songbird.     

Migration strategies and annual space‐use in an Afro‐Palaearctic aerial insectivore – the European nightjar Caprimulgus europaeus

Obligate insectivorous birds breeding in high latitudes travel thousands of kilometres during annual movements to track the local seasonal peaks of food abundance in a continuously fluctuating resource landscape. Avian migrants use an array of strategies when conducting these movements depending on e.g. morphology, life history traits and environmental factors encountered en route. Here we used geolocators to derive data on the annual space‐use, temporal pattern and migratory strategies in an Afro‐Palaearctic aerial insectivorous bird species – the European nightjar Caprimulgus europaeus. More specifically, we aimed to test a set of hypothesises pertaining to the migration of a population of nightjars breeding in south‐eastern Sweden. We found that the birds wintered across the central and western parts of the southern tropical Africa almost entirely outside the currently described wintering range of the species. The nightjars performed a narrow loop migration across Sahara, with spring Sahel stopovers significantly to the west of autumn stops indicative to an adaptive response to winds during migration. To our surprise, the migration speed was faster in the autumn (119 km d^{? 1}) than in the spring (99 km d^{? 1}), possibly due to the prevailing wind regimes over the Sahara. The estimated flight fraction in both autumn (14%) and spring (12%) was almost exactly as the theoretically predicted 1:7 time relationship between flights and stopovers for small birds. The temporal patterns within the annual cycle indicate that individuals follow alternative spatiotemporal schedules that converge towards the breeding season. The positive relationship between the spatially and temporally distant winter departure and breeding arrival suggests that individuals´ temporal fine‐tuning to breeding may be constrained, leading to potential negative fitness consequences.     

Acid dissociation constants and pH values for standard “bes” and “tricine” buffer solutions in 30, 40, and 50 mass% dimethyl sulfoxide/water between 25 and −25 °C

Information is given concerning two standard buffer solutions suitable as pH references in 30, 40, and 50 mass% dimethyl sulfoxide (DMSO)/H₂O mixed solvents at subzero temperatures from −20 to 0 °C, with the intention of establishing a pH (designated pH^*) scale. The two buffers selected were the ampholytes N,N-bis(2-hydroxyethyl)-2-aminoethane sulfonic acid (“bes”) and N-tris(hydroxymethyl)methylglycine (“tricine”), and the reference standard consisted of equal molal quantities of the buffer and its respective sodium salt. The assignment of pH^* values was based on measurements of the emf of cells without liquid junction of the type: Pt;H₂(g,1 atm) ¦Bes, Na Besate, NaCl ¦ AgCl;Ag and Pt;H₂(g,1 atm) ¦Tricine, Na Tricinate, NaCl ¦AgCl;Ag and the pH^* was derived from a determination of K₂, the equilibrium constant for the dissociation process (Buffer)^±/ai (Buffer)⁻ + H⁺.     

Migrating songbirds on stopover prepare for,and recover from,oxidative challenges posed by long‐distance flight

Managing oxidative stress is an important physiological function for all aerobic organisms, particularly during periods of prolonged high metabolic activity, such as long‐distance migration across ecological barriers. However, no previous study has investigated the oxidative status of birds at different stages of migration and whether that oxidative status depends on the condition of the birds. In this study, we compared (1) energy stores and circulating oxidative status measures in (a) two species of Neotropical migrants with differing migration strategies that were sampled at an autumn stopover site before an ecological barrier; and (b) a species of trans‐Saharan migrant sampled at a spring stopover site after crossing an ecological barrier; and (2) circulating oxidative measures and indicators of fat metabolism in a trans‐Saharan migrant after stopovers of varying duration (0–8 nights), based on recapture records. We found fat stores to be positively correlated with circulating antioxidant capacity in Blackpoll Warblers and Red‐eyed Vireos preparing for fall migration on Block Island, USA, but uncorrelated in Garden Warblers on the island of Ponza, Italy, after a spring crossing of the Sahara Desert and Mediterranean Sea. In all circumstances, fat stores were positively correlated with circulating lipid oxidation levels. Among Garden Warblers on the island of Ponza, fat anabolism increased with stopover duration while oxidative damage levels decreased. Our study provides evidence that birds build antioxidant capacity as they build fat stores at stopover sites before long flights, but does not support the idea that antioxidant stores remain elevated in birds with high fuel levels after an ecological barrier. Our results further suggest that lipid oxidation may be an inescapable hazard of using fats as the primary fuel for flight. Yet, we also show that birds on stopover are capable of recovering from the oxidative damage they have accrued during migration, as lipid oxidation levels decrease with time on stopover. Thus, the physiological strategy of migrating songbirds may be to build prophylactic antioxidant capacity in concert with fuel stores at stopover sites before a long‐distance flight, and then repair oxidative damage while refueling at stopover sites after long‐distance flight.     

Garden Warbler Sylvia borin migration in sub-Saharan West Africa: phenology and body mass changes

The Garden Warbler is a classic subject for the study of Palaearctic–African bird migration strategies. Most studies have considered the situation close to the breeding areas, while the African and especially the sub‐Saharan part of the species’ migration have received comparatively little attention. Here we use autumn and spring ringing data from Nigeria and The Gambia to study the movements and energetics of the species in West Africa during the non‐breeding season. The first Garden Warblers arrive south of the desert around the beginning of September, roughly at the same time as the median date for their passage through the Baltic Sea region and c. 3 weeks before their median passage date through southern Italy. In the Nigerian Sahel savannahs, where, owing to the rainy season and its associated increase in food availability, many more Garden Warblers stop over in autumn than in the dry spring, the median date of passage is 1 October. The body mass on arrival south of the desert is normally only a few grams more than the lean body mass (LBM; 15 g) – with a mean of 16.6 g (sd = ±1.8 g) in The Gambia and 17.4 g (sd = ±1.8 g) in the Nigerian Sahel. After resting and refuelling in the Sahel, Sudan and Guinea‐type savannahs the Garden Warblers depart during November–December for wintering areas further south. Before leaving, they again increase their body mass, with an average fuel load of c. 20%, and often more than 50% relative to LBM. Some of the birds passing through Nigeria probably spend midwinter around the Congo Basin. During spring they return northwards to the Guinea savannah zone in April and fuel‐up there for the trans‐Sahara passage. At this time they normally increase their body reserves to around 50% of the LBM, but c. 10% of the birds gain 100%, thus doubling their mass. The passage there peaks around 20 April and continues well into May. That the main take‐off northwards is directly from the Guinea savannahs is indicated by the very low numbers trapped in the Sahel during spring.     

WEIGHTS OF SOME PALAEARCTIC MIGRANTS IN SOUTHERN UGANDA

Between March 1966 and May 1968 Palaearctic passerines were mist-netted in thick bush and lightly wooded savannah habitats near Kampala, on the northern shore of Lake Victoria. This paper reports weights of the seven principal species involved. Most migrants appeared to be in a lean condition during the winter months, when weights were relatively low and varied little in each species. Birds were not particularly light on arrival. In fact, autumn Garden Warblers Sylvia borin and Willow Warblers Phylloscopus trochilus were sometimes markedly heavy, and for the former species there was some evidence that the individuals concerned were passage migrants. Autumn weights of Swallows Hirundo rustica, Reed Warblers Acrocephalus scirpaceus and Yellow Wagtails Motacilla flava were similar to those recorded in winter. The mean weight of all species rose during late March or early April. Although most Garden Warblers and Willow Warblers trapped at the time of spring migration were within the normal winter weight range, many Acrocephalus warblers and the majority of Sand Martins Riparia riparia and Yellow Wagtails were rather heavy. Spring weights 40% or more above mean winter weight were not uncommon in the Sedge Warbler Acrocephalus schoenobaenus, but were recorded only occasionally in other species. Although most passerine migrants evidently left Kampala with substantial fat reserves, it was concluded that a considerable number of warblers departed at rather low weight. High spring weights were mainly confined to a period of two or three weeks in each of the warbler species. Locally wintering Acrocephalus warblers must have attained full premigratory weights within three weeks, and a number of spring retraps showed substantial gains at minimum mean rates of between 0–1 and 0–35 g per day. Most heavy Garden Warblers were probably on passage. Significant correlations between weight and wing-length were obtained for all species investigated, regressions of weight on wing-length being in the range 011-0-25 g/mm. Spring weights are briefly compared with data from Nigeria, and the northward migration of passerines from Lake Victoria is discussed.     

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.     

Foraging behavior of Cerulean Warblers during the breeding and non‐breeding seasons: evidence for the breeding currency hypothesis

Birds require additional resources for raising young, and the breeding currency hypothesis predicts that insectivorous species exploit large soft‐bodied prey during the breeding season, but shift to small, likely hard‐bodied, prey during the non‐breeding season. To test this hypothesis, we examined prey use by Cerulean Warblers (Setophaga cerulea), foliage‐gleaning Nearctic‐Neotropical migrants, during the breeding and non‐breeding seasons. We collected data on foraging behavior during the breeding season (including observations of prey items fed to young) in upland mixed‐oak forest in southeastern Ohio in 2009 and 2010 and, during the non‐breeding season, in shade coffee in the Cordillera de Merida, Venezuela, in 2008–2009. Cerulean Warblers captured 7% more large prey (visible prey extending beyond the bill) during the breeding than the non‐breeding season, but foraged at similar rates during both seasons. Large, soft‐bodied prey appeared to be especially important for feeding young. We found that adults delivered large prey on >50% of provisioning visits to nests and 69% of identifiable large prey fed to nestlings were greenish larvae (likely Lepidoptera or caterpillars) that camouflage against leaves where they would tend to be captured by foliage‐gleaning birds. Availability of specific taxa appeared to influence tree species foraging preferences. As reported by other researchers, we found that Cerulean Warblers selected trees in the genus Carya for foraging and our examination of caterpillar counts from the central Appalachian Mountains (Butler and Strazanac 2000 ) showed that caterpillars with greenish coloration, especially Baileya larvae, may be almost twice as abundant on Carya than Quercus. Our results provide evidence for the breeding currency hypothesis, and highlight the importance of caterpillars to a foliage‐gleaning migrant warbler of conservation concern.     

Song in two cryptic species: comparative analysis of Large-billed Reed Warblers Acrocephalus orinus and Blyth’s Reed Warblers Acrocephalus dumetorum

Capsule: Songs of Large-billed Reed Warblers Acrocephalus orinus and Blyth’s Reed Warblers Acrocephalus dumetorum differed in quantitative parameters. Blyth’s Reed Warbler used different modes of singing on breeding grounds and migration stopovers.

Aims: To compare the songs of two cryptic species of reed warblers. To compare Blyth’s Reed Warbler songs in different parts of the breeding range and on migration stopovers in central Asia. To investigate the status of Blyth’s Reed Warblers in central Asia.

Methods: We analysed song recordings of individual singing males and conducted field observations of singing behaviour at migration stopover and breeding sites. The status of Blyth’s Reed Warblers in central Asia was reviewed from the historical Russian literature and other sources.

Results: We found differences between six out of eight variables in songs of Large-billed and Blyth’s Reed Warblers on breeding sites. Blyth’s Reed Warbler songs recorded on migration stopovers showed differences in five variables from Large-billed Reed Warblers and in six variables from breeding Blyth’s Reed Warblers. On migration stopovers, Blyth’s Reed Warblers sang actively but did not stay more than 1–2 days. The evidence suggests that Blyth’s Reed Warbler does not breed in central Asia.

Conclusion: Songs of Large-billed and Blyth’s Reed Warblers could be distinguished by quantitative analysis. Blyth’s Reed Warblers used different modes of singing on breeding sites and migration stopovers. We believe vocalizations of Blyth’s Reed Warblers on migration to be plastic song.     

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 strength of migratory connectivity for birds en route to breeding through the Gulf of Mexico

The strength of migratory connectivity is a measure of the cohesion of populations among phases of the annual cycle, including breeding, migration, and wintering. Many Nearctic‐Neotropical species have strong migratory connectivity between breeding and wintering phases of the annual cycle. It is less clear if this strength persists during migration when multiple endogenous and exogenous factors may decrease the cohesion of populations among routes or through time along the same routes. We sampled three bird species, American redstart Setophaga ruticilla, ovenbird Seiurus aurocapilla, and wood thrush Hylocichla mustelina, during spring migration through the Gulf of Mexico region to test if breeding populations differentiate spatially among migration routes or temporally along the same migration routes and the extent to which within‐population timing is a function of sex, age, and carry‐over from winter habitat, as measured by stable carbon isotope values in claws (δ¹³C). To make quantitative comparisons of migratory connectivity possible, we developed and used new methodology to estimate the strength of migratory connectivity (MC) from probabilistic origin assignments identified using stable hydrogen isotopes in feathers (δ²H). We found support for spatial differentiation among routes by American redstarts and ovenbirds and temporal differentiation along routes by American redstarts. After controlling for breeding origin, the timing of American redstart migration differed among ages and sexes and ovenbird migration timing was influenced by carry‐over from winter habitat. The strength of migratory connectivity did not differ among the three species, with each showing weak breeding‐to‐spring migration MC relative to prior assessments of breeding‐wintering connectivity. Our work begins to fill an essential gap in methodology and understanding of the extent to which populations remain together during migration, information critical for a full annual cycle perspective on the population dynamics and conservation of migratory animals.     

Diurnal space use and nocturnal roost‐site selection by male Cerulean Warblers during the breeding season

Detailed information about space use during the breeding season is limited for most Nearctic‐Neotropical migratory species of songbirds because of their small size and often cryptic behaviors. We monitored male Cerulean Warblers (Setophaga cerulea), a species of conservation concern, using radio‐telemetry during the 2006–2008 breeding seasons in northern Alabama to better understand their space use and habitat selection. We estimated diurnal home range and core areas using information theoretic criteria, located nocturnal roost sites, and related day and evening locations to surrounding landscape habitat, including features representative of canopy disturbances. Mean home range size was 6.7 ha (N = 10), and home ranges included an average of at least 2 core areas encompassing 0.7 ha. We located 53 nocturnal roost sites that were an average 159.0 m from the center of the nearest core area. More than one‐third (36.6%) of roost sites were located outside the diurnal home ranges of male Cerulean Warblers; only 13.6% were located in core areas. Males in our study moved much farther than reported in previous studies, with some singing in areas > 300 m from previously used song perches, a behavior suggesting pursuit of extra‐pair copulations. Cerulean Warblers in our study preferentially selected a heavily forested landscape composed of mesic, floodplain bottomlands with little man‐made disturbance. Within their home ranges, diurnal locations of males in core areas were located significantly closer to a creek than locations outside of core areas. Our results suggest that male Cerulean Warblers require much larger areas than previously reported and underscore the importance of a predominately forested landscape in their habitat selection process. Although edge habitats appeared to influence space use by male Cerulean Warblers in our study, the extent to which this is an essential requirement is unclear. Our results and those of previous studies suggest that specific habitat requirements of this species can vary at the local scale throughout its breeding range.