Stopover duration,movement patterns and temporary home ranges of fall migrant yellow‐rumped warblers Setophaga coronata in native and anthropogenic woodlands of the Northern Prairie region,USA

Stopover behavior of migrant birds is influenced by their energetic condition, but also by extrinsic factors, including weather conditions and habitat attributes such as vegetation structure, microclimates, predation pressure, competition, and food availability. Anthropogenic habitats may differ from natural habitats in these attributes, which could promote differing stopover behaviors for migrants in the two habitat types and affect overall habitat suitability. We used radio‐telemetry to measure stopover behaviors of fall migrant yellow‐rumped warblers Setophaga coronata in native riparian corridor woodlands (corridors) and anthropogenic woodlots (woodlots) in the Northern Prairie region. We measured stopover duration, movement rate, and temporary home range size for birds in both habitat types by attaching radio‐transmitters and relocating birds to either corridor (n = 17) or woodlot (n = 16) study sites. We used AIC_C to rank null, global, and reduced models, which included habitat type, energetic condition, habitat size, year, date, and movement rate (for stopover duration analyses only) as explanatory variables. Model rankings showed that habitat type was not included in any of the top models (ΔAIC_C < 2) for movement behavior, temporary home range size, or stopover duration, which suggests similar functional habitat quality between the two habitat types. These data add similar behavioral responses for birds in the two habitat types to similar fattening rates and stress physiology, further confirming similar suitability of native and anthropogenic woodland habitats in this region as stopover habitat. We also applied logistic regression with a model selection approach, including cloud cover, tail wind component, temperature, and barometric pressure as independent variables, and departure decision as the dependent variable, to evaluate the effects of weather variables on departure. Model selection suggested that cloud cover is a prominent factor affecting departure decisions and the other variables may also influence departure decisions of yellow‐rumped warblers from inland stopover sites.     

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.     

Carry‐over effects of winter habitat quality on en route timing and condition of a migratory passerine during spring migration

We examined how conditions prior to migration influenced migration performance of two breeding populations of black‐and‐white warblers Mniotilta varia by linking information on the migrant's winter habitat quality, measured via stable carbon isotopes, with information on their breeding destination, measured via stable hydrogen isotopes. The quality of winter habitat strongly influenced the timing of migration when we accounted for differential timing of migration between breeding populations. Among birds migrating to the same breeding destination, males and females arriving early to the stopover site originated from more mesic habitat than later arriving birds, suggesting that the benefits of occupying high‐quality mesic habitat during the winter positively influence the timing of migration. However, male warblers arriving early to the stopover site were not in better migratory condition than later arriving conspecifics that originated from poor‐quality xeric winter habitat, regardless of breeding destination. The two breeding populations stopover at the study site during different time periods, suggesting that the lower migratory condition of early birds is not a function of the time of season, but potentially a migrant's migration strategy. Strong selection pressures to arrive early on the breeding grounds to secure high‐quality breeding territories may drive males from high‐quality winter habitat to minimize time at the expense of energy. This migration strategy would result in a smaller margin of safety to buffer the effects of adverse weather or scarcity of food, increasing the risk of mortality. The migratory condition of females was the same regardless of the timing of migration or breeding destination, suggesting that females adopt a strategy that conserves energy during migration. This study fills an important gap in our understanding of the linkages between winter habitat quality and factors that influence the performance of migration, the phase of the annual cycle thought to be limiting most migratory bird populations.     

Connecting the dots: Stopover strategies of an intercontinental migratory songbird in the context of the annual cycle

The phases of the annual cycle for migratory species are inextricably linked. Yet, less than five percent of ecological studies examine seasonal interactions. In this study, we utilized stable hydrogen isotopes to geographically link individual black‐and‐white warblers (Mniotilta varia) captured during spring migration with breeding destinations to understand a migrant's stopover strategy in the context of other phases of the annual cycle. We found that stopover strategy is not only a function of a bird's current energetic state, but also the distance remaining to breeding destination and a bird's time‐schedule, which has previously been linked to habitat conditions experienced in the preceding phase of the annual cycle. Birds in close proximity to their breeding destination accumulate additional energy reserves prior to arrival on the breeding grounds, as reflected by higher migratory condition upon arrival, higher refueling rates measured via blood plasma metabolites, and longer stopover durations compared to birds migrating to breeding destinations farther from the stopover site. However, late birds near their breeding destination were more likely to depart on the day of arrival (i.e., transients), and among birds that stopped over at the site, the average duration of stopover was almost half the time of early conspecifics, suggesting late birds are trying to catch‐up with the overall time‐schedule of migration for optimal arrival time on the breeding grounds. In contrast, birds with long distances remaining to breeding destinations were more likely to depart on the day of arrival and primarily used stopover to rest before quickly resuming migration, adopting similar strategies regardless of a bird's time‐schedule. Our study demonstrates that migrants adjust their en route strategies in relation to their time‐schedule and distance remaining to their breeding destination, highlighting that strategies of migration should be examined in the context of other phases of the annual cycle.     

Individual consistency of long‐distance migration in a songbird: significant repeatability of autumn route,stopovers and wintering sites but not in timing of migration

Through new tracking techniques, data on timing and routes of migration in long‐distance migrant birds are accumulating. However, studies of the consistency of migration of the same individuals between years are still rare in small‐sized passerine birds. This type of information is important to understand decisions and migration abilities at the individual level, but also for life history theory, for understanding carry over effects between different annual cycle stages and for conservation. We analysed individual repeatability of migration between years in great reed warblers Acrocephalus arundinaceus; a medium‐sized European songbird migrating to sub‐Saharan Africa. In seven males, with geolocator data from 2–4 yr per bird, we found low to moderate (non significant) repeatability in timing of migration parameters (R ≤ 0.41), but high (and significant) repeatability for most spatial parameters, i.e. autumn route (R = 0.64) and stopover sites (R = 0.59–0.87) in Europe, and wintering sites (R = 0.77–0.99) in sub‐Saharan Africa. This pattern of high spatial but low temporal within‐individual repeatability of migration between years contrasts other tracking studies of migrating birds that generally have found consistency in timing but flexibility in routes. High spatial consistency of migration in the great reed warbler may be due to it being a specialist in wetlands, an unevenly distributed habitat, favouring a strategy of recurrence at previously visited sites. Low temporal repeatability may be caused by large between‐year variation in carry‐over effects from the breeding season, high flexibility in decision rules during migration or high sensitivity to environmental factors (weather, wind) during migration.     

The effects of dietary macronutrients on flight ability,energetics, and fuel metabolism of yellow‐rumped warblers Setophaga coronata

The catabolism of protein from organs and muscles during migratory flight is necessary to produce glucose, key metabolic intermediates, and water, but may have negative effects on flight range and refueling at stopovers. We tested the hypothesis, suggested by previous studies, that birds that eat high‐protein insect diets use more protein for fuel in flight than those that eat high‐carbohydrate fruits. First, we fed migratory yellow‐rumped warblers synthetic fruit or mixed insect/fruit diets, and measured metabolic rates and fuel mixture under basal conditions and during exercise in a hop/hover wheel respirometer. Birds eating the fruit diet had greater plasma triglyceride and non‐esterified fatty acid concentrations, and the higher protein mixed diet increased plasma uric acid only during feeding. Diet did not affect metabolic rates or the fuel mixture under resting or exercise conditions. We then fed yellow‐rumped warblers synthetic diets that differed only in the relative proportion of carbohydrate and protein (60:15 versus 15:60 as % dry mass) and tested them in wind tunnel flights lasting up to six hours. Birds fed the high carbohydrate diet became heavier and fatter than when fed the high protein diet. Plasma uric acid concentration was increased and plasma phospholipid concentration was decreased by the high protein diet in the pre‐flight state (after a 3 h fast), but diet only affected plasma phospholipids during flight (lower in high protein birds). Neither diet nor amount of body fat affected the rate of loss of lean mass or fat during flight. Inter‐individual or seasonal differences in diet do not appear to influence the amount of protein catabolized during endurance flight. However, birds fed the high carbohydrate diet had greater voluntary flight duration, independent of body fatness, suggesting that there may be other performance benefits of high carbohydrate diets for migratory birds.     

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.     

Breeding latitude and timing of spring migration in songbirds crossing the Gulf of Mexico

Each spring, millions of songbirds migrate across the Gulf of Mexico on their way to breeding sites in North America. Data from radar and migration monitoring stations have revealed broad patterns in the spatial and temporal course of trans-Gulf migration. Unfortunately, we have limited information on where these birds have previously spent the winter and where they are migrating to breed. Here we measure stable-hydrogen isotopes in feathers (δD_f) to infer the breeding latitude of five species of songbirds – hooded warblers Wilsonia citrina , American redstarts Setophaga ruticilla , black-and-white warblers Mniotilta varia , ovenbirds Seiurus aurocapilla , and northern waterthrushes S. noveboracensis – that were captured at a stopover site along the coast of southwestern Louisiana in spring 2004. Values of δD_f across all species ranged from −163 to −35‰ (n=212), and within most species the range was consistent with the latitudinal extent of known breeding sites in central and eastern North America. Individuals that arrived first along the northern Gulf coast had δD_f values indicative of southerly breeding sites in hooded warblers, American redstarts, black-and-white warblers, and ovenbirds, but no relationship was found between passage timing and δD_f for northern waterthrushes. Our findings suggest that spring passage is often timed to coincide with the emergence of suitable conditions on breeding areas, with southern breeding birds migrating first.     

Body condition and feather molt of a migratory shorebird during the non‐breeding season

Migratory shorebirds have some of the highest fat loads among birds, especially species which migrate long distances. The upland sandpiper Bartramia longicauda makes long‐distance migrations twice a year, but variation in body condition or timing of feather molt during the non‐breeding season has not been studied. Molt is an important part of the annual cycle of migratory birds because feather condition determines flight performance during migration, and long‐distance movements are energetically costly. However, variation in body condition during molt has been poorly studied. The objective of our field study was to examine the timing and patterns of feather molt of a long distance migratory shorebird during the non‐breeding season and test for relationships with body size, fat depots, mass, and sex. Field work was conducted at four ranches in the Northern Campos of Uruguay (Paysandú and Salto Departments). We captured and marked 62 sandpipers in a 2‐month period (Nov–Jan) during four non‐breeding seasons (2008–2012). Sex was determined by genetic analyses of blood samples taken at capture. Molt was measured in captured birds using rank scores based on published standards. Body mass and tarsus length measurements showed female‐biased sexual size dimorphism with males smaller than females. Size‐corrected body mass (body condition) showed a U‐shaped relationship with the day of the season, indicating that birds arrived at non‐breeding grounds in relatively good condition. Arriving in good body condition at non‐breeding grounds is probably important because of the energetic demands due to physiological adjustments after migration and the costs of feather molt.     

Migration distance does not predict blood parasitism in a migratory songbird

Migration can influence host–parasite dynamics in animals by increasing exposure to parasites, by reducing the energy available for immune defense, or by culling of infected individuals. These mechanisms have been demonstrated in several comparative analyses; however, few studies have investigated whether conspecific variation in migration distance may also be related to infection risk. Here, we ask whether autumn migration distance, inferred from stable hydrogen isotope analysis of summer‐grown feathers (δ²H_f) in Europe, correlates with blood parasite prevalence and intensity of infection for willow warblers (Phylloscopus trochilus) wintering in Zambia. We also investigated whether infection was correlated with individual condition (assessed via corticosterone, scaled mass index, and feather quality). We found that 43% of birds were infected with Haemoproteus palloris (lineage WW1). Using generalized linear models, we found no relationship between migration distance and either Haemoproteus infection prevalence or intensity. There was spatial variation in breeding ground origins of infected versus noninfected birds, with infected birds originating from more northern sites than noninfected birds, but this difference translated into only slightly longer estimated migration distances (~214 km) for infected birds. We found no relationship between body condition indices and Haemoproteus infection prevalence or intensity. Our results do not support any of the proposed mechanisms for migration effects on host–parasite dynamics and cautiously suggest that other factors may be more important for determining individual susceptibility to disease in migratory bird species.     

Across a migratory divide: divergent migration directions and non‐breeding grounds of Eurasian reed warblers revealed by geolocators and stable isotopes

Migratory divides represent narrow zones of overlap between parapatric populations with distinct migration directions and, consequently, expected divergent non‐breeding distributions. The composition of the mixed population at a migratory divide and the corresponding non‐breeding ranges remain, however, unknown for many Palaearctic‐African migrants. Here, we used light‐level geolocation to track migration direction and non‐breeding grounds of Eurasian reed warblers Acrocephalus scirpaceus from three breeding populations across the species’ migratory divide. Moreover, by using feathers grown at non‐breeding grounds, we quantified stable isotope composition for individuals with known southwestern (SW) and southeastern (SE) migration directions. On a larger sample per population, we then assessed the proportions of SW‐ and SE‐migrating phenotypes in each of the three populations. All tracked reed warblers from Germany and two thirds of the birds tagged from the Czech population headed initially SW. Nevertheless, about one third of the birds from the Czech site migrated towards SE. No tracking data have been obtained for the Bulgarian population. The initial migration direction determined by geolocators was a strong predictor of the non‐breeding region, with SW migrants staying in west Africa and SE migrants in central Africa. Feather δ³⁴S and δ¹⁵N values confirmed the predominance of SW migrants in the German population, the co‐occurrence of SW and SE migrants in the Czech population, and indicated a high (72%) proportion of SE migrants in the Bulgarian population. Thus, the combined approach of geolocator tracking and stable isotopic assignments provided clear evidence for the existence of a migratory divide in the southeast of central Europe and predicted non‐breeding range in central and central‐eastern Africa for the eastern population.     

An ‘ecological trap’ for yellow warbler nest microhabitat selection

Contrary to assumptions of habitat selection theory, field studies frequently detect ‘ecological traps’, where animals prefer habitats conferring lower fitness than available alternatives. Evidence for traps includes cases where birds prefer breeding habitats associated with relatively high nest predation rates despite the importance of nest survival to avian fitness. Because birds select breeding habitat at multiple spatial scales, the processes underlying traps for birds are likely scale‐dependent. We studied a potential ecological trap for a population of yellow warblers Dendroica petechia while paying specific attention to spatial scale. We quantified nest microhabitat preference by comparing nest‐ versus random‐site microhabitat structure and related preferred microhabitat features with nest survival. Over a nine‐year study period and three study sites, we found a consistently negative relationship between preferred microhabitat patches and nest survival rates. Data from experimental nests described a similar relationship, corroborating the apparent positive relationship between preferred microhabitat and nest predation. As do other songbirds, yellow warblers select breeding habitat in at least two steps at two spatial scales; (1) they select territories at a coarser spatial scale and (2) nest microhabitats at a finer scale from within individual territories. By comparing nest versus random sites within territories, we showed that maladaptive nest microhabitat preferences arose during within‐territory nest site selection (step 2). Furthermore, nest predation rates varied at a fine enough scale to provide individual yellow warblers with lower‐predation alternatives to preferred microhabitats. Given these results, tradeoffs between nest survival and other fitness components are unlikely since fitness components other than nest survival are probably more relevant to territory‐scale habitat selection. Instead, exchanges of individuals among populations facing different predation regimes, the recent proliferation of the parasitic brown‐headed cowbird Molothrus ater, and/or anthropogenic changes to riparian vegetation structure are more likely explanations.     

Digestive response to restricted feeding in migratory yellow-rumped warblers

Smaller guts and slow initial mass gains at stopover sites have led to the idea that digestive physiology limits refueling rates in migrating birds. We tested the digestive-limitation hypothesis in yellow-rumped warblers using food restriction to simulate infrequent feeding during migration, which may cause a reduction in alimentary tract mass. Restricted birds had small intestine, pancreas, and liver masses 18%-22% lower than ad lib.-fed controls. Total activities of sucrase, maltase, aminopeptidase, and amylase were significantly lower in restricted birds, while those of trypsin and chymotrypsin were not. Only aminopeptidase mass-specific activity was significantly lower in restricted birds. Previously restricted birds were able to feed and digest at a high rate immediately following return to ad lib. feeding. Digestive efficiency did not differ between groups. These results suggest that before migration yellow-rumped warblers have some spare digestive capacity to compensate for declines in their digestive organ masses during migration.     

Adcyap1 polymorphism covaries with breeding latitude in a Nearctic migratory songbird,the Wilson's warbler (Cardellina pusilla)

Understanding the genetic background of complex behavioral traits, showing multigenic control and extensive environmental effects, is a challenging task. Among such traits, migration is known to show a large additive genetic component. Yet, the identification of specific genes or gene regions explaining phenotypic variance in migratory behavior has received less attention. Migration ultimately depends on seasonal cycles, and polymorphism at phenological candidate genes may underlie variation in timing of migration or other aspects of migratory behavior. In this study of a Nearctic–Neotropical migratory songbird, the Wilson's warbler (Cardellina pusilla), we investigated the association between polymorphism at two phenological candidate genes, Clock and Adcyap1, and two aspects of the migratory phenotype, timing of spring migration through a stopover site and inferred latitude of the breeding destination. The breeding destination of migrating individuals was identified using feather deuterium ratio (δ²H), which reliably reflects breeding latitude throughout the species' western breeding range. Ninety‐eight percent of the individuals were homozygous at Clock, and the rare heterozygotes did not deviate from homozygous migration phenology. Adcyap1 was highly polymorphic, and allele size was not significantly associated with migration date. However, Adcyap1 allele size significantly positively predicted the inferred breeding latitude of males but not of females. Moreover, we found a strong positive association between inferred breeding latitude and Adcyap1 allele size in long‐distance migrating birds from the northern sector of the breeding range (western Canada), while this was not the case in short‐distance migrating birds from the southern sector of the breeding range (coastal California). Our findings support previous evidence for a role of Adcyap1 in shaping the avian migratory phenotype, while highlighting that patterns of phenological candidate gene–phenotype associations may be complex, significantly varying between geographically distinct populations and even between the sexes.     

Differentiating between stopover and staging sites: functions of the southern and northern Yellow Sea for long‐distance migratory shorebirds

Evidence‐based protection of migratory birds at flyway levels requires a solid understanding of their use of ‘stopping sites’ during migration. To characterize the site use of northward‐migration great knots Calidris tenuirostris in China, we compared length of stay and fuel deposition during northward migration at areas in the south and the north of the Yellow Sea, a region critical for migrating shorebirds. Radio‐tracking showed that at the southern site great knots stayed for only short periods (2.3 ± 1.9 d, n = 40), and bird captures showed that they did not increase their mean body mass while there. In the north birds stayed for 1 month (31.0 ± 13.6 d, n = 22) and almost doubled their mean body mass. Fuel consumption models suggest that great knots departing from the northern Yellow Sea should be able to fly nonstop to the breeding grounds, whereas those from the south would require a refueling stop further north. These results indicate that the study sites in the northern and southern Yellow Sea serve different roles: the southern site acts as a temporary stopover area that enables birds with low fuel stores to make it to main staging areas further north, while the northern site serves as the critical staging site where birds refuel for the next leg of their migration. The rapid turnover rate in the southern Yellow Sea indicates that many more birds use that area than are indicated by peak counts. Differential use of the southern and northern sites indicates that both play crucial roles in the ability of great knots to migrate successfully.     

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.     

MIGRATION,MITOCHONDRIA, AND THE YELLOW‐RUMPED WARBLER

Discordance between mitochondrial and nuclear DNA has been noted in many systems. Asymmetric introgression of mitochondria is a common cause of such discordances, although in most cases the drivers of introgression are unknown. In the yellow‐rumped warbler, evidence suggests that mtDNA from the eastern, myrtle warbler, has introgressed across much of the range of the western form, the Audubon's warbler. Within the southwestern United States myrtle mtDNA comes into contact with another clade that occurs in the Mexican black‐fronted warbler. Both northern forms exhibit seasonal migration, whereas black‐fronted warblers are nonmigratory. We investigated the link between mitochondrial introgression, mitochondrial function, and migration using novel genetic, isotopic, biochemical, and phenotypic data obtained from populations in the transition zone. Isotopes suggest the zone is coincident with a shift in migration, with individuals in the south being resident and populations further north becoming increasingly more migratory. Mitochondrial respiration in flight muscles demonstrates that myrtle‐type individuals have a significantly greater acceptor control ratio of mitochondria, suggesting it may be more metabolically efficient. To our knowledge this is the first time this type of intraspecific variation in mitochondrial respiration has been measured in wild birds and we discuss how such mitochondrial adaptations may have facilitated introgression.     

Carry‐over effects of winter habitat vary with age and sex in yellow warblers Setophaga petechia

We use stable isotope data to investigate the role of winter habitat use in altering the breeding phenology of yellow warblers Setophaga petechia. We first confirm that δ¹³C and δ¹⁵N isotopic signatures vary with winter habitat use in this species. We then examine the relationship between winter habitat use, breeding phenology and productivity within four age‐sex‐classes, since life history theory would predict that carry‐over effects should vary with age and gender. The δ¹³C signatures of yellow warblers using riparian habitats over winter were more depleted than the signatures of those using agricultural or scrub habitat. Individuals on the Pacific coast of Mexico were also more δ¹⁵N enriched than those on the southern Gulf of Mexico. δ¹³C and δ¹⁵N signatures were only correlated with earlier clutch initiation and subsequent higher productivity in first‐breeding‐season females. We estimate that shifts in δ¹³C equivalent to a shift from scrub to riparian winter habitat would be associated with the production of 0.8 more fledglings by yearling females. Pre‐breeding events that influence the timing of breeding could also influence the reproductive performance of older males and females, but we found little evidence that winter habitat use influenced breeding season phenology in these birds.     

Stable isotope ratios indicate that body condition in migrating passerines is influenced by winter habitat

Although predicted some time ago, there has been little success in demonstrating that the overall fitness of migratory birds depends on the combined influences of their experiences over all seasons. We used stable carbon isotope signatures (delta13C) in the claws of migrating black-throated blue warblers Dendroica caerulescens to infer their wintering habitats and investigated whether winter habitat selection can be linked to condition during migration. Resident bird species with low delta13C corresponded to selection of more mesic habitats, and migrating birds with low delta13C were in better condition than conspecifics with higher delta13C signatures. These findings concur with empirical observations on the wintering grounds, where dominants (mostly males) tend to exclude subordinates from mesic areas (considered to be high-quality habitats). We believe that variation in condition during migration may be one of the key factors determining differences in arrival times and condition at the breeding areas, which in turn have a major influence on reproductive success.     

Cold tolerance,and not earlier arrival on breeding grounds,explains why males winter further north in an Arctic‐breeding songbird

Sex biases in distributions of migratory birds during the non‐breeding season are widespread; however, the proximate mechanisms contributing to broad‐scale sex‐ratio variation are not well understood. We analyzed a long‐term winter‐banding dataset in combination with spring migration data from individuals tracked by using geolocators to test three hypotheses for observed variation in sex‐ratios in wintering flocks of snow buntings Plectrophenax nivalis. We quantified relevant weather conditions in winter (temperature, snowfall and snow depth) at each banding site each year and measured body size and condition (fat scores) of individual birds (n > 5500). We also directly measured spring migration distance for 17 individuals by using light‐level geolocators. If the distribution pattern of birds in winter is related to interactions between individual body size and thermoregulation, then larger bodied birds (males) should be found in colder sites (body size hypothesis). Males may also winter closer to breeding grounds to reduce migration distance for early arrival at breeding sites (arrival timing hypothesis). Finally, males may be socially dominant over females, and thus exclude females from high‐quality wintering sites (social dominance hypothesis). We found support for the body size hypothesis, in that colder and snowier weather predicted both larger body size and higher proportions of males banded. Direct tracking revealed that males did not winter significantly closer to their breeding site, despite being slightly further north on average than females from the same breeding population. We found some evidence for social dominance, in that females tended to carry more fat than males, potentially indicating lower habitat quality for females. Global climatic warming may reduce temperature constraints on females and smaller‐bodied males, resulting in broad‐scale changes in distributional patterns. Whether this has repercussions for individual fitness, and therefore population demography, is an important area of future research.