Dynamic occupancy models reveal within‐breeding season movement up a habitat quality gradient by a migratory songbird

The timing of settlement decisions likely influences the quality of breeding site choices.This is particularly the case in migratory birds, because the conditions that enhance breeding success are often not apparent upon arrival after migration. A strategy that addresses this problem is to adjust settlement decisions when reliable information becomes available. We used a new indirect method – dynamic site occupancy modeling – to estimate apparent movement of black‐throated blue warblers Dendroica caerulescens among sites within a breeding season. Because individuals should disperse to sites that maximize their fitness, we hypothesized that warblers would move up a habitat quality gradient when opportunities arose. For our study species, that would involve moving into sites with greater shrub density and at higher elevation within northern hardwoods forest, as these two features are positively correlated with reproduction and apparent survival in this species. Although the probability of site occupancy in our study landscape remained consistent throughout the breeding season (range: 0.66–0.69), occupancy models revealed substantial support for apparent movement of individuals within the breeding season. The mean probability of emigration from a point count site was 0.21 (±0.03 SE), and the mean probability of immigration to a site not previously occupied was 0.51 (±0.05 SE). The spatial distribution of this movement was a function of habitat quality. A portion of the black‐throated blue warbler population appears to arrive on the breeding grounds and settle initially in sub‐optimal habitat, moving subsequently into high quality densely shrubbed habitat at higher elevations. This modeling approach provides a new means to test hypotheses about habitat selection and movement by using presence–non‐detection data.     

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.     

Annual variation in long‐distance dispersal driven by breeding and non‐breeding season climatic conditions in a migratory bird

Long‐distance dispersal is a fundamental process in ecology and evolution but the factors that influence these movements remain poorly understood in most species. We used stable hydrogen isotopes to quantify the rate and direction of long‐distance immigration in a breeding population of American redstarts and to test whether the settlement decisions that result in long‐distance dispersal are driven by habitat saturation or by the phenology of breeding‐season resources. Our results provide evidence that both natal dispersal and breeding dispersal were influenced by the timing of breeding‐season phenology, with both age classes more likely to disperse north in years when the onset of breeding‐season phenology occurs earlier than normal. Yearlings were also more likely to disperse north following winters with poor habitat quality on their non‐breeding grounds, demonstrating that carry‐over effects from the non‐breeding season influence natal dispersal in this species. Collectively, these results are consistent with the hypothesis that American redstarts use the phenology of breeding season resources as a cue to select breeding sites. Our results suggest that long‐distance dispersal may allow individuals to rapidly respond to advancing phenology caused by global climate change, though their ability to do so may be constrained by long‐term decline in habitat quality predicted for their tropical non‐breeding grounds.     

Late‐season snowfall is associated with decreased offspring survival in two migratory arctic‐breeding songbird species

While the effect of weather on reproduction has been studied for many years in avian taxa, the rapid pace of climate change in arctic regions has added urgency to this question by changing the weather conditions species experience during breeding. Given this, it is important to understand how factors such as temperature, rain, snowfall, and wind affect reproduction both directly and indirectly (e.g. through their effects on food availability). In this study, we ask how weather factors and food availability influence daily survival rates of clutches in two arctic‐breeding migratory songbirds: the Lapland longspur Calcarius lapponicus, a circumpolar breeder, and Gambel's white‐crowned sparrow Zonotrichia leucophrys gambelii, which breeds in shrubby habitats across tundra, boreal and continental climates. To do this, we monitored clutch survival in these two species from egg‐lay through fledge at field sites located near Toolik Field Station (North Slope, Alaska) across 5 yr (2012–2016). Our results indicate that snowfall and cold temperatures decreased offspring survival rates in both species; although Lapland longspurs were more susceptible to snowfall. Food availability, quantified by pitfall sampling and sweep‐net sampling methods, had minimal effects on offspring survival. Some climate models predict increased precipitation for the Arctic with global warming, and in the Toolik region, total snow accumulation may be increasing. Placed in this context, our results suggest that changes in snow storms with climate change could have substantial consequences for reproduction in migratory songbirds breeding in the North American Arctic.     

Faster migration in autumn than in spring: seasonal migration patterns and non‐breeding distribution of Icelandic whimbrels Numenius phaeopus islandicus

Migration is fundamental in the life of many birds and entails significant energetic and time investments. Given the importance of arrival time in the breeding area and the relatively short period available to reproduce (particularly at high latitudes), it is expected that birds reduce spring migration duration to a greater extent than autumn migration, assuming that pressure to arrive into the wintering area might be relaxed. This has previously been shown for several avian groups, but recent evidence from four tracked Icelandic whimbrels Numenius phaeopus islandicus, a long distance migratory wader, suggests that this subspecies tends to migrate faster in autumn than in spring. Here, we 1) investigate differences in seasonal migration duration, migration speed and ground speed of whimbrels using 56 migrations from 19 individuals tracked with geolocators and 2) map the migration routes, wintering and stopover areas for this population. Tracking methods only provide temporal information on the migration period between departure and arrival. However, migration starts with the fuelling that takes place ahead of departure. Here we estimate the period of first fuelling using published fuel deposition rates and thus explore migration speed using tracking data. We found that migration duration was shorter in autumn than in spring. Migration speed was higher in autumn, with all individuals undertaking a direct flight to the wintering areas, while in spring most made a stopover. Wind patterns could drive whimbrels to stop in spring, but be more favourable during autumn migration and allow a direct flight. Additionally, the stopover might allow the appraisal of weather conditions closer to the breeding areas and/or improve body condition in order to arrive at the breeding sites with reserves.     

Quantifying full phenological event distributions reveals simultaneous advances,temporal stability and delays in spring and autumn migration timing in long‐distance migratory birds

Phenological changes in key seasonally expressed life‐history traits occurring across periods of climatic and environmental change can cause temporal mismatches between interacting species, and thereby impact population and community dynamics. However, studies quantifying long‐term phenological changes have commonly only measured variation occurring in spring, measured as the first or mean dates on which focal traits or events were observed. Few studies have considered seasonally paired events spanning spring and autumn or tested the key assumption that single convenient metrics accurately capture entire event distributions. We used 60 years (1955–2014) of daily bird migration census data from Fair Isle, Scotland, to comprehensively quantify the degree to which the full distributions of spring and autumn migration timing of 13 species of long‐distance migratory bird changed across a period of substantial climatic and environmental change. In most species, mean spring and autumn migration dates changed little. However, the early migration phase (≤10th percentile date) commonly got earlier, while the late migration phase (≥90th percentile date) commonly got later. Consequently, species' total migration durations typically lengthened across years. Spring and autumn migration phenologies were not consistently correlated within or between years within species and hence were not tightly coupled. Furthermore, different metrics quantifying different aspects of migration phenology within seasons were not strongly cross‐correlated, meaning that no single metric adequately described the full pattern of phenological change. These analyses therefore reveal complex patterns of simultaneous advancement, temporal stability and delay in spring and autumn migration phenologies, altering species' life‐history structures. Additionally, they demonstrate that this complexity is only revealed if multiple metrics encompassing entire seasonal event distributions, rather than single metrics, are used to quantify phenological change. Existing evidence of long‐term phenological changes detected using only one or two metrics should consequently be interpreted cautiously because divergent changes occurring simultaneously could potentially have remained undetected.     

Determining fine‐scale migratory connectivity and habitat selection for a migratory songbird by using new GPS technology

Migratory aerial insectivores are among the fastest declining avian groups, but our understanding of these trends has been limited by poor knowledge of migratory connectivity and the identification of critical habitat across the vast distances they travel annually. Using new, archival GPS loggers, we tracked individual purple martins Progne subis from breeding colonies across North America to determine precise (< > 10 m) locations of migratory and overwintering roost locations in South America and to test hypotheses for fine‐scale migratory connectivity and habitat use. We discovered weak migratory connectivity at the roost scale, and extensive, fine‐scale mixing of birds in the Amazon from distant (> 2000 km) breeding sites, with some individuals sharing the same roosting trees. Despite vast tracts of contiguous forest in this region, birds occupied a much more limited habitat, with most (56%) roosts occurring on small habitat islands that were strongly associated with water. Only 17% of these roosts were in current protected areas. These data reflect a critical advance in our ability to remotely determine precise migratory connectivity and habitat selection across vast spatial scales, enhancing our understanding of population dynamics and enabling more effective conservation of species at risk.     

Experimental dissociation of individual quality, food and timing of breeding effects on double-brooding in a migratory songbird

Annual reproductive success in many species is influenced by the number of breeding attempts within a season. Although previous studies have shown isolated effects of female quality, food, and timing of breeding on the probability of female birds producing second broods, to our knowledge, none have tested the relative importance of multiple factors and their interactions using simultaneous manipulations within populations of free-living birds. In this study, we show that individual quality and timing of breeding interact to affect the probability of double-brooding in female mountain bluebirds (Sialia currucoides). High-quality females (those that naturally initiated clutches early in the season) were more likely to double-brood, regardless of whether their hatching date was advanced or delayed, whereas later breeding, lower quality females were much less likely to double-brood when their first attempt was delayed. This indicates that annual fecundity of poorer quality (or younger) female bluebirds may be more sensitive to seasonal variation in environmental conditions. In addition, birds that were provided with supplemental food throughout first breeding attempts were more likely to double-brood in one of the study years, suggesting that female bluebirds may be energetically limited in their capacity to initiate a second brood. Females that had their first brood delayed also had a shorter inter-brood interval and were moulting fewer feathers during second broods compared to controls, while females in better condition showed more advanced moult in second breeding attempts. Taken together, our results demonstrate the combined effects of age- or individual quality-mediated energetic trade-offs between current and future reproduction, and between investments in offspring and self-maintenance, on annual fecundity of female birds.     

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.     

Recent population declines in Afro‐Palaearctic migratory birds: the influence of breeding and non‐breeding seasons

Aim

Recent, rapid population declines in many Afro‐Palaearctic migratory bird species have focussed attention on changing conditions within Africa. However, processes influencing population change can operate throughout the annual cycle and throughout migratory ranges. Here, we explore the evidence for impacts of breeding and non‐breeding conditions on population trends of British breeding birds of varying migratory status and wintering ecology.

Location

Great Britain (England & Scotland).

Methods

Within‐ and between‐species variation in population trends is quantified for 46 bird species with differing migration strategies.

Results

Between 1994 and 2007, rates of population change in Scotland and England differed significantly for 19 resident and 15 long‐distance migrant species, but were similar for 12 short‐distance migrant species. Of the six long‐distance migrant species that winter in the arid zone of Africa, five are increasing in abundance throughout Britain. In contrast, the seven species wintering in the humid zone of Africa are all declining in England, but five of these are increasing in Scotland. Consequently, populations of both arid and humid zone species are increasing significantly faster in Scotland than England, and only the English breeding populations of species wintering in the humid zone are declining.

Main conclusions

Population declines in long‐distance migrants, especially those wintering in the humid zone, but not residents or short‐distance migrants suggest an influence of non‐breeding season conditions on population trends. However, the consistently less favourable population trends in England than Scotland of long‐distance migrant and resident species strongly suggest that variation in the quality of breeding grounds is influencing recent population changes. The declines in humid zone species in England, but not Scotland, may result from poorer breeding conditions in England exacerbating the impacts of non‐breeding conditions or the costs associated with a longer migration, while better conditions in Scotland may be buffering these impacts.

     

Radio‐tags have no behavioral or physiological effects on a migratory songbird during breeding and molt

ABSTRACT Radio‐tags are commonly used in studies of avian behavior and ecology, even though many studies have revealed negative impacts of radio‐tags. Other studies of songbirds have revealed no negative effects of radio‐tags on parental care, nesting success, or annual return rates. However, such studies have not addressed the potential physiological costs of carrying radio‐tags or determined if such costs affect birds during the energetically costly molt period. Simultaneously investigating the behavioral, physiological, and fitness consequences of radio‐tags is important for evaluating possible costs. We studied Wood Thrushes (Hylocichla mustelina) in northwestern Pennsylvania and used a repeated‐measures design to determine if radio‐tags affected parental care behavior and nesting success, and measured levels of plasma lipid metabolites (β‐hydroxybutyrate, free glycerol, and triglycerides) to evaluate physiological condition. We also examined the possible physiological effects of radio‐tags during the energetically demanding period of molt in late summer. Radio‐tags did not affect male or female feeding rates, nesting success, number of nesting attempts, number of eggs laid, fledging success, or hatching success, and we found no significant effects of radio‐tag use on levels of plasma lipid metabolites of nesting females. During molt, Wood Thrushes with radio‐tags had lower levels of β‐hydroxybutyrate and higher levels of trigylcerides, suggesting they were in better, not worse, energetic condition. Our results suggest that radio‐tags do not have negative effects on the behavior or physiology of Wood Thrushes. Similar studies on a wider range of songbirds are needed before concluding radio‐tags have little or no impact during the molting period.     

Finish with a sprint: Evidence for time‐selected last leg of migration in a long‐distance migratory songbird

Under time‐selected migration, birds should choose a strategy for outcompeting rivals over securing access to prime resources at the final destination. Thus, migration can be viewed as a race among individuals where winners are arriving first when conditions are suitable. The sprint migration hypothesis predicts that individuals shift from maximum sustained speed to a final burst of sprint to shorten the transition from migration to breeding (Alerstam, 2006). In this study, we test the hypothesis of a final sprint migration in a long‐distance Afro‐Palearctic migrant, the collared flycatcher Ficedula albicollis, during autumn and spring, and compare migration strategies between the seasons. In both seasons, collared flycatchers evidently exhibited sprint migration by increasing their overall speed over the last leg of migration after the Sahara crossing. This phenomenon was more pronounced in spring, contributing to overall faster spring migration and possibly highlighting higher importance for early arrival at the breeding grounds. In both seasons and particularly in spring, late departing individuals flew at a faster rate, partially being able to catch up with their early departing conspecifics. Differential fueling strategies may play an important role in determining migration speed, especially during the early stages of the migration, and might explain the observed differences in migration speeds between late and early departing individuals. Our findings suggest competition for early arrival at the breeding and at the nonbreeding destinations alike. Sprint migration might be an appropriate strategy to gain advantage over conspecifics and settle in prime territories as well as to cope with the increasingly earlier springs at high latitudes.     

Androgen‐metabolizing enzymes show region‐specific changes across the breeding season in the brain of a wild songbird

The Lapland longspur (Calcarius lapponicus) is an arctic‐breeding songbird that shows rapid behavioral changes during a short breeding season. Changes in plasma testosterone (T) in the spring are correlated with singing but not territorial aggression in males. Also, T treatment increases song but not aggression in this species. In contrast, in temperate‐zone breeders, song and aggression are highly correlated, and both increase after T treatment. We asked whether regional or temporal differences in androgen‐metabolizing enzymes in the longspur brain explain hormone‐behavior patterns in this species. We measured the activities of aromatase, 5α‐reductase and 5β‐reductase in free‐living longspur males. Aromatase and 5α‐reductase convert T into the active steroids 17β‐estradiol (E₂) and 5α‐dihydrotestosterone (5α‐DHT), respectively. 5β‐Reductase deactivates T via conversion to 5β‐DHT, an inactive steroid. We examined seven brain regions at three stages in the breeding season. Overall, aromatase activity was high in the hypothalamus, hippocampus, and ventromedial telencephalon (containing nucleus taeniae, the avian homologue to the amygdala). 5β‐Reductase activity was high throughout the telencephalon. Activities of all three enzymes changed over time in a region‐specific manner. In particular, aromatase activity in the rostral hypothalamus was decreased late in the breeding season, which may explain why T treatment at this time does not increase aggression. Changes in 5β‐reductase do not explain the effects of plasma T on aggressive behavior. © 1999 John Wiley & Sons, Inc. J Neurobiol 41: 176–188, 1999     

Territorial occupancy and breeding performance in a migratory raptor do not follow ideal despotic distribution patterns

It has commonly been argued that many territorial species select their breeding sites following an ideal despotic distribution model, in which the most productive, high-quality territories are more frequently occupied. Theoretical and empirical studies have shown that this occupancy pattern may have population regulatory consequences, leading to density dependence in heterogeneous habitats. During a 9-year research project in a forested area of south-eastern Spain, we tested some of the predictions of the ideal despotic distribution model and the site-dependent population regulation theory in a migratory raptor species, the booted eagle Hieraaetus pennatus . Contrary to the predictions of the despotic model, our results showed that the temporal pattern of territorial occupation did not differ from randomness, and that the territory occupancy rate was not significantly related to the reproductive parameters considered. At population level, the breeding variables were density independent, suggesting the absence of site-dependent regulation. In addition, we were unable to find significant differences in the habitat characteristics between high-quality and low-quality territories, classified according to the criteria of both occupancy frequency and average productivity. Overall, our results suggest that booted eagles select their territories at random, probably due to the lack of strong environmental heterogeneity, and that occupancy rate is not a good measure of territory quality for the population studied.     

Mapping migration in a songbird using high‐resolution genetic markers

Neotropic migratory birds are declining across the Western Hemisphere, but conservation efforts have been hampered by the inability to assess where migrants are most limited—the breeding grounds, migratory stopover sites or wintering areas. A major challenge has been the lack of an efficient, reliable and broadly applicable method for measuring the strength of migratory connections between populations across the annual cycle. Here, we show how high‐resolution genetic markers can be used to identify genetically distinct groups of a migratory bird, the Wilson's warbler (Cardellina pusilla), at fine enough spatial scales to facilitate assessing regional drivers of demographic trends. By screening 1626 samples using 96 highly divergent single nucleotide polymorphisms selected from a large pool of candidates (~450 000), we identify novel region‐specific migratory routes and timetables of migration along the Pacific Flyway. Our results illustrate that high‐resolution genetic markers are more reliable, precise and amenable to high throughput screening than previously described intrinsic marking techniques, making them broadly applicable to large‐scale monitoring and conservation of migratory organisms.     

Short‐ and long‐term costs of reproduction in a migratory songbird

Costs of reproduction represent a common life‐history trade‐off. Critical to understanding these costs in migratory species is the ability to track individuals across successive stages of the annual cycle. We assessed the effects of total number of offspring fledged and date of breeding completion on pre‐migratory body condition, the schedule of moult and annual survival in a migratory songbird, the Savannah Sparrow Passerculus sandwichensis. Between 2008 and 2010, moult was delayed for individuals that finished breeding later in the breeding period and resulted in reduced lean tissue mass during the pre‐migratory period, suggesting an indirect trade‐off between the timing of breeding completion and condition just prior to migration. Lean tissue mass decreased as the number of offspring fledged increased in 2009, a particularly cool and wet year, illustrating a direct trade‐off between reproductive effort and condition just prior to migration in years when weather is poor. However, using a 17‐year dataset from the same population, we found that parents that fledged young late in the breeding period had the highest survival and that number of offspring fledged did not affect survival, suggesting that individuals do not experience long‐term trade‐offs between reproduction and survival. Taken together, our results suggest that adult Savannah Sparrows pay short‐term costs of reproduction, but that longer‐term costs are mitigated by individual quality, perhaps through individual variation in resource acquisition.     

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.     

Patterns and correlates of songbird movements at an ecological barrier during autumn migration assessed using landscape‐ and regional‐scale automated radiotelemetry

Departure decisions of songbirds at ecological barriers they encounter en route can strongly influence time, energy and survival costs of migration. To date, most field studies of departure decisions and their correlates have used indirect methods and followed migrants at a single stopover site, with limited information on what happens to individuals after they depart from the site. We used an automated radiotelemetry array extending 350 km from southwest Nova Scotia to southern Maine to study the migratory and stopover movements of Northern Waterthrushes Parkesia noveboracensis, Red‐eyed Vireos Vireo olivaceus and Yellow‐rumped Warblers Setophaga coronata in relation to fuel load and weather at the northeastern edge of the Gulf of Maine. From the 105 radio‐transmitters we deployed in southwest Nova Scotia, we recorded 42 landscape‐scale stopover flights and 47 migratory flights by 75 individuals. Of the migratory flights, 57% were orientated southwest, a trajectory that, if held, would require individuals to complete a 350–440 km overwater flight. The remaining 43% of migratory flights were orientated northwest, away from the Gulf of Maine, and 15 individuals were confirmed to have detoured around all or a portion of the barrier, as evidenced by their being re‐detected over the Bay of Fundy and/or along the coast of Maine between 4 h and 15 days later. Across all individuals, initial fat score had a positive effect on departure probability, especially for individuals that made stopover flights. Among weather variables, tailwind assistance was the best predictor of migratory departures but did not appear to be the main factor determining whether individuals orientated towards or away from the Gulf of Maine. Weather had little effect on departure decisions of individuals that made stopover flights. These differences in the correlates of migratory departures and stopover flights would probably not have been distinguishable had our study been restricted to a local scale. Therefore, our findings highlight the importance of expanding the scale at which departure decisions and the ecology of stopover in general are studied.     

Potential age differences in the migratory behaviour of a nocturnal songbird migrant during autumn and spring

In migratory songbirds, older individuals are thought to be more efficient migrants than younger individuals. Age‐specific differences in migratory efficiency have been reported mainly in respect of arrival timing, energy stores, rate of energy accumulation, departure behaviour, and departure direction. Yet, these traits were rarely assessed simultaneously in a single species. We further lack information whether age‐specific differences in behavioural traits present in autumn still manifest to the same degree in spring. Here we used the northern wheatear Oenanthe oenanthe, a long‐distance nocturnal songbird migrant, and investigated age‐specific differences in energy stores at capture (autumn: 1059 birds/spring: 803 birds), rate of energy accumulation (168/147), nocturnal departure timing (126/105), and departure direction (94/77) for both seasons. We found that in autumn, young northern wheatears departed significantly later in the night than older birds. This difference was not observed in spring. The resulting advance in nocturnal departure timing by young birds from autumn to spring may be due to learning based on experience gained during autumn, and/or to selective disappearance of those young individuals showing late departure times during autumn. We found no age‐specific difference in any of the other migratory traits investigated. To get a better understanding of age effects in songbird migrants, we need to study the potential adjustments of migratory behaviour within the individual over its life time. By individually tracking songbirds throughout their lifetime, we could start estimating whether a certain migratory decision (fuelling, departure, orientation) translates into higher (or lower) fitness, and whether individuals adjust their migratory behaviour based on learning from ‘wrong decisions’.