Long-distance Migratory Songbird Research Articles

Asynchronous movement patterns between breeding and stopover locations in a long-distance migratory songbird

Asynchronous movement patterns between breeding and stopover locations in a long-distance migratory songbird

Spring departure date, not en route conditions, drive migration rate and arrival timing in a long-distance migratory songbird

Long-distance avian migrants are threatened by a rapidly changing environment, which can induce a mismatch between favorable environmental conditions and the timing of breeding. Therefore, migratory birds must rely on phenotypic plasticity within migration timing to remain synchronized with earlier occurring resources over the shorter term. However, how this plasticity responds to en route environmental factors and whether or not it may allow migratory birds to remain synchronized with advancing springs is largely unknown. The objective of this study was to investigate how extrinsic factors, such as those related to the weather (e.g., temperature, precipitation, and wind speed) and daylength experienced along spring migration routes, and intrinsic factors, such as sex and age, migration departure date from wintering grounds, and migration destination (i.e., breeding site), influence the migration rate and spring arrival date at breeding grounds in purple martins (Progne subis). We investigated arrival timing and rate using 307 individual migration tracks sampled across the birds’ eastern breeding range. We found that the departure date and total stopover time explained most of the variation in migration rate, with birds departing later traveling faster but taking more stopover days, leading to a slower overall migration rate. To a lesser extent, facilitating southern winds and lower precipitation also induced a faster migration rate. We found that birds with a faster migration rate arrived earlier at their breeding sites. We also found that female birds traveling to more northern breeding sites arrived earlier than male birds, and that they migrated at a faster rate. Overall, our results suggest that the migration rate may be most impacted by intrinsic factors tied to spring departure dates that are aligned with the phenology of breeding sites rather than by conditions experienced en route. These findings support growing evidence that selection for spring departure date may be needed for the advancement of spring arrival date to match advancing springs with climate change. Future research investigating how environmental conditions at stopovers contribute to the overall migration rate would provide further insight into the factors influencing migration timing.

Long-duration wind tunnel flights reveal exponential declines in protein catabolism over time in short- and long-distance migratory warblers

During migration, long-distance migratory songbirds may fly nonstop for days, whereas shorter-distance migrants complete flights of 6 to 10 h. Fat is the primary fuel source, but protein is also assumed to provide a low, consistent amount of energy for flight. However, little is known about how the use of these fuel sources differs among bird species and in response to flight duration. Current models predict that birds can fly until fat stores are exhausted, with little consideration of protein's limits on flight range or duration. We captured two related migratory species-ultra long-distance blackpoll warblers (Setophaga striata) and short-distance yellow-rumped warblers (Setophaga coronata)-during fall migration and flew them in a wind tunnel to examine differences in energy expenditure, overall fuel use, and fuel mixture. We measured fat and fat-free body mass before and after flight using quantitative magnetic resonance and calculated energy expenditure from body composition changes and doubly labeled water. Three blackpolls flew voluntarily for up to 28 h-the longest wind tunnel flight to date-and ended flights with substantial fat reserves but concave flight muscle, indicating that protein loss, rather than fat, may actually limit flight duration. Interestingly, while blackpolls had significantly lower mass-specific metabolic power in flight than that of yellow-rumped warblers and fuel use was remarkably similar in both species, with consistent fat use but exceptionally high rates of protein loss at the start of flight that declined exponentially over time. This suggests that protein may be a critical, dynamic, and often overlooked fuel for long-distance migratory birds.

Genomic architecture of migration timing in a long-distance migratory songbird

The impact of climate change on spring phenology poses risks to migratory birds, as migration timing is controlled predominantly by endogenous mechanisms. Despite recent advances in our understanding of the underlying genetic basis of migration timing, the ways that migration timing phenotypes in wild individuals may map to specific genomic regions requires further investigation. We examined the genetic architecture of migration timing in a long-distance migratory songbird (purple martin, Progne subis subis) by integrating genomic data with an extensive dataset of direct migratory tracks. A moderate to large amount of variance in spring migration arrival timing was explained by genomics (proportion of phenotypic variation explained by genomics = 0.74; polygenic score R2 = 0.24). On chromosome 1, a region that was differentiated between migration timing phenotypes contained genes that could facilitate nocturnal flights and act as epigenetic modifiers. Overall, these results advance our understanding of the genomic underpinnings of migration timing.

Departure, routing and landing decisions of long-distance migratory songbirds in relation to weather.

Migrating birds flexibly adjust their individual migratory decisions, i.e. departing, routing and landing, based on intrinsic (e.g. energy stores) and extrinsic (e.g. landscape features and weather) factors modulating the endogenous stimuli. So far, these decisions have mostly been studied separately. Notably, we lack information on which factors landing decisions during active flight are based on. Therefore, we simultaneously recorded all three decisions in free-flying long-distance migratory songbirds in a coastal stopover area via regional-scale radio-telemetry and related them to the prevailing weather. Birds departed under favourable weather conditions resulting in specific nights with increased departure probability. Once departed, birds could either fly offshore or take a route along the coast, which was predicted by wind support. Radio-tracking revealed that departed individuals more likely interrupted their migratory endurance flight under overcast or headwind conditions. Studying departure, routing and landing decisions in concert, we highlight the importance of weather as a common driver across all migratory decisions. By radio-tracking individuals between stopovers, we provide evidence that avoidance of adverse weather conditions is an important function of stopover. Understanding how birds adjust migratory decisions and how they affect the timing of migration and survival is key to link migration performance to individual fitness.

The structure of songbird MHC class I reveals antigen binding that is flexible at the N-terminus and static at the C-terminus.

Long-distance migratory animals such as birds and bats have evolved to withstand selection imposed by pathogens across the globe, and pathogen richness is known to be particularly high in tropical regions. Immune genes, so-called Major Histocompatibility Complex (MHC) genes, are highly duplicated in songbirds compared to other vertebrates, and this high MHC diversity has been hypothesised to result in a unique adaptive immunity. To understand the rationale behind the evolution of the high MHC genetic diversity in songbirds, we determined the structural properties of an MHC class I protein, Acar3, from a long-distance migratory songbird, the great reed warbler Acrocephalus arundinaceus (in short: Acar). The structure of Acar3 was studied in complex with pathogen-derived antigens and shows an overall antigen presentation similar to human MHC class I. However, the peptides bound to Acar3 display an unusual conformation: Whereas the N-terminal ends of the peptides display enhanced flexibility, the conformation of their C-terminal halves is rather static. This uncommon peptide-binding mode in Acar3 is facilitated by a central Arg residue within the peptide-binding groove that fixes the backbone of the peptide at its central position, and potentially permits successful interactions between MHC class I and innate immune receptors. Our study highlights the importance of investigating the immune system of wild animals, such as birds and bats, to uncover unique immune mechanisms which may neither exist in humans nor in model organisms.

Activity patterns throughout the annual cycle in a long-distance migratory songbird, the red-backed shrike Lanius collurio

BackgroundLong-distance migratory birds undergo complex annual cycles during which they must adjust their behaviour according to the needs and conditions encountered throughout the year. Yet, variation in activity throughout the entire annual cycle has rarely been studied in wild migratory birds.MethodsWe used multisensor data loggers to evaluate the patterns of activity throughout the complete annual cycle of a long-distance migratory bird, the red-backed shrike Lanius collurio. Accelerometer data was used to identify life-history stages and to estimate levels of activity during various phases of the annual cycle. In this study, we analysed the variation in daytime activity along the annual cycle and between migratory and non-migratory days.ResultsThe birds’ daytime activity varied throughout the annual cycle while night-time activity was almost exclusively restricted to migratory flights. The highest daytime activity levels were observed during the breeding season, while it remained low during autumn migration and the winter period. Daytime activity differed between sexes during the breeding period, when the males showed the highest level in activity. During migratory periods, both sexes exhibited a higher daytime activity in spring compared to autumn migration, being particularly high in the final migratory leg towards the breeding ground. The birds showed a lower daytime activity on migratory days (days when a migratory flight took place during the succeeding night) than on non-migratory days during both migratory seasons.ConclusionsActivity measured during daytime results from a combination of several behaviours, and a high daytime activity during spring migration and the breeding period is possibly reflecting particularly energy-demanding periods in the annual cycle of migratory birds. The use of multisensor data loggers to track annual activity provides us with a full annual perspective on variation in activity in long-distance migratory species, an essential approach for understanding possible critical life-history stages and migration ecology.

Variation in body size and plumage does not explain apparent survival for a long-distance migratory songbird, the Bobolink (Dolichonyx oryzivorus)

El tordo charlatán (Dolichonyx oryzivorus) es un ave canora de pastizal migratoria de larga distancia cuya población global está en declive a largo plazo, principalmente debido a la pérdida de hábitat y a la intensificación de las prácticas agrícolas. Para entender mejor los factores que afectan su ciclo anual, utilizamos una población con edad conocida de tordos charlatanes machos (n = 121) que se reproducían en pastizales agrícolas de Vermont y exploramos cómo la variación en su plumaje (la extensión del amarillo de su capucha) y la morfología corporal (masa corporal y longitud del ala) explicaban la variación en la sobrevivencia aparente. La masa corporal del tordo charlatán y la longitud del ala variaron entre los años 2–5. El tamaño de la capucha del tordo charlatán no tuvo variaciones con la edad. Sin embargo, ninguno de los 3 caracteres morfológicos explicaba la variación de sobrevivencia aparente. Nuestros resultados resaltan los desafíos asociados a entender la variación en la calidad de los individuos relativa a la edad, particularmente con respecto a factores que afectan procesos demográficos de especies en declive. Palabras clave: ala, Icteridae, masa, morfología, plumaje, Programa MARK, Vermont.

Arrival condition and refueling performance of long-distance migratory songbirds at a spring stopover site in the eastern Gulf of Mexico

Barrier islands along the northern Gulf coast provide critical stopover habitats for Nearctic–Neotropical migratory songbirds using a spring trans-Gulf migration strategy. Islands bordering Apalachicola Bay, Florida, project southward into the Gulf of Mexico providing an opportunity for first landfall. In 2013 and 2014, we studied spring stopover ecology on St. George Island, Florida. We provide a detailed description of the arrival body condition and fuel deposition rates of 8 species of migratory birds. We used 3 approaches to measure fuel deposition rate: (1) estimated an hourly rate of mass change by regressing body mass or scaled body mass by time of day, (2) estimated a daily rate of mass change by examining body mass of recaptured individuals, and (3) for 4 species, measured plasma triglyceride concentrations as a proxy of individual refueling rate. The majority of migratory birds in the eastern Gulf of Mexico arrived in poor body condition with depleted fat and muscle stores. We found no significant changes in body mass or scaled body mass as the day progressed, except for Indigo Bunting (Passerina cyanea), which gained mass throughout the day. Recapture rate was low on St. George Island (<5%), but most recaptured individuals tended to lose mass. Using plasma triglyceride concentration, we found that refueling rate for Gray Catbird (Dumetella carolinensis) declined throughout the day. We found no relationship between body condition and refueling rate in Gray Catbird or Catharus thrushes. Our results suggests that long-distance migratory birds arriving on St. George Island after a nonstop migratory flight may be physiologically challenged due to poor body condition. Our results provide the first quantitative assessment of songbird refueling performance on Florida panhandle barrier islands and reinforce the importance of Gulf of Mexico barrier islands as critical stopover sites for long-distance migrants in poor body condition.

Миграция североазиатских воробьиных

The East Asian-Australasian bird migration system is one of the most species-rich migration systems, nevertheless, we have very little information on the migration of the species that use the Asian-Australasian Flyway. Most knowledge is available about waterfowls (cranes, ducks). However, very little is known about songbirds, mainly due to the lack of large-scale, long-term ringing activities. Most of what we know about the migration of these species is based primarily on field observations and the results of the Migratory Animal Pathological Survey (MAPS) conducted in the 1960s and 1970s. In the 2010s, however, several local ringing projects started. They produced considerable knowledge about the migration of songbirds. More recently, geolocators have also aided researchers in their work, providing even more accurate data on the migratory routes and migratory habits of species. The present study summarises the data we have obtained over the past decade about the migration of long-distance migratory songbirds nesting in North Asia. The article is based primarily on the data collected at ringing stations in the Far Eastern Russia, complemented by research from other areas in East and Southeast Asia. This review highlights the need for further research to ensure long-term protection of species that, at times, show a drastic decline in numbers.

The Influence of Different Light Wavelengths of Anthropogenic Light at Night on Nestling Development and the Timing of Post-fledge Movements in a Migratory Songbird

Many different aspects of an animal’s lifecycle such as its behavior, patterns of hormone activity, and internal clock time, can be affected by anthropogenic light at night (ALAN). Exposing an organism to ALAN during its early life could also have an impact on its development. Since photoperiod can trigger or schedule the migration timing of long-distance migratory birds, there is great potential for anthropogenic light to interact with photoperiod to affect timing. However, very little has been investigated regarding the impacts of ALAN on post-hatching development and migration timing. We investigated the impact of ALAN during nestling development in a long-distance migratory songbird to determine the potential impact on the timing of post-breeding movements in the wild. We experimentally manipulated the light by using programmable lighting, in the nest boxes of free-living nestlings of purple martin (Progne subis) in Manitoba, Canada. We exposed two groups of developing nestlings, from hatch to fledge date, to green or white LED lights (5 lux) during the night. We also included a control group that experienced natural, ambient light at night. We found that some adults abandoned their nests shortly after starting the experiment (4 of 15 nests in the white light treatment). For the nests that remained active, nestlings exposed to the white light treatment had higher weights (at day 20 or 22), later fledge dates (1.54 ± 0.37, 95% CI 0.80–2.28), and later colony departure date (2.84 ± 1.00, 95% CI 0.88–4.81), than young of the control group. Moreover, nestlings of both white and green light groups had longer nesting duration than nestlings of the control group. This study demonstrates the impact of ALAN on the development of post-breeding movement timing in nestlings of wild migratory birds. However, our results also indicate that green light may have less of an impact as compared to white light.

When to depart from a stopover site? Time since arrival matters more than current weather conditions

AbstractOn the journey to wintering sites, most migratory birds alternate between flights and stopovers, where they rest and refuel. In contrast to the time-minimization strategy commonly assumed to drive the pre-breeding migration, birds are rather expected to follow an energy minimization during post-breeding migration. It is the cumulative duration of flights and stopovers that determines the total energy requirements and duration of the journey. Since migrating birds actually spend most of the time at stopovers sites, selection to minimize the amount of energy or time spent on migration is likely to operate on the effectiveness of stopover rest and refueling. Here, we address the relative contribution of factors acting on departure decisions from a stopover site during the post-breeding migration in a long-distance migratory songbird. When capture probability is low, it is impossible to measure fattening over the entire duration of the stopover. To get around this limitation, we use time since arrival (TSA) as a proxy for the progressive temporal change occurring in the internal state of an individual (i.e. rest, physiological recovery, and fuel loading) during the stopover. We develop a capture–recapture model to address the respective effects of estimated TSA and of weather conditions on departure probability. Using a 20-year dataset for Sedge Warblers (Acrocephalus schoenobaenus), we show that TSA served as a surrogate of the most important information that birds use when deciding to depart from a stopover site, while low humidity and rising atmospheric pressure only slightly increase daily departure probability. Hence, a bird would resume migration mainly according to the time it had to rest and refuel, and then fine-tuning departure decision according to weather conditions. The generality of these results needs to be assessed by applying this modeling framework to other migratory species and at sites or times with greater weather variability.

Experimental manipulation of photoperiod influences migration timing in a wild, long-distance migratory songbird.

Previous laboratory studies have demonstrated the role of photoperiod in cueing the migration timing of small land birds; however, how migration timing of young birds in wild environments develops in relation to these cues have rarely been investigated. Such investigations can make important contributions to our developing understanding of the phenotypic plasticity of migration timing to new conditions with climate change, where changes in the timing of nesting may expose juvenile birds to different photoperiods. We investigated the impact of manipulating photoperiod during nestling development in a long-distance migratory songbird on the timing of post-breeding movements in the wild. Using programmable lighting installed in the nest-boxes of purple martins (Progne subis), we exposed developing nestlings, from hatch to fledge date, to an extended photoperiod that matched the day length of the summer solstice in Manitoba, Canada. We found that birds with a simulated, earlier photoperiod had a longer nesting period and later fledge and autumn departure dates than control group birds. This study demonstrates the phenotypic plasticity of first-year birds to the ontogenetic effect of their hatch date in the formation of the timing of their first post-breeding movements. Further, we discuss how these results have implications for the potential use of assisted evolution approaches to alter migration timing to match new conditions with climate change.

Breeding latitude is associated with the timing of nesting and migration around the annual calendar among Purple Martin (Progne subis) populations

Timing is critical for species that migrate seasonally, as timing mismatches can lead to reduced reproductive success and population declines. We investigated the effect of breeding latitude on the migration timing of a long-distance migratory songbird (Purple Martin, Progne subis subis) by using light-level geolocators deployed across breeding sites between 26° and 53° latitude (n = 329). We used nesting data (1st egg dates; n = 28,165) to explore latitudinal patterns in the timing of breeding. Using generalized linear mixed models (GLMMs) we found that breeding latitude was highly associated with timing around the calendar, explaining population-level timing across both spring and fall migration. Populations breeding at more northern latitudes migrated later in fall (mean days/degree latitude = 2), in spring (mean days/degree latitude = 3), and nested later (days/degree latitude = 3) than populations breeding at more southern latitudes. Generalized additive mixed models (GAMMs) revealed significant non-linearity between breeding latitude and spring and fall migration timing, and the timing of nesting. In spring, the most northern breeders (> 40 degrees N) were more synchronous in their timing, departing their nonbreeding sites and arriving earlier at their breeding sites relative to their breeding latitude. These most northern breeders also had similar nest timing and fall migration departure timing. Breeding latitude was also associated with seasonal partitioning of migratory and sedentary periods. The most northern breeders spent 10% more time on fall migration, 2% more at wintering grounds, 1% more on spring migration, and 13% less time at their breeding sites. Our results support the notion that selection for timing at breeding latitudes can have year-round effects on the timing of seasonal events in long-distance migratory songbirds. Future studies should examine whether these timing patterns associated with latitude limit phenotypic responses to climate change.

Habitat loss on the breeding grounds is a major contributor to population declines in a long-distance migratory songbird.

Many migratory species are declining and for most, the proximate causes of their declines remain unknown. For many long-distance Neotropical migratory songbirds, it is assumed that habitat loss on breeding or non-breeding grounds is a primary driver of population declines. We integrated data collected from tracking technology, community science and remote sensing data to quantify migratory connectivity (MC), population trends and habitat loss. We quantified the correlation between forest change throughout the annual cycle and population declines of a long-distance migratory songbird, the Connecticut warbler (Oporornis agilis, observed decline: -8.99% yr-1). MC, the geographic link between populations during two or more phases of the annual cycle, was stronger between breeding and autumn migration routes (MC = 0.24 ± 0.23) than between breeding and non-breeding locations (MC = -0.2 ± 0.14). Different Connecticut warbler populations tended to have population-specific fall migration routes but overlapped almost completely within the northern Gran Chaco ecoregion in South America. Cumulative forest loss within 50 km of breeding locations and the resulting decline in the largest forested patch index was correlated more strongly with population declines than forest loss on migratory stopover regions or on wintering locations in South America, suggesting that habitat loss during the breeding season is a driver of observed population declines for the Connecticut warbler. Land-use practices that retain large, forested patches within landscapes will likely benefit breeding populations of this declining songbird, but further research is needed to help inform land-use practices across the full annual cycle to minimize the impacts to migratory songbirds and abate ongoing population declines.

Weak evidence of carry-over effects of overwinter climate and habitat productivity on spring passage of migratory songbirds at a northern stopover site in Ontario

Abstract Reduced rainfall in tropical regions decreases the availability of food resources for overwintering songbirds, with negative impacts on their body condition. Increasingly dry conditions at tropical wintering sites as a result of climate change may impact the ability of temperate-breeding songbirds to prepare for and execute their spring migration. We studied the carry-over effects of natural climatic fluctuations created by the El Niño–Southern Oscillation (ENSO) in tropical overwintering areas on 7 long-distance migratory songbirds at a Canadian stopover site. We used the Normalized Difference Vegetation Index as a proxy for tropical habitat productivity and resource availability and a 34-year bird banding dataset from Long Point, Ontario, Canada to assess migration timing and stopover body condition. To increase our ability to detect potential carry-over effects, we employed a novel approach of using recent migratory connectivity studies to restrict the wintering ranges to areas most likely used by individuals passing through southern Ontario. Using linear models, we found that overwinter habitat productivity was significantly negatively influenced by dry ENSO events in the overwintering ranges in 3 of 7 species, with a fourth near-significant. Subregional differences in the effect of ENSO on precipitation patterns may explain why we did not detect a consistent effect of ENSO on overwinter habitat productivity for all species. Despite restricting the wintering range and using a robust dataset for species with diverse life histories, we detected only weak and often conflicting evidence of population-level carry-over effects from dry ENSO events and overwinter habitat productivity. Negative carry-over effects may be strongest and most evident during the earlier stages of migration because birds may be able to compensate to some extent for poor departure condition and late departure while en route.

Warmer April Temperatures on Breeding Grounds Promote Earlier Nesting in a Long-Distance Migratory Bird, the Prothonotary Warbler

Global climate change and warming are altering hemispheric and local weather patterns. Altered weather patterns have great potential to affect the phenology of life history events, such as the initiation of breeding in organisms that reproduce seasonally. Long-distance migratory birds may be particularly challenged by changes in local weather on breeding grounds because they arrive from distant locations and must commence breeding when conditions are appropriate. Here we explore the effects of local temperature on first egg dates and annual productivity in a long-distance Neotropical migratory songbird, the prothonotary warblerProtonotaria citrea. We present results from a 20-year (1994 to 2013) study documenting the detailed nesting activities of a color-marked population (average of 155 individual females each year) of warblers in southern Illinois, United States. The warblers typically arrive in April and start breeding in late April and May in our study system. We tested for an effect of local average April daily temperature and female age on first egg dates, total number of offspring produced per female, and the probability of fledging two broods. We found that warmer April temperatures promoted earlier first egg dates and higher average annual productivity in the warblers. On average, older females had earlier first egg dates than 1-year-old females, but both age groups responded similarly to local April temperatures. The reproductive gains associated with earlier first egg dates in warmer years stemmed from an increased probability of successfully fledging two broods, suggesting that earlier first egg dates do not currently create a mismatch with food (insect) resources. Earliest arrival dates of warblers to the region of our study system were not affected by local April temperatures, suggesting that females vary their first egg date based on conditions they experience/assess after their arrival. Whereas these birds currently adjust the timing of their breeding and actually produce more offspring in warmer years, continued global warming may eventually upset the current balance between arrival dates, food resources, and the commencement of nesting.

Pre-migration artificial light at night advances the spring migration timing of a trans-hemispheric migratory songbird.

Artificial light at night (ALAN) is increasing at a high rate across the globe and can cause shifts in animal phenology due to the alteration of perceived photoperiod. Birds in particular may be highly impacted due to their use of extra-retinal photoreceptors, as well as the use of photoperiodic cues to time life events such as reproduction, moult, and migration. For the first time, we used light-logging geolocators to determine the amount of ALAN experienced by long-distance migratory songbirds (purple martin; Progne subis) while at their overwintering sites in South America to measure its potential relationship with spring migration timing. Almost a third of birds (48/155; 31%) were subjected to at least one night with ALAN over 30 days prior to spring migration. Birds that experienced the highest number of nights (10+) with artificial light departed for spring migration on average 8 days earlier and arrived 8 days earlier at their breeding sites compared to those that experienced no artificial light. Early spring migration timing due to pre-migration ALAN experienced at overwintering sites could lead to mistiming with environmental conditions and insect abundance on the migratory route and at breeding sites, potentially impacting survival and/or reproductive success. Such effects would be particularly detrimental to species already exhibiting steep population declines such as purple martins and other migratory aerial insectivores.

Population-specific assessment of carry-over effects across the range of a migratory songbird

Annual cycle events may be interlinked, influence following annual cycle stages, and may alter performance of individuals. Such links, called carry-over effects, can explain individual variation in timing or reproductive success in migratory species. Identifying the key links affecting fitness may reveal the mechanisms of species population dynamics but the current evidence for the strongest carry-over effects is equivocal. Here, we aim to assess the carry-over effects in great reed warblers Acrocephalus arundinaceus, a long-distance migratory songbird, using 103 full-annual tracks from three European and two Asian breeding populations. Our results showed strong positive relationships within autumn and spring migration periods and buffering capacity of the non-breeding period preventing events to carry over between these periods. Moreover, we found no profound relation between the non-breeding habitat quality or seasonality (quantified using stable isotopes and remote sensing data) and the timing of spring migration. The strongest carry-over effects occurred in individuals from the southern European breeding population compared to the northern and the central European populations. A moderate relationship between the habitat seasonality during moult and the spring migration timing indicates the importance of the complete moult. The overall weak carry-over effects of non-breeding habitat conditions found in this study contrast with previous results and imply between-species differences in these crucial relationships. Moreover, the population-specific carry-over effects highlight the importance of multi-population approach and advise caution in interpretation of results from single-population studies. Finally, the carry-over effect from the moulting period indicates the significance of a so-far neglected link in the species. Environmental conditions vary in space and time. Therefore, migratory species adjust the timing of migration in order to maximise their fitness. However, the links between annual cycle events in multiple populations and the consequences of environmental conditions outside the breeding range are scarcely known. In this study, we used tracking data of the great reed warbler, an insectivorous bird species breeding across western Eurasia and wintering in Africa, to study a complex system of links between annual events. We found that the strength of these links differed between geographically distinct populations but not between sexes. Moreover, harsh environmental conditions during moult delayed the timing of subsequent events. Our findings could help explain large-scale differences in population size changes observed in some species and highlight the importance of energetically demanding moult period for the life of migratory species. Finally, our results demonstrate the need for multi-population approach in studies on seasonal interactions.

Earlier and slower or later and faster: Spring migration pace linked to departure time in a Neotropical migrant songbird.

Migratory birds travel vast distances and the timing of migratory flights can affect survival and the ability to reproduce. For Neotropical migrant songbirds, early spring departure from wintering sites, early arrival to the breeding grounds and higher reproductive success have been related to the use of suitable habitats and environmental conditions during the non-breeding season. However, how migratory strategies are shaped by winter habitat choice is largely unknown due to the general inability to track birds from specific wintering habitats to stopovers or breeding destinations. We assessed how winter habitat (native forest vs. shade-grown coffee plantations) relates to spring departure date and migration pace in Swainson's Thrush Catharus ustulatus. We also determined the effect of departure date and total migration duration on the arrival date of birds detected near or within their breeding range. We used a novel application of Motus radiotelemetry arrays to track individuals from their wintering grounds in the Andes of South America along their migratory journey to North America. We found variation in migratory strategies between habitats, with birds wintering in native forest departing later than birds in coffee. We present isotopic evidence for native forest being of higher quality than shade-coffee for Swainson's Thrush and hypothesize that moister conditions in forest, as shown by stable isotope (δ13 C) analysis of thrush whole blood, provides favourable pre-migratory conditions allowing birds to delay departure from wintering grounds. Habitat, between-site and -year variation in departure date, suggests that birds made facultative adjustments to winter habitat quality and environmental conditions. Independent of habitat, birds that departed later migrated faster and this pattern was maintained along the migration route (n=44). Migrating earlier and slower or later and faster was unlikely to result in significant differences in arrival time to breeding destinations. Our findings reveal underappreciated complexity in migratory decisions by long-distance migrants that contrast with the current paradigm of earlier departures and arrival from optimal habitats. The next step is to understand the relative fitness benefits of early versus late schedules or whether each strategy is an equally good response to experienced conditions.