A refined magnetic pulse treatment method for magnetic navigation experiments with adequate sham control: a case study on free-flying songbirds.

Migratory songbirds may navigate by extracting positional information from the geomagnetic field, potentially with a magnetic-particle-based receptor. Previous studies assessed this hypothesis experimentally by exposing birds to a strong but brief magnetic pulse aimed at remagnetizing the particles and evoking an altered behaviour. Critically, such studies were not ideally designed because they lacked an adequate sham treatment controlling for the induced electric field that is fundamentally associated with a magnetic pulse. Consequently, we designed a sham-controlled magnetic-pulse experiment, with sham and treatment pulse producing a similar induced electric field, while limiting the sham magnetic field to a value that is deemed insufficient to remagnetize particles. We tested this novel approach by pulsing more than 250 wild, migrating European robins (Erithacus rubecula) during two autumn seasons. After pulsing them, five traits of free-flight migratory behaviour were observed, but no effect of the pulse could be found. Notably, one of the traits, the migratory motivation of adults, was significantly affected in only one of the two study years. Considering the problem of reproducing experiments with wild animals, we recommend a multi-year approach encompassing large sample size, blinded design and built-in sham control to obtain future insights into the role of magnetic-particle-based magnetoreception in bird navigation.

A conceptual framework on the role of magnetic cues in songbird migration ecology.

Migrating animals perform astonishing seasonal movements by orienting and navigating over thousands of kilometres with great precision. Many migratory species use cues from the sun, stars, landmarks, olfaction and the Earth's magnetic field for this task. Among vertebrates, songbirds are the most studied taxon in magnetic-cue-related research. Despite multiple studies, we still lack a clear understanding of when, where and how magnetic cues affect the decision-making process of birds and hence, their realised migratory behaviour in the wild. This understanding is especially important to interpret the results of laboratory experiments in an ecologically appropriate way. In this review, we summarise the current findings about the role of magnetic cues for migratory decisions in songbirds. First, we review the methodological principles for orientation and navigation research, specifically by comparing experiments on caged birds with experiments on free-flying birds. While cage experiments can show the sensory abilities of birds, studies with free-flying birds can characterise the ecological roles of magnetic cues. Second, we review the migratory stages, from stopover to endurance flight, in which songbirds use magnetic cues for their migratory decisions and incorporate this into a novel conceptual framework. While we lack studies examining whether and when magnetic cues affect orientation or navigation decisions during flight, the role of magnetic cues during stopover is relatively well studied, but mostly in the laboratory. Notably, many such studies have produced contradictory results so that understanding the biological importance of magnetic cues for decisions in free-flying songbirds is not straightforward. One potential explanation is that reproducibility of magnetic-cue experiments is low, probably because variability in the behavioural responses of birds among experiments is high. We are convinced that parts of this variability can be explained by species-specific and context-dependent reactions of birds to the study conditions and by the bird's high flexibility in whether they include magnetic cues in a decision or not. Ultimately, this review should help researchers in the challenging field of magnetoreception to design experiments meticulously and interpret results of such studies carefully by considering the migration ecology of their focal species.

Pre-migratory flights in migrant songbirds: the ecological and evolutionary importance of understudied exploratory movements.

Across the animal kingdom, from honeybees to cranes to beavers, exploratory movements to exploit resources, scout prospective territories, or otherwise gain valuable experiences and information that promote fitness have been documented. For example, exploratory movements to investigate potential dispersal targets have been observed in roe deer, Northern cardinals, and tigers alike. However, despite how widespread these movements are, a cohesive definition of exploratory movements has been lacking. We first provide a clear definition of exploratory movements, and use one particular group-migratory songbirds-to catalogue exploratory movements across the annual cycle. The exceptional mobility of migratory songbirds results in exploratory movements not only at a local scale, but also on a regional scale, both in and out of the breeding season. We review the extent to which these movements are made within this group, paying particular attention to how such movements confer fitness benefits, as by securing high-quality territories, prospecting for extra-pair paternity, or even exploiting ephemeral resources. We then zoom in one step further to a particular exploratory movement that has been, to date, almost completely overlooked within this group: that of pre-migratory flights. These flights, which occur during the transitional period between the stationary breeding period and the onset of migration, occur at night and may not be made by all individuals in a population-reasons why these flights have been heretofore critically understudied. We provide the first definition for this behaviour, summarise the current knowledge of this cryptic movement, and hypothesise what evolutionary/ecological advantages conducting it may confer to the individuals that undertake it. As these flights provide experience to the individuals that undertake them, we expect that birds that make pre-migratory flights are better equipped to survive migration (direct fitness benefits) and, due to orientation/navigation abilities, may also reach preferred territories on breeding and wintering grounds faster (indirect fitness benefits). We hope to encourage ecologists to consider such hidden movements in their research concepts and to enhance the framework of movement ecology by this behaviour due to its presumed high biological importance to the annual cycle of birds.

The urge to breed early: Similar responses to environmental conditions in short- and long-distance migrants during spring migration.

Birds migrating different distances experience different temporal, energetic, physiological, and physical constraints throughout migration, which is reflected in their migration strategy. Consequently, we predict different behavioral decisions to similar environmental cues between short- and long-distance migrants, which has been documented for autumn migration. Here, we focus on the question whether trade-off decisions regarding departure, routing, and landing when alternating between migratory endurance flights and stopovers also differ during spring migration. As early arrivals at the breeding grounds should be ultimately favored regardless of migration distance, selection may favor more similar behavioral decisions in spring than in autumn. We radio-tagged short- and long-distance migratory songbirds at stopover sites along the German North Sea coast during spring and automatically tracked their migratory behavior using a large-scale network of receiver stations. Once departed, birds could either cross the sea or detour along the coast. We corrected for spatially biased detection data, using a hierarchical multistate model to assess how birds respond to variation in environmental conditions in their day-to-day departure decisions and route selection. The day-to-day departure probability was higher in long-distance migrants independently of the routing decision. Irrespective of migration distance, all species more likely departed under light winds and rainless conditions, while the influence of air pressure change and relative humidity was species-specific. By accounting for detection probabilities, we estimated that about half of all individuals of each species crossed the sea but did not find differences between short- and long-distance migrants. Offshore flights were more likely when winds blew offshore and began earlier within the night compared with onshore flights. Our results suggest that selection more similarly affects birds of different migration distances in spring than in autumn. These findings put the focus toward how ultimate mechanisms may shape departure and routing decisions differently between migration seasons.

Gauge-and-compass migration: inherited magnetic headings and signposts can adapt to changing geomagnetic landscapes.

BACKGROUND: For many migratory species, inexperienced (naïve) individuals reach remote non-breeding areas independently using one or more inherited compass headings and, potentially, magnetic signposts to gauge where to switch between compass headings. Inherited magnetic-based migration has not yet been assessed as a population-level process, particularly across strong geomagnetic gradients or where long-term geomagnetic shifts (hereafter, secular variation) could create mismatches with magnetic headings. Therefore, it remains unclear whether inherited magnetic headings and signposts could potentially adapt to secular variation under natural selection. METHODS: To address these unknowns, we modelled migratory orientation programs using an evolutionary algorithm incorporating global geomagnetic data (1900-2023). Modelled population mixing incorporated both natal dispersal and trans-generational inheritance of magnetic headings and signposts, including intrinsic (stochastic) variability in inheritance. Using the model, we assessed robustness of trans-hemispheric migration of a migratory songbird whose Nearctic breeding grounds have undergone rapid secular variation (mean 34° clockwise drift in declination, 1900-2023), and which travels across strong geomagnetic gradients via Europe to Africa. RESULTS: Model-evolved magnetic-signposted migration was overall successful throughout the 124-year period, with 60-90% mean successful arrival across a broad range in plausible precision in compass headings and gauging signposts. Signposted migration reduced trans-Atlantic flight distances and was up to twice as successful compared with non-signposted migration. Magnetic headings shifted plastically in response to the secular variation (mean 16°-17° among orientation programs), whereas signpost latitudes were more constrained (3°-5° mean shifts). This plasticity required intrinsic variability in inheritance (model-evolved σ ≈ 2.6° standard error), preventing clockwise secular drift from causing unsustainable open-ocean flights. CONCLUSIONS: Our study supports the potential long-term viability of inherited magnetic migratory headings and signposts, and illustrates more generally how inherited migratory orientation programs can both mediate and constrain evolution of routes, in response to global environmental change.

The oxidative balance and stopover departure decisions in a medium- and a long-distance migrant.

BACKGROUND: Birds have extremely elevated metabolic rates during migratory endurance flight and consequently can become physiologically exhausted. One feature of exhaustion is oxidative damage, which occurs when the antioxidant defense system is overwhelmed by the production of damaging reactive oxygen species (ROS). Migrating birds have been shown to decrease the amount of oxidative lipid damage during stopovers, relatively stationary periods in between migratory flights. It has therefore been argued that, in addition to accumulating fuel, one of the functions of stopover is to restore the oxidative balance. If this is so, we would expect that migrating birds are unlikely to resume migration from stopover when they still have high amounts of lipid damage. METHODS: To test this hypothesis, we measured parameters of the oxidative balance and related these to stopover departure decisions of song thrushes (Turdus philomelos) and northern wheatears (Oenanthe oenanthe), a medium- and long-distance songbird migrant, respectively. We measured malondialdehyde (MDA) concentration, a biomarker for oxidative lipid damage, and total non-enzymatic antioxidant capacity (AOX), an overall biomarker of protection against ROS. Stopover departure decisions were determined using a fully automated telemetry system set-up on our small island study site. RESULTS: The decision to resume migration was not related with MDA concentration in either study species, also not when this was corrected for circulating fatty acid concentrations. Similarly, AOX did not affect this decision, also not when corrected for uric-acid concentration. The time within the night when birds departed also was not affected by MDA concentration or AOX. However, confirming earlier observations, we found that in both species, fat individuals were more likely to depart than lean individuals, and fat northern wheatears departed earlier within the night than lean conspecifics. Northern wheatears additionally departed earlier in spring with more southerly winds. CONCLUSIONS: We found no support for the idea that stopovers departure decisions are influenced by parameters of the oxidative balance. We discuss possible reasons for this unexpected finding.

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 Yellow-browed Warbler (Phylloscopus inornatus) as a model to understand vagrancy and its potential for the evolution of new migration routes.

Why and how new migration routes emerge remain fundamental questions in ecology, particularly in the context of current global changes. In its early stages, when few individuals are involved, the evolution of new migration routes can be easily confused with vagrancy, i.e. the occurrence of individuals outside their regular breeding, non-breeding or migratory distribution ranges. Yet, vagrancy can in theory generate new migration routes if vagrants survive, return to their breeding grounds and transfer their new migration route to their offspring, thus increasing a new migratory phenotype in the population. Here, we review the conceptual framework and empirical challenges of distinguishing regular migration from vagrancy in small obligate migratory passerines and explain how this can inform our understanding of migration evolution. For this purpose, we use the Yellow-browed Warbler (Phylloscopus inornatus) as a case study. This Siberian species normally winters in southern Asia and its recent increase in occurrence in Western Europe has become a prominent evolutionary puzzle. We first review and discuss available evidence suggesting that the species is still mostly a vagrant in Western Europe but might be establishing a new migration route initiated by vagrants. We then list possible empirical approaches to check if some individuals really undertake regular migratory movements between Western Europe and Siberia, which would make this species an ideal model for studying the links between vagrancy and the emergence of new migratory routes.

Predicting performance of naïve migratory animals, from many wrongs to self-correction.

Migratory orientation of many animals is inheritable, enabling inexperienced (naïve) individuals to migrate independently using a geomagnetic or celestial compass. It remains unresolved how naïve migrants reliably reach remote destinations, sometimes correcting for orientation error or displacement. To assess naïve migratory performance (successful arrival), we simulate and assess proposed compass courses for diverse airborne migratory populations, accounting for spherical-geometry effects, compass precision, cue transfers (e.g., sun to star compass), and geomagnetic variability. We formulate how time-compensated sun-compass headings partially self-correct, according to how inner-clocks are updated. For the longest-distance migrations simulated, time-compensated sun-compass courses are most robust to error, and most closely resemble known routes. For shorter-distance nocturnal migrations, geomagnetic or star-compass courses are most robust, due to not requiring nightly cue-transfers. Our predictive study provides a basis for assessment of compass-based naïve migration and mechanisms of self-correction, and supports twilight sun-compass orientation being key to many long-distance inaugural migrations.

A magnetic pulse does not affect free-flight navigation behaviour of a medium-distance songbird migrant in spring.

Current evidence suggests that migratory animals extract map information from the geomagnetic field for true navigation. The sensory basis underlying this feat is elusive, but presumably involves magnetic particles. A common experimental manipulation procedure consists of pre-treating animals with a magnetic pulse, with the aim of re-magnetising particles to alter the internal representation of the external field prior to a navigation task. Although pulsing provoked deflected bearings in caged songbirds, analogous studies with free-flying songbirds yielded inconsistent results. Here, we pulsed European robins (Erithacus rubecula) at an offshore stopover site during spring migration and monitored their free-flight behaviour with a regional-scale network of radio-receiving stations. We found no pulse effect on departure probability, nocturnal departure timing departure direction or consistency of flight direction. This suggests either no use of the geomagnetic map by our birds, or that magnetic pulses do not affect the sensory system underlying geomagnetic map detection.

Can differential fatty acid composition help migrating birds to limit oxidative lipid damage?

During migratory endurance flights, which are energetically very demanding, migrants have to deal with prolonged elevated generation of reactive oxygen species (ROS). To limit the damaging actions that ROS have on lipids and proteins, migrating birds are known to upregulate their antioxidant defence system. However, there may be additional ways to limit oxidative damage incurred from flying. Migratory endurance flights are fuelled mainly with fatty acids (FAs), and the risk of their peroxidation (resulting in oxidative lipid damage) increases with the number of double bonds in a FA, with polyunsaturated FAs (2 or more double bonds, PUFAs) being most peroxidation-prone. By fuelling their flights with relatively few PUFAs, migratory birds could thus limit oxidative lipid damage. Within migratory birds, there is considerable variation in the length of their flights, with nocturnal migrants making lengthier flight bouts, thus more likely to experience lengthier periods of elevated ROS production, than diurnal migrants. However, whether migrants making lengthier flights incur more oxidative lipid damage is unknown. Neither is it known whether flight length and FA composition are associated. Therefore, we determined plasmatic malondialdehyde level, a marker of oxidative lipid damage, and FA composition of three nocturnal and two diurnal migrant species caught at an autumn stopover site. We found little inter-specific variation in malondialdehyde level, indicating that the amount of oxidative lipid damage was comparable across the species. In contrast, the species strongly differed in their plasmatic FA composition. The nocturnal migrants had significantly lower relative PUFA levels than both diurnal migrants, an effect mainly attributable to linoleic acid, an essential (strictly dietary) FA. Consequently, the susceptibility of plasmatic FAs to lipid peroxidation was significantly lower in the nocturnal than diurnal migrants. Because in birds, energy expenditure during flight decreases with the degree of FA unsaturation, we interpret our observation of lower PUFA levels in nocturnal migrants as support for the idea that utilizing PUFA-poor fuel can help migrating birds to curb oxidative lipid damage.

Understanding the ecological and evolutionary function of stopover in migrating birds.

Global movement patterns of migratory birds illustrate their fascinating physical and physiological abilities to cross continents and oceans. During their voyages, most birds land multiple times to make so-called 'stopovers'. Our current knowledge on the functions of stopover is mainly based on the proximate study of departure decisions. However, such studies are insufficient to gauge fully the ecological and evolutionary functions of stopover. If we study how a focal trait, e.g. changes in energy stores, affects the decision to depart from a stopover without considering the trait(s) that actually caused the bird to land, e.g. unfavourable environmental conditions for flight, we misinterpret the function of the stopover. It is thus important to realise and acknowledge that stopovers have many different functions, and that not every migrant has the same (set of) reasons to stop-over. Additionally, we may obtain contradictory results because the significance of different traits to a migrant is context dependent. For instance, late spring migrants may be more prone to risk-taking and depart from a stopover with lower energy stores than early spring migrants. Thus, we neglect that departure decisions are subject to selection to minimise immediate (mortality risk) and/or delayed (low future reproductive output) fitness costs. To alleviate these issues, we first define stopover as an interruption of migratory endurance flight to minimise immediate and/or delayed fitness costs. Second, we review all probable functions of stopover, which include accumulating energy, various forms of physiological recovery and avoiding adverse environmental conditions for flight, and list potential other functions that are less well studied, such as minimising predation, recovery from physical exhaustion and spatiotemporal adjustments to migration. Third, derived from these aspects, we argue for a paradigm shift in stopover ecology research. This includes focusing on why an individual interrupts its migratory flight, which is more likely to identify the individual-specific function(s) of the stopover correctly than departure-decision studies. Moreover, we highlight that the selective forces acting on stopover decisions are context dependent and are expected to differ between, e.g. K-/r-selected species, the sexes and migration strategies. For example, all else being equal, r-selected species (low survival rate, high reproductive rate) should have a stronger urge to continue the migratory endurance flight or resume migration from a stopover because the potential increase in immediate fitness costs suffered from a flight is offset by the expected higher reproductive success in the subsequent breeding season. Finally, we propose to focus less on proximate mechanisms controlling landing and departure decisions, and more on ultimate mechanisms to identify the selective forces shaping stopover decisions. Our ideas are not limited to birds but can be applied to any migratory species. Our revised definition of stopover and the proposed paradigm shift has the potential to stimulate a fruitful discussion towards a better evolutionary ecological understanding of the functions of stopover. Furthermore, identifying the functions of stopover will support targeted measures to conserve and restore the functionality of stopover sites threatened by anthropogenic environmental changes. This is especially important for long-distance migrants, which currently are in alarming decline.

No apparent effect of a magnetic pulse on free-flight behaviour in northern wheatears (Oenanthe oenanthe) at a stopover site.

Naïve migrants reach their wintering grounds following a clock-and-compass strategy. During these inaugural migrations, birds internalise, among others, cues from the Earth's magnetic field to create a geomagnetic map, with which they navigate to destinations familiar to them on subsequent migrations. Geomagnetic map cues are thought to be sensed by a magnetic-particle-based receptor, which can be specifically affected by a magnetic pulse. Indeed, the orientation of experienced but not naïve birds was compromised after magnetic pulsing, indicating geomagnetic map use. Little is known about the importance of this putative magnetoreceptor for navigation and decision-making in free-flying migrants. Therefore, we studied in unprecedented detail how a magnetic pulse would affect departure probability, nocturnal departure timing, departure direction and consistency in flight direction over 50-100 km in experienced and naïve long-distant migrant songbirds using a large-scale radio-tracking system. Contrary to our expectations and despite a high sample size (ntotal = 137) for a free-flight study, we found no significant after-effect of the magnetic pulse on the migratory traits, suggesting the geomagnetic map is not essential for the intermediate autumn migration phase. These findings warrant re-thinking about perception and use of geomagnetic maps for migratory decisions within a sensory and ecological context.

High individual repeatability of the migratory behaviour of a long-distance migratory seabird.

BACKGROUND: Understanding the evolution of migration requires knowledge of the patterns, sources, and consequences of variation in migratory behaviour, a need exacerbated by the fact that many migratory species show rapid population declines and require knowledge-based conservation measures. We therefore need detailed knowledge on the spatial and temporal distribution of individuals across their annual cycle, and quantify how the spatial and temporal components of migratory behaviour vary within and among individuals. METHODS: We tracked 138 migratory journeys undertaken by 64 adult common terns (Sterna hirundo) from a breeding colony in northwest Germany to identify the annual spatiotemporal distribution of these birds and to evaluate the individual repeatability of eleven traits describing their migratory behaviour. RESULTS: Birds left the breeding colony early September, then moved south along the East Atlantic Flyway. Wintering areas were reached mid-September and located at the west and south coasts of West Africa as well as the coasts of Namibia and South Africa. Birds left their wintering areas late March and reached the breeding colony mid-April. The timing, total duration and total distance of migration, as well as the location of individual wintering areas, were moderately to highly repeatable within individuals (repeatability indexes: 0.36-0.75, 0.65-0.66, 0.93-0.94, and 0.98-1.00, respectively), and repeatability estimates were not strongly affected by population-level inter-annual variation in migratory behaviour. CONCLUSIONS: We found large between-individual variation in common tern annual spatiotemporal distribution and strong individual repeatability of several aspects of their migratory behaviour.

The avian lightweights: Trans-Saharan migrants show lower lean body mass than short-/medium-distance migrants.

Avian trans-Saharan migrants travelling long distances and crossing ecological barriers experience different constraints in terms of time, energy and safety than short-/medium-distance migrants without barrier-crossings. As such, natural selection shapes the aerodynamic properties of these groups differently. Yet, to the best of our knowledge, we lack information on whether natural selection has contributed to reducing energetic flight costs through generally lower body mass in trans-Saharan migrants. To fill parts of this gap, we investigated this eco-morphological pattern in 5,410 individuals of 22 Palearctic songbird species ranging from short-/medium-distance to trans-Saharan migrants. We used individual size-independent scaled lean body mass values based on wing length as a measure of body size and, for the first time, precisely determined lean body mass values by direct measurements via quantitative magnetic resonance technology. Scaled lean body mass for a given body size was significantly higher in short-/medium-distance migrants than in trans-Saharan migrants. Although scaled lean body mass significantly decreased with increasing migration distance in short-/medium-distance migrants, no such effect was found in trans-Saharan migrants. Our results thus show an eco-morphological pattern relating species' lean body mass not only to migration distance but also to migration group. This suggests that selective effects of the presence/absence of ecological barriers and/or of a threshold level for migration distance on migrant birds may be more important than the linear continuum of migration distance per se.

Temperature change is an important departure cue in nocturnal migrants: controlled experiments with wild-caught birds in a proof-of-concept study.

The decision-making process of migrating birds at stopover sites is a complex interplay of the innate migration program and both intrinsic and extrinsic factors. While it is well studied how variation in precipitation, wind and air pressure influence this process, there is less evidence of the effects of temperature changes on the departure decision. Thus, we lack knowledge on how the predicted changes due to global climate change in temperature alone may affect the decision-making process during migration. Aiming to fill parts of this gap, we conducted a proof-of-concept study by manipulating the ambient temperature of temporarily confined wild-caught migrant songbirds under constant feeding conditions. In spring, departure probability increased with a 20°C rise in temperature for both a medium-distance migrant (European robin, Erithacus rubecula) and a long-distance migrant (northern wheatear, Oenanthe oenanthe), and in autumn, departure probabilities of the long-distance migrant both decreased with a 20°C rise and increased with a 20°C drop. Consequently, the temperature is an important departure cue influencing the decision-making process of migrating songbirds. Incorporating causal relationships between changes in temperature and departure probability in migration models could substantially improve our ability to predict the effects of climate change on the phenology of migratory birds.

A quasi-experimental approach using telemetry to assess migration-strategy-specific differences in the decision-making processes at stopover.

BACKGROUND: Migrant birds travel between their breeding areas and wintering grounds by alternating energetically and physiologically demanding flights with periods of rest and fuelling, so-called stopovers. An important intrinsic factor influencing the decision to resume migration is the amount of energy stores available for the next flight. Correlative studies with free-flying birds and experimental studies with caged birds have shown that the amount of energy stores affects the day-to-day, within-day and the directional decision of departure. The methodological advantages of both the correlative and experimental approach are combined when radio-tagging many individuals on the same day and subsequently determining the departure decisions at a high spatiotemporal resolution. Making use of such a quasi-experimental approach with an automated radio-tracking system at stopover, we studied the effect of energy stores on departure decisions and whether they vary between species of different migration strategies experiencing contrasting time constraints. For this, we chose a long-distance migrant, the common redstart (Phoenicurus phoenicurus), and a medium-distance migrant, the European robin (Erithacus rubecula), because the former has to travel at relatively higher speed to reach its wintering ground in a reasonable time at the expense of relatively higher energetic costs for travelling than the latter. RESULTS: Common redstarts with higher energy stores were more likely to resume migration than their conspecifics with lower energy stores, whereas this pattern was absent in the European robins. The amount of energy stores significantly affected the timing of departure within the day, with large energy stores yielding early departures in both species. Departure directions from the stopover site during the first night after capture were oriented towards the seasonally appropriate direction but were not affected by variation in energy stores. CONCLUSIONS: We demonstrate the importance of variation in energy stores on the departure decisions and that it may affect species with different migration strategies dissimilarly in autumn. Nevertheless, knowledge of other intrinsic factors, such as feeding conditions, health status and physiological consequences of previous flights, is additionally required to better understand the departure decisions of migrants, as this is the key to providing an overall assessment of the decision-making process.

Can Mitogenomes of the Northern Wheatear (Oenanthe oenanthe) Reconstruct Its Phylogeography and Reveal the Origin of Migrant Birds?

The Northern Wheatear (Oenanthe oenanthe, including the nominate and the two subspecies O. o. leucorhoa and O. o. libanotica) and the Seebohm's Wheatear (Oenanthe seebohmi) are today regarded as two distinct species. Before, all four taxa were regarded as four subspecies of the Northern Wheatear. Their classification has exclusively been based on ecological and morphological traits, while their molecular characterization is still missing. With this study, we used next-generation sequencing to assemble 117 complete mitochondrial genomes covering O. o. oenanthe, O. o. leucorhoa and O. seebohmi. We compared the resolution power of each individual mitochondrial marker and concatenated marker sets to reconstruct the phylogeny and estimate speciation times of three taxa. Moreover, we tried to identify the origin of migratory wheatears caught on Helgoland (Germany) and on Crete (Greece). Mitogenome analysis revealed two different ancient lineages that separated around 400,000 years ago. Both lineages consisted of a mix of subspecies and species. The phylogenetic trees, as well as haplotype networks are incongruent with the present morphology-based classification. Mitogenome could not distinguish these presumed species. The genetic panmixia among present populations and taxa might be the consequence of mitochondrial introgression between ancient wheatear populations.