Eco-evolutionary drivers of avian migratory connectivity.

Migratory connectivity, reflecting the extent by which migrants tend to maintain their reciprocal positions in seasonal ranges, can assist in the conservation and management of mobile species, yet relevant drivers remain unclear. Taking advantage of an exceptionally large (~150,000 individuals, 83 species) and more-than-a-century-long dataset of bird ringing encounters, we investigated eco-evolutionary drivers of migratory connectivity in both short- and long-distance Afro-Palearctic migratory birds. Connectivity was strongly associated with geographical proxies of migration costs and was weakly influenced by biological traits and phylogeny, suggesting the evolutionary lability of migratory behaviour. The large intraspecific variability in avian migration strategies, through which most species geographically split into distinct migratory populations, explained why most of them were significantly connected. By unravelling key determinants of migratory connectivity, our study improves knowledge about the resilience of avian migrants to ecological perturbations, providing a critical tool to inform transboundary conservation and management strategies at the population level.

Weather at the winter and stopover areas determines spring migration onset, progress, and advancements in Afro-Palearctic migrant birds.

Climate change causes changes in the timing of life cycle events across all trophic groups. Spring phenology has mostly advanced, but large, unexplained, variations are present between and within species. Each spring, migratory birds travel tens to tens of thousands of kilometers from their wintering to their breeding grounds. For most populations, large uncertainties remain on their exact locations outside the breeding area, and the time spent there or during migration. Assessing climate (change) effects on avian migration phenology has consequently been difficult due to spatial and temporal uncertainties in the weather potentially affecting migration timing. Here, we show for six trans-Saharan long-distance migrants that weather at the wintering and stopover grounds almost entirely (∼80%) explains interannual variation in spring migration phenology. Importantly, our spatiotemporal approach also allows for the systematic exclusion of influences at other locations and times. While increased spring temperatures did contribute strongly to the observed spring migration advancements over the 55-y study period, improvements in wind conditions, especially in the Maghreb and Mediterranean, have allowed even stronger advancements. Flexibility in spring migration timing of long-distance migrants to exogenous factors has been consistently underestimated due to mismatches in space, scale, time, and weather variable type.

Transcriptome signature changes in the liver of a migratory passerine.

The liver plays a principal role in avian migration. Here, we characterised the liver transcriptome of a long-distance migrant, the Northern Wheatear (Oenanthe oenanthe), sampled at different migratory stages, looking for molecular processes linked with adaptations to migration. The analysis of the differentially expressed genes suggested changes in the periods of the circadian rhythm, variation in the proportion of cells in G1/S cell-cycle stages and the putative polyploidization of this cell population. This may explain the dramatic increment in the liver's metabolic capacities towards migration. Additionally, genes involved in anti-oxidative stress, detoxification and innate immune responses, lipid metabolism, inflammation and angiogenesis were regulated. Lipophagy and lipid catabolism were active at all migratory stages and increased towards the fattening and fat periods, explaining the relevance of lipolysis in controlling steatosis and maintaining liver health. Our study clears the way for future functional studies regarding long-distance avian migration.

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.

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.

The role of ketogenesis in the migratory fattening of the northern wheatear Oenanthe oenanthe.

The fuelling capacity of migratory birds and their ability to avoid health conditions derived from the subsequent fat overload are exceptional among vertebrates. In this work, we screen the expression of the genes involved in the production of ketone bodies (KB) in the liver of northern wheatears (Oenanthe oenanthe) during the development and resolution of migratory fattening. Thirteen genes were found to be regulated among the migratory stages. Based on the dynamics of gene expression, we concluded that KB play a versatile role in wheatears' energy metabolism homeostasis. The ketogenic pathway can adaptively: (i) provide carbon equivalents for lipogenesis, speeding up fuelling; (ii) replace glucose during long-distance flights using lipids as the substrate; (iii) act as a floodgate to avoid steatosis; and (iv) might provide a metabolic solution to defatting in captive birds.

Autumn bird migration phenology: A potpourri of wind, precipitation and temperature effects.

Climate change has caused a clear and univocal trend towards advancement in spring phenology. Changes in autumn phenology are much more diverse, with advancement, delays, and 'no change' all occurring frequently. For migratory birds, patterns in autumn migration phenology trends have been identified based on ecological and life-history traits. Explaining interspecific variation has nevertheless been challenging, and the underlying mechanisms have remained elusive. Radar studies on non-species-specific autumn migration intensity have repeatedly suggested that there are strong links with weather. In long-term species-specific studies, the variance in autumn migration phenology explained by weather has, nevertheless, been rather low, or a relationship was even lacking entirely. We performed a spatially explicit time window analysis of weather effects on mean autumn passage of four trans-Saharan and six intra-European passerines to gain insights into this apparent contradiction. We analysed data from standardized daily captures at the Heligoland island constant-effort site (Germany), in combination with gridded daily temperature, precipitation and wind data over a 55-year period (1960-2014), across northern Europe. Weather variables at the breeding and stopover grounds explained up to 80% of the species-specific interannual variability in autumn passage. Overall, wind conditions were most important. For intra-European migrants, wind was even twice as important as either temperature or precipitation, and the pattern also held in terms of relative contributions of each climate variable to the temporal trends in autumn phenology. For the trans-Saharan migrants, however, the pattern of relative trend contributions was completely reversed. Temperature and precipitation had strong trend contributions, while wind conditions had only a minor impact because they did not show any strong temporal trends. As such, understanding species-specific effects of climate on autumn phenology not only provides unique insights into each species' ecology but also how these effects shape the observed interspecific heterogeneity in autumn phenological trends.

An exception to the rule: Captivity does not stress wild migrating northern wheatears.

Wild animals typically suffer from stress when brought into captivity. This stress is characterized by elevated circulating glucocorticoid levels and weight loss. We here describe for the first time a case where a wild animal, the long-distance migrating northern wheatear, does not show signs of stress when caged. We captured these birds on a stopover site during their spring migration and caged them individually with ad libitum access to food and water. The birds were divided into four groups and were blood-sampled immediately in the field, a few hours after caging, one day after caging, or three days after caging, respectively. From these blood-samples we determined circulating corticosterone level. Food intake and body mass were also monitored. We found that, with very few exceptions, corticosterone levels were low and did not differ among the groups. Accordingly, almost all birds consumed huge quantities of food and substantially increased their body mass. Together these results clearly show that caging does not result in indications of stress in wild migrating northern wheatears. Confinement-specific conditions such as restricted movement normally stress animals. We suggest migratory birds may not perceive such conditions as stressors due to their hyperphagic state, a notion that requires further testing.

The influence of weather on avian spring migration phenology: What, where and when?

Over the past decades, spring arrival and passage of most short- and medium-distance migrating birds in the Northern Hemisphere have advanced. Changes in spring temperature at the passage or arrival area have been most frequently shown to be related to these changes in spring migration phenology. In most studies, preliminary assumptions are made on both the spatial location and the specific time frame of the weather influencing spring migration phenology. We performed a spatially explicit time-window analysis of the effect of weather on mean spring passage dates of nine short- and medium-distance passerines. We analysed data from standardized daily captures at the Helgoland (Germany) constant-effort site, in combination with gridded daily temperature, precipitation and wind data from the NCEP data set over a 55-year period (1960-2014), across the whole of West Europe and North Africa. Although we allowed for a time window of any length at any location, nevertheless incorporating various measures to avoid spurious correlations, time windows at the likely wintering or spring stopover grounds were almost exclusively selected as the best predicting variables (96%-100% of identified variables). The weather variables at the wintering and stopover grounds explain up to 77% of the interannual variability in spring passage. Yet, the response of spring migration phenology to weather at the winter or stopover areas does not fully explain the observed trends. Spring migration phenology is, hence, strongly driven by weather at the wintering and stopover grounds, but additional mechanisms are needed to fully explain the advancement of spring migration. Our results also clearly show that previously illustrated correlations, or the lack thereof, between spring migration phenology and weather at the passage or arrival location are due to spatio-temporal correlations in the weather data. This spatial mismatch might have led to false conclusions, especially the further away the wintering or stopover sites are.

Challenging a 15-year-old claim: The North Atlantic Oscillation index as a predictor of spring migration phenology of birds.

Many migrant bird species that breed in the Northern Hemisphere show advancement in spring arrival dates. The North Atlantic Oscillation (NAO) index is one of the climatic variables that have been most often investigated and shown to be correlated with these changes in spring arrival. Although the NAO is often claimed to be a good predictor or even to have a marked effect on interannual changes in spring migration phenology of Northern Hemisphere breeding birds, the results on relations between spring migration phenology and NAO show a large variety, ranging from no, over weak, to a strong association. Several factors, such as geographic location, migration phase, and the NAO index time window, have been suggested to partly explain these observed differences in association. A combination of a literature meta-analysis, and a meta-analysis and sliding time window analysis of a dataset of 23 short- and long-distance migrants from the constant-effort trapping garden at Helgoland, Germany, however, paints a completely different picture. We found a statistically significant overall effect size of the NAO on spring migration phenology (coefficient = -0.14, SE = 0.054), but this on average only explains 0%-6% of the variance in spring migration phenology across all species. As such, the value and biological meaning of the NAO as a general predictor or explanatory variable for climate change effects on migration phenology of birds, seems highly questionable. We found little to no definite support for previously suggested factors, such as geographic location, migration phenology phase, or the NAO time window, to explain the heterogeneity in correlation differences. We, however, did find compelling evidence that the lack of accounting for trends in both time series has led to strongly inflated (spurious) correlations in many studies (coefficient = -0.13, SE = 0.019).

Flexible reaction norms to environmental variables along the migration route and the significance of stopover duration for total speed of migration in a songbird migrant.

BACKGROUND: Predicting the consequences of continuing anthropogenic changes in the environment for migratory behaviours such as phenology remains a major challenge. Predictions remain particularly difficult, because our knowledge is based on studies from single-snapshot observations at specific stopover sites along birds' migration routes. However, a general understanding on how birds react to prevailing environmental conditions, e.g. their 'phenotypic reaction norm', throughout the annual cycle and along their entire migration routes is required to fully understand how migratory birds respond to rapid environmental change. RESULTS: Here, we provide direct evidence that northern wheatears (Oenanthe oenanthe) from a breeding population in Alaska adjusted their probability to resume migration as well as the distance covered per night, i.e. travel speed, to large-scale environmental conditions experienced along their 15,000 km migratory route on both northwards and southwards migrations. These adjustments were found to be flexible in space and time. At the beginning of autumn migration, northern wheatears showed high departure probabilities and high travel speeds at low surface air temperatures, while far away from Alaska both traits decreased with increasing air temperatures. In spring, northern wheatears increasingly exploited flow assistance with season, which is likely a behavioural adjustment to speed up migration by increasing the distance travelled per night. Furthermore, the variation in total stopover duration but not in travel speed had a significant effect on the total speed of migration, indicating the prime importance of total stopover duration in the overall phenology of bird migration. CONCLUSION: Northern wheatears from Alaska provide evidence that the phenotypic reaction norm to a set of environmental conditions cannot be generalized to universal and persistent behavioural reaction pattern across entire migratory pathways. This highlights the importance of full annual-cycle studies on migratory birds to better understand their response to the environment. Understanding the mechanisms behind phenotypic plasticity during migration is particularly important in the assessment of whether birds can keep pace with the potentially increasing phenological mismatches observed on the breeding grounds.

Endogenous control of fuelling in a migratory songbird.

The Northern wheatear (Oenanthe oenanthe) is a small long-distance migratory songbird that breeds throughout the Northern hemisphere and winters in sub-Saharan Africa. The main components of its migratory behaviour, i.e. seasonal migratory restlessness and body mass changes, have been shown to be under endogenous control. However, it is still unknown whether the disposition to accumulate fuel reserves is an inherited trait. We cross-bred Northern wheatears from two populations known to accumulate different amounts of fuel in a common-garden setup, and measured their maximum fuelling as the difference between the lightest and heaviest body mass recorded over each year for 4 years or longer. We used the largest value as a measure of maximum voluntary fuelling potential. F1-generation "hybrids" showed intermediate values to those of the parent populations. It was previously shown that in the wheatear the amount of fuel accumulated is closely linked to the presence of large ecological barriers to cross. This study shows that this adaptation has been fixed at the genetic level, and that intermediate traits are transferred to the next generation, with possible implications on the viability of such individuals in nature.

Baseline corticosterone levels are higher in migrating than sedentary common blackbirds in autumn, but not in spring.

Corticosterone at baseline levels is thought to be mainly involved in the regulation of uptake, storage and release of energy, processes central to avian migration. Consequently, corticosterone levels are thought to be upregulated during migration, but the temporal pattern of its secretion during migration is not well defined. For example, although it appears that corticosterone levels decrease from flight to stopover, it is unknown if levels at stopover are still elevated and it is largely unclear how these levels compare to non-migratory life-history stages. Furthermore, what role corticosterone plays in crucial migratory processes, such as refueling and departure from stopover, is far from understood. We here determined baseline corticosterone levels in migrating and resident common blackbirds (Turdus merula), sampled simultaneously on Helgoland, a stopover site that also supports a sedentary breeding population. In autumn, migrants had higher corticosterone levels than residents, but in spring levels did not differ between the two groups. Corticosterone levels of migrants were very similar in spring and autumn, whereas in residents levels tended to be higher in spring than autumn. Higher levels in residents in spring than autumn most likely reflect the higher daily workload faced by birds during the pre-breeding than the post-breeding period. Our study thus indicates that, relative to the levels observed in residents in autumn, in spring baseline corticosterone levels were moderately elevated in both migrants and residents and that in autumn levels were moderately elevated in migrants only. Currently, corticosterone's main function at stopover is thought to lie in the regulation of departure. Because most migrant blackbirds stay only one or two days on Helgoland, our results are in line with this idea and suggest that migrating blackbirds up-regulated their corticosterone level in anticipation of an oncoming flight bout.

Migratory connectivity and population-specific migration routes in a long-distance migratory bird.

Knowledge about migratory connectivity, the degree to which individuals from the same breeding site migrate to the same wintering site, is essential to understand processes affecting populations of migrants throughout the annual cycle. Here, we study the migration system of a long-distance migratory bird, the Montagu's harrier Circus pygargus, by tracking individuals from different breeding populations throughout northern Europe. We identified three main migration routes towards wintering areas in sub-Saharan Africa. Wintering areas and migration routes of different breeding populations overlapped, a pattern best described by 'weak (diffuse) connectivity'. Migratory performance, i.e. timing, duration, distance and speed of migration, was surprisingly similar for the three routes despite differences in habitat characteristics. This study provides, to our knowledge, a first comprehensive overview of the migration system of a Palaearctic-African long-distance migrant. We emphasize the importance of spatial scale (e.g. distances between breeding populations) in defining patterns of connectivity and suggest that knowledge about fundamental aspects determining distribution patterns, such as the among-individual variation in mean migration directions, is required to ultimately understand migratory connectivity. Furthermore, we stress that for conservation purposes it is pivotal to consider wintering areas as well as migration routes and in particular stopover sites.

Corticosterone, food intake and refueling in a long-distance migrant.

Elevated baseline corticosterone levels function to mobilize energy in predictable life-history stages, such as bird migration. At the same time, baseline corticosterone has a permissive effect on the accumulation of fat stores (fueling) needed for migratory flight. Most migrants alternate flight bouts with stopovers, during which they replenish the fuel used during the preceding flight (refueling). The role of corticosterone in refueling is currently unclear. In a fasting-re-feeding experiment on northern wheatears (Oenanthe oenanthe) in autumn we found that baseline total and free corticosterone levels were negatively related with both food intake and the rate of fuel deposition after fasting. This confirms our earlier findings in wild conspecifics in spring and indicates that corticosterone does not stimulate stopover refueling. Whether the negative relationship between baseline corticosterone level and fuel deposition rate is causal is questionable, because within-individual comparison of corticosterone metabolite levels in droppings did not reveal differences between refueling and control periods. In other words, corticosterone does not appear to be down-regulated during refueling, which would be expected if it directly hampers refueling. We discuss possible correlates of corticosterone level that may explain the negative association between corticosterone and stopover refueling. Additionally, we found that fasting decreases total corticosterone level, which contrasts with previous studies. We propose that the difference is due to the other studies being conducted outside of the migration life-history stage, and provide a possible explanation for the decrease in corticosterone during fasting in migrating birds.

When and where does mortality occur in migratory birds? Direct evidence from long-term satellite tracking of raptors.

Information about when and where animals die is important to understand population regulation. In migratory animals, mortality might occur not only during the stationary periods (e.g. breeding and wintering) but also during the migration seasons. However, the relative importance of population limiting factors during different periods of the year remains poorly understood, and previous studies mainly relied on indirect evidence. Here, we provide direct evidence about when and where migrants die by identifying cases of confirmed and probable deaths in three species of long-distance migratory raptors tracked by satellite telemetry. We show that mortality rate was about six times higher during migration seasons than during stationary periods. However, total mortality was surprisingly similar between periods, which can be explained by the fact that risky migration periods are shorter than safer stationary periods. Nevertheless, more than half of the annual mortality occurred during migration. We also found spatiotemporal patterns in mortality: spring mortality occurred mainly in Africa in association with the crossing of the Sahara desert, while most mortality during autumn took place in Europe. Our results strongly suggest that events during the migration seasons have an important impact on the population dynamics of long-distance migrants. We speculate that mortality during spring migration may account for short-term annual variation in survival and population sizes, while mortality during autumn migration may be more important for long-term population regulation (through density-dependent effects).

Migratory restlessness in captive individuals predicts actual departure in the wild.

In captivity, migratory birds show increased activity during the time that they would normally migrate. The phenology and intensity of such 'migratory restlessness' has been shown to mirror species- and population-specific migration patterns observed in the wild and has consequently been used as a proxy for the motivation to migrate. Many studies doing so, however, were aiming to explain among-individual variation in migratory behaviour or traits, and not species- or population-specific traits. These studies thus assumed that, also at the level of the individual, migratory restlessness is an accurate proxy for the motivation to migrate. We tested this assumption for the first time and found that it holds; individuals showing very little migratory restlessness remained at stopover for longer than one night, whereas most individuals showing more restlessness departed sooner. This finding validates the use of migratory restlessness as a proxy for the motivation to migrate, thereby justifying the conclusions made in a large body of research on avian migration.

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.

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.

Stopover optimization in a long-distance migrant: the role of fuel load and nocturnal take-off time in Alaskan northern wheatears (Oenanthe oenanthe).

INTRODUCTION: In long-distance migrants, a considerably higher proportion of time and energy is allocated to stopovers rather than to flights. Stopover duration and departure decisions affect consequently subsequent flight stages and overall speed of migration. In Arctic nocturnal songbird migrants the trade-off between a relatively long migration distance and short nights available for travelling may impose a significant time pressure on migrants. Therefore, we hypothesize that Alaskan northern wheatears (Oenanthe oenanthe) use a time-minimizing migration strategy to reach their African wintering area 15,000 km away. RESULTS: We estimated the factors influencing the birds' daily departure probability from an Arctic stopover before crossing the Bering Strait by using a Cormack-Jolly-Seber model. To identify in which direction and when migration was resumed departing birds were radio-tracked. Here we show that Alaskan northern wheatears did not behave as strict time minimizers, because their departure fuel load was unrelated to fuel deposition rate. All birds departed with more fuel load than necessary for the sea crossing. Departure probability increased with stopover duration, evening fuel load and decreasing temperature. Birds took-off towards southwest and hence, followed in general the constant magnetic and geographic course but not the alternative great circle route. Nocturnal departure times were concentrated immediately after sunset. CONCLUSION: Although birds did not behave like time-minimizers in respect of the optimal migration strategies their surplus of fuel load clearly contradicted an energy saving strategy in terms of the minimization of overall energy cost of transport. The observed low variation in nocturnal take-off time in relation to local night length compared to similar studies in the temperate zone revealed that migrants have an innate ability to respond to changes in the external cue of night length. Likely, birds maximized their potential nightly flight range by taking off early in the night which in turn maximizes their overall migration speed. Hence, nocturnal departure time may be a crucial parameter shaping the speed of migration indicating the significance of its integration in future migration models.