Jason W. 2013 Multi-generational longdistance migration of insects: studying the painted lady butterfly in the Western Palaearctic. Ecography, 36 (4). 474-486. 10.1111/j.1600-0587.2012.07738.x Contact CEH NORA team at noraceh@ceh.ac.ukThe NERC and CEH trademarks and logos ('the Trademarks') are registered trademarks of NERC in the UK and other countries, and may not be used without the prior written consent of the Trademark owner. (up to 60 degrees of latitude). The cycle comprises an annual poleward advance of the 73 populations in spring followed by an equatorward return movement in autumn, with returning 74 individuals potentially flying thousands of kilometres. We show that many long-distance 75 migrants take advantage of favourable winds, moving downwind at high elevation (from 76 some tens of metres from the ground to altitudes over 1,000 m), pointing at strong similarities 77 in the flight strategies used by V. cardui and other migrant Lepidoptera. Our results reveal the 78 highly successful strategy that has evolved in these insects, and provide a useful framework 79 for a better understanding of long-distance seasonal migration in the temperate regions 80 worldwide. 81 82 5

The northern wheatear ( Oenanthe oenanthe ) is a small (approx. 25 g), insectivorous migrant with one of the largest ranges of any songbird in the world, breeding from the eastern Canadian Arctic across Greenland, Eurasia and into Alaska (AK). However, there is no evidence that breeding populations in the New World have established overwintering sites in the Western Hemisphere. Using light-level geolocators, we demonstrate that individuals from these New World regions overwinter in northern sub-Sahara Africa, with Alaskan birds travelling approximately 14 500 km each way and an eastern Canadian Arctic bird crossing a wide stretch of the North Atlantic (approx. 3500 km). These remarkable journeys, particularly for a bird of this size, last between one to three months depending on breeding location and season (autumn/spring) and result in mean overall migration speeds of up to 290 km d −1 . Stable-hydrogen isotope analysis of winter-grown feathers sampled from breeding birds generally support the notion that Alaskan birds overwinter primarily in eastern Africa and eastern Canadian Arctic birds overwinter mainly in western Africa. Our results provide the first evidence of a migratory songbird capable of linking African ecosystems of the Old World with Arctic regions of the New World.

Billions of songbirds breeding in the Western Palaearctic cross the largest desert of the world, the Sahara, twice a year. While crossing Europe, the vast majority use an intermittent flight strategy, i.e. fly at night and rest or feed during the day. However, it was long assumed that they overcome the Sahara in a 40 h non-stop flight. In this study, we observed bird migration with radar in the plain sand desert of the Western Sahara (Mauritania) during autumn and spring migration and revealed a clear prevalence of intermittent migration. Massive departures of songbirds just after sunset independent of site and season suggests strongly that songbirds spent the day in the plain desert. Thus, most songbirds cross the Sahara predominantely by the intermittent flight strategy. Autumn migration took place mainly at low altitudes with high temperatures, its density decreased abruptly before sunrise, followed by very little daytime migration. Migration was highly restricted to night-time and matched perfectly the intermittent flight strategy. However, in spring, when migratory flights occurred at much higher altitudes than in autumn, in cool air, about 17% of the songbird migration occurred during the day. This suggests that flying in high temperatures and turbulent air, as is the case in autumn, may lead to an increase in water and/or energy loss and may prevent songbirds from prolonged flights into the day.

In birds, accumulating energy is far slower than spending energy during flight. During migration, birds spend, therefore, most of the time at stopover refueling energy used during the previous flight. This elucidates why current energy stores and actual rate of accumulating energy are likely crucial factors influencing bird's decision when to resume migration in addition to other intrinsic (sex, age) and extrinsic (predation, weather) factors modulating the decision within the innate migration program. After first summarizing how energy stores and stopover durations are generally determined, we critically review that high-energy stores and low rates of accumulating energy were significantly related to high departure probabilities in several bird groups. There are, however, also many studies showing no effect at all. Recent radio-tracking studies highlighted that migrants leave a site either to resume migration or to search for a better stopover location, so-called "landscape movements". Erroneously treating such movements as departures increases the likelihood of type II errors which might mistakenly suggest no effect of either trait on departure. Furthermore, we propose that energy loss during the previous migratory flight in relation to bird's current energy stores and migration strategy significantly affects its urge to refuel and hence its departure decision.

No abstract

Summary1. An innate migration strategy guides birds through space and time. Environmental variation further modulates individual behaviour within a genetically determined frame. In particular, ecological barriers could influence departure direction and its timing. A shift in the migratory direction in response to an ecological barrier could reveal how birds adjust their individual trajectories to environmental cues and body condition. 2. Northern wheatears of the Greenland ⁄ Iceland subspecies Oenanthe oenanthe leucorhoa arrive in Western Europe en route from their West African winter range. They then undergo an endogenously controlled shift in migratory direction from north to north-west to cross a large ecological barrier, the North Atlantic. We radiotracked these songbirds departing from Helgoland, a small island in the North Sea, over an unprecedented range of their journey. 3. Here, we show that both birds' body condition and the wind conditions that they encountered influenced the departure direction significantly. Jointly high fuel loads and favourable wind conditions enabled migrants to cross large stretches of sea. Birds in good condition departed early in the night heading to the sea towards their breeding areas, while birds with low fuel loads and ⁄ or flying in poor weather conditions departed in directions leading towards nearby mainland areas during the entire night. These areas could be reached even after setting off late at night. 4. Behavioural adjustment of migratory patterns is a critical adaptation for crossing ecological barriers. The observed variation in departure direction and time in relation to fuel load and wind revealed that these birds have an innate ability to respond by jointly incorporating internal information (body condition) and external information (wind support).

Background Songbirds following distinct migration strategies (e.g. long- vs. short- to medium-distance migrants) often differ in their speed of migration during autumn and, thus, are assumed to face different time constraints. During migration, most songbird species alternate migratory flights with stopover periods. Many of them restrict these migratory flights to the night, i.e., they are nocturnal migrants. At stopover, nocturnal migrants need to select a specific night (night-to-night decision) and time of night (within-night decision) to resume migration. These departure decisions, which largely determine the speed of migration, are jointly affected by a set of intrinsic and extrinsic factors, i.e., departure cues. Here we aim to assess whether the set of intrinsic and extrinsic factors and the magnitude of their respective effects on stopover departure decisions differs between nocturnally migrating songbird species, depending on their migration strategy and associated time constraints. Methods We radio-tracked migrating Northern Wheatears (Oenanthe oenanthe; long-distance migrant), European robins (Erithacus rubecula) and Common Blackbirds (Turdus merula; both medium-distance migrants) during autumn stopover and analysed their night-to-night and within-night departure timing in relation to intrinsic and extrinsic factors. Results Species generally differed in their departure timing on both temporal scales, with shortest stopovers and earliest nocturnal departures in the long-distance migrant. Some factors, such as day of year, fuel load, cloud cover and crosswind, had consistent effects on stopover departure decisions in all three species. However, species differed in the effects of tailwind assistance, change in atmospheric pressure and air temperature on their stopover departure decisions. Whereas night-to-night decisions were affected by these extrinsic factors in either both or one of the medium-distance migrants, such effects were not found in the long-distance migrant. Conclusions Our results suggest that the general timing of departures in songbirds is affected by the species-specific migration strategy and associated time constraints. Further, they imply that the assessment and usage of specific extrinsic factors, i.e., weather conditions, as departure cues is adjusted based on this migration strategy, with the long-distance migrants being least selective at departure. Other intrinsic and extrinsic factors, however, seem to be used as departure cues independent of migration strategy.

Besides the scientific interest in the quantification of bird migration, there is an increasing need to quantify bird movements for the assessment of bird collision risk with artificial structures. In many environmental impact studies, the radar method is used in an inappropriate manner. The processing of echoes consists often of counting blips within defined screen fields, and the surveyed volume is estimated without reference to the detection probabilities of different ‘target sizes’ (radar cross‐sections). The aim of this paper is to present a procedure to quantify bird migration reliably using radar by stating the theoretical requirements of every single step of this procedure and presenting methodological solutions using our own radar data from extensive field studies. Our methodological solutions can be applied to various radar systems, including widely used ship radar. The procedure presented involves discriminating the echoes of birds and insects and estimating the different detection probabilities of differently ‘sized’ birds (radar cross‐sections). By ignoring the different detection probabilities, density estimations may be wrong by as much as 400%. We fear that quantification of bird migration and predicted bird numbers affected by collisions with artificial structures are in many cases based on unreliable estimates.