Diffuse, short and slow migration among Blue Tits

The knowledge of migration systems in long-distance regular migrants is in many cases extensive. Our understanding of the migratory characteristics of partial migrants, on the other hand, is far more rudimentary. We investigated migratory characteristics of partially migratory Blue Tits Cyanistes caeruleus using ringing recoveries of Swedish birds, to answer questions about geographic migration patterns, age-specific migrations, migration speeds and synchrony of movements. Median migration distance of Swedish Blue Tits was 82 km, with a main autumn direction in the sector between S and W (large directional scatter). Northerly and southerly populations did not differ in migration directions or distances, suggesting chain migration to be the general pattern. A larger proportion of adult Blue Tits remained near the breeding grounds during winter than was the case for juveniles. Some of the migrating birds (17%) seemed not to return in spring but stayed to breed closer to the winter area. Swedish Blue Tits show an exceptionally slow migration speed (median 13 km/day), among the slowest speeds recorded for any migrant bird. The Blue Tit represents an extreme case of diffuse, short and slow bird migration.     

Flight by night or day? Optimal daily timing of bird migration

Many migratory bird species fly mainly during the night (nocturnal migrants), others during daytime (diurnal migrants) and still others during both night and day. Need to forage during the day, atmospheric structure, predator avoidance and orientation conditions have been proposed as explanations for the widespread occurrence of nocturnal migration. However, the general principles that determine the basic nocturnal-diurnal variation in flight habits are poorly known. In the present study optimal timing of migratory flights, giving the minimum total duration of the migratory journey, is evaluated in a schematic way in relation to ecological conditions for energy gain in foraging and for energy costs in flight. There exists a strong and fundamental advantage of flying by night because foraging time is maximized and energy deposition can take place on days immediately after and prior to the nocturnal flights. The increase in migration speed by nocturnal compared with diurnal migration will be largest for birds with low flight costs and high energy deposition rates. Diurnal migration will be optimal if it is associated with efficient energy gain immediately after a migratory flight because suitable stopover/foraging places have been located during the flight or if energy losses during flight are substantially reduced by thermal soaring and/or by fly-and-forage migration. A strategy of combined diurnal and nocturnal migration may be optimal when birds migrate across regions with relatively poor conditions for energy deposition (not only severe but also soft barriers). Predictions about variable timing of migratory flights depending on changing foraging and environmental conditions along the migration route may be tested for individual birds by analysing satellite tracking results with respect to daily travel routines in different regions. Documenting and understanding the adaptive variability in daily travel schedules among migrating animals constitute a fascinating challenge for future research.     

Migration comes of age

The Age of Migration has been uniquely successful as a student text on international migration, not only filling a niche but defining the field. This review article documents the evolution of the five editions of the book in terms of its core structure and enlarging scope. In a more critical vein, I note some omissions and potential shortcomings, while acknowledging the subjectivity of my perspective. More attention could have been given to the mobilities paradigm, transnationalism, internal and return migration, and quantitative analyses. Nevertheless, this book has done more than any other to ensure that the academic study of migration now occupies a central place in the social sciences.     

Regional wind patterns likely shape a seasonal migration detour

Migrating animals should optimise time and energy use when migrating, travelling directly to their destination. Detours from the most direct route may arise however because of barriers and weather conditions. Identifying how such situations arise from variable weather conditions is crucial to understand population response in the light of increased anthropogenic climate change. Here we used light-level geolocators to follow Cyprus wheatears for their full annual cycle in two separate years migrating between Cyprus, over the Mediterranean and the Sahara to winter in north–east sub-Saharan Africa. We predicted that any route detours would be related to wind conditions experienced during migration. We found that spring migration for all birds included an eastern detour, whilst autumn migrations were direct across the Sahara. The direct autumn migration was likely a consequence of consistent tail-winds, whilst the eastern detour in spring is likely to be more efficient given the wind conditions which are against a direct route. Such variable migration routes shaped by coincidence with prevailing winds are probably common suggesting that some birds may be able to adapt to future changes in wind conditions.     

Where the wild birds go: explaining the differences in migratory destinations across terrestrial bird species

Given their large movement capacities, migratory birds have in principle a wide range of possible geographical locations for their breeding and non‐breeding destinations, yet each species migrates between consistent breeding and non‐breeding ranges. In this study, we use a macroecological approach to search for the general factors explaining the location of the seasonal ranges of migratory bird species across the globe. We develop a null model to test the hypotheses that access to resources, geographical distance, tracking of temperature, and habitat conditions (separately as well as considered together) have a major influence in the location of species’ migratory destinations, once each species’ geographical constraints are taken into account. Our results provide evidence for a trade‐off between costs associated with distance travelled and gains in terms of better access to resources. We also provide strong support to the hypotheses that all factors tested, with the exception of habitat, have a strong and additive effect on the global geography of bird migration. Indeed, our results indicate that species’ contemporary migratory destinations (i.e. the combination of their breeding and non‐breeding ranges) are such that they allow species to track a temperature regime throughout the year, to escape local competition and reach areas with better access to resources, and to minimise the spatial distance travelled, within the limitations imposed by the geographical location of each species. Our study thus sheds light on the mechanisms underpinning bird migration and provides a strong basis for predicting how migratory species will respond to future change.     

Danger management and the seasonal adjustment of migratory speed by sandpipers

The behavior of migrating birds is governed by time‐, energy‐ and danger‐minimizing strategies. The adjustment of migration speed (i.e. the rate at which distance is covered during a migration) is a behavioral tactic that might contribute to these strategic goals. Shorter stopovers and greater fuel loads increase migration speed, but both require more intensive foraging at stopovers, making migrants more vulnerable to predators. A simple numerical model shows how seasonal alterations in migration speed can lower the exposure of western sandpipers to peregrine falcons, their most important predator. The ‘caution–speed–caution’ pattern of higher migration speed in the mid‐passage period, observed in earlier work, requires that the intensive foraging necessary heightens vulnerability, and that migrants are exposed to both migrant predators as well as predators resident at migratory stopovers.     

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

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

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

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

A fence barrier method of leading edge cell capture for explorative biochemical research

The scratch or wound-healing assay is used ubiquitously for investigating re-epithelialisation and has already revealed the importance of cells comprising the leading edge of healing epithelial wounds. However it is currently limited to studying the effect of known biochemical agents on the tissue of choice. Here we present an adaptation that extends the utility of this model to encompass the collection of cells from the leading edge of migrating epithelial sheets making available explorative biochemical analyses. The method is scalable and does not require expensive apparatus, making it suitable for large and small laboratories alike. We detail the application of our method and exemplify proof of principle data derived from primary human keratinocyte cultures.     

Considerations of varied thermoregulatory expressions in migration theory

Optimal migration theory has been used for three decades to generate predictions of stopover behavior and understand migration ecology. Yet, to date, there have been no attempts to understand the impacts of thermoregulation on migration theory predictions of stopover behavior. Though most migrants are homeothermic, a diverse group of migrants from bats to hummingbirds and warblers make use of some degree of heterothermy. We consider how thermoregulation influences stopover fuel deposition rates, and thus alters optimal migration theory predictions of stopover behavior using a hypothetical migratory bat as a model organism. We update the analytical models of optimal migration theory by considering scenarios of fixed metabolic rate (the current assumption of optimal migration theory) and three different mass-specific metabolic rates including homeothermy, shallow torpor heterothermy and deep torpor heterothermy. Our results predict that heterotherms will make shorter stopovers, have a decreased departure fuel load, and reduce the overall time and energy costs associated with stopovers relative to homeotherms, highlighting that thermoregulation can drastically influence stopover behavior and ultimately play a critical role in population level patterns of migration.     

The annual cycle of a trans-equatorial Eurasian-African passerine migrant: different spatio-temporal strategies for autumn and spring migration

The small size of the billions of migrating songbirds commuting between temperate breeding sites and the tropics has long prevented the study of the largest part of their annual cycle outside the breeding grounds. Using light-level loggers (geolocators), we recorded the entire annual migratory cycle of the red-backed shrike Lanius collurio, a trans-equatorial Eurasian-African passerine migrant. We tested differences between autumn and spring migration for nine individuals. Duration of migration between breeding and winter sites was significantly longer in autumn (average 96 days) when compared with spring (63 days). This difference was explained by much longer staging periods during autumn (71 days) than spring (9 days). Between staging periods, the birds travelled faster during autumn (356 km d(-1)) than during spring (233 km d(-1)). All birds made a protracted stop (53 days) in Sahelian sub-Sahara on southbound migration. The birds performed a distinct loop migration (22 000 km) where spring distance, including a detour across the Arabian Peninsula, exceeded the autumn distance by 22 per cent. Geographical scatter between routes was particularly narrow in spring, with navigational convergence towards the crossing point from Africa to the Arabian Peninsula. Temporal variation between individuals was relatively constant, while different individuals tended to be consistently early or late at different departure/arrival occasions during the annual cycle. These results demonstrate the existence of fundamentally different spatio-temporal migration strategies used by the birds during autumn and spring migration, and that songbirds may rely on distinct staging areas for completion of their annual cycle, suggesting more sophisticated endogenous control mechanisms than merely clock-and-compass guidance among terrestrial solitary migrants. After a century with metal-ringing, year-round tracking of long-distance migratory songbirds promises further insights into bird migration.     

Revealing patterns of nocturnal migration using the European weather radar network

Nocturnal avian migration flyways remain an elusive concept, as we have largely lacked methods to map their full extent. We used the network of European weather radars to investigate nocturnal bird movements at the scale of the European flyway. We mapped the main migration directions and showed the intensity of movement across part of Europe by extracting biological information from 70 weather radar stations from northern Scandinavia to Portugal, during the autumn migration season of 2016. On average, over the 20 nights and all sites, 389 birds passed per 1 km transect per hour. The night with highest migration intensity showed an average of 1621 birds km^–1 h^–1 passing the radar stations, but there was considerable geographical and temporal variation in migration intensity. The highest intensity of migration was seen in central France. The overall migration directions showed strong southwest components. Migration dynamics were strongly related to synoptic wind conditions. A wind‐related mass migration event occurred immediately after a change in wind conditions, but quickly diminished even when supporting winds continued to prevail. This first continental‐scale study using the European network of weather radars demonstrates the wealth of information available and its potential for investigating large‐scale bird movements, with consequences for ecosystem function, nutrient transfer, human and livestock health, and civil and military aviation.     

Animal migration: linking models and data beyond taxonomic limits

An international workshop on animal migration was held at the Lorentz Center in Leiden, The Netherlands, 2–6 March 2009, bringing together leading theoreticians and empiricists from the major migratory taxa, aiming at the identification of cutting-edge questions in migration research that cross taxonomic borders.     

Evolutionary aspects of migratory behavior in European warblers

During the past 20 years, European Sylvia warblers have been used for a model study of the control mechanisms of bird migration and of evolutionary aspects of migratory behavior. Endogenous annual rhythms (‘circannual’ rhythms) and photo-period have proved to be the essential internal and external controlling factors. It is unknown whether this basic system, that also controls migration in other bird species, is currently evolutionarily stable or is instead adapting birds to the present slightly changing environmental conditions. Using the Blackcap, the control of partial migration in a bird species was analyzed. Two-way selective breeding experiments demonstrated a large selection response and high heritability values. These experiments have also indicated that a partially migratory population can become either almost completely migratory, or sedentary, in two to five generations. Hence, genetic influences are very important and presumably dominant over environmental factors in the expression of migratory or sedentary behavior. The large selection response implies a strikingly high evolutionary potential with respect to strong selection pressures. Further, in the Blackcap, migratory orientation behavior (in addition to migratory activity) was immediately transmitted into a F₁-generation when a cross-breeding experiment was performed using birds from a migratory and a resident population. The hybrids displayed their migratory activity along an axis that is used by their migratory parents. Finally, a rapidly developing novel migratory habit (new migratory direction to new wintering areas) in the Blackcap is discussed with respect to a positive feedback-mechanism, possibly including a series of advantages leading to above-average fitness.     

标记回收法确认我国北方地区草地螟的迁飞

为了证实草地螟Loxostege sticticalis L.（鳞翅目： 螟蛾科）由我国华北越冬区向东北迁飞危害的假说并提供直接的证据, 采用网捕成虫喷雾染料标记释放回收的方法,于2009年越冬代草地螟成虫盛发期间（6月1-4日）分别在内蒙古武川县（41.07°N, 111.42°E）和察右中旗（41.13°N, 112.58°E）的6个标放点共标记当地越冬代草地螟成虫约181万头并自然释放。标记释放后, 在东北、 西北和华北11个草地螟常发省（市、 区）组织各级监测预警部门利用虫情测报灯、 捕虫网和性诱剂诱捕器等多种器械进行回收, 先后于2009年6月7-29日在内蒙古科右中旗、 乌兰浩特、 扎兰屯分别回收到由察右中旗标记释放的成虫9头, 在辽宁省北票市回收到由武川县标记释放的成虫1头。标记释放点至回收点直线距离为725~1 117 km。这是迄今为止世界上确认的草地螟最远迁飞距离。这些证据不仅证实了我国越冬代草地螟成虫能从华北迁往东北危害, 而且为进一步了解草地螟的种群动态规律, 改善草地螟的预测预报技术提供了重要的科学依据。     

Spring phenology of ecological productivity contributes to the use of looped migration strategies by birds

Migration is a common strategy used by birds that breed in seasonal environments. The patterns and determinants of migration routes, however, remain poorly understood. Recent empirical analyses have demonstrated that the locations of two North America migration flyways (eastern and western) shift seasonally, reflecting the influence of looped migration strategies. For the eastern but not western flyway, seasonal variation in atmospheric circulation has been identified as an explanation. Here, we test an alternative explanation based on the phenology of ecological productivity, which may be of greater relevance in western North America, where phenology is more broadly dictated by elevation. Migrants in the western flyway selected lower-elevation spring routes that were wetter, greener and more productive, and higher-elevation autumn routes that were less green and less productive, but probably more direct. Migrants in the eastern flyway showed little season variation but maintained associations with maximum regional greenness. Our findings suggest the annual phenology of ecological productivity is associated with en route timing in both flyways, and the spring phenology of ecological productivity contributes to the use of looped strategies in the western flyway. This fine-tuned spatial synchronization may be disrupted when changing climate induces a mismatch between food availability and needs.     

The contested politics of naming in Europe’s “refugee crisis”

How we label, categorize and, in turn, differentiate between those on the move, for example those who cross the Mediterranean on unseaworthy boats, has enormous implications on the kind of legal and moral obligations receiving states and societies feel towards them. This is perhaps even more salient nowadays in the context of the process of reform of the global governance of migration initiated with the 2016 New York Declaration for Refugees and Migrants.     

Speciation with gene flow could be common

The likelihood of speciation in the face of homogenizing gene flow (i.e. without complete geographical isolation) is one of the most debated topics in evolutionary biology. Demonstrating this phenonemon is hampered by the difficulty of isolating the effects of time since population divergence vs. gene flow on levels of molecular genetic differentiation. For example, weak genetic differentiation between taxa could be due to recent divergence, gene flow, or a combination of these factors. Nonetheless, a number of convincing examples of speciation with gene flow have recently emerged, owing in part to the development of new analytical methods designed to estimate gene flow specifically. A recent example of speciation with gene flow in salamanders (Niemiller et al. 2008) further advances our understanding of this phenonemon, by showing that gene flow between cave and spring salamanders was ongoing during speciation, rather than having occurred after a long period of allopatric divergence. Future work on the ecological and genetic factors reducing gene flow will likely increase our understanding of the conditions that faciliate divergence in the face of gene flow.