Abstract Hundreds of millions of birds reach the Mediterranean islands or Mediterranean coast of Europe every spring after having crossed the Sahara Desert and the Mediterranean Sea. Using data from three small insular stopover sites, we calculated body mass without fuel for 18 trans-Saharan passerine migrants. We subsequently used arrival fuel loads coupled with potential flight range estimates to assess the percentage of birds that are forced to perform an obligatory stopover after crossing the Mediterranean Sea due to fuel depletion. Average arrival fuel loads were among the lowest ever recorded in the Mediterranean region and minimum body mass values recorded for several species were lower than any other individual value reported. The percentage of birds that needed to replenish their energy stores before resuming their northward migration journey varied from 0% to 50% depending on the species and locality studied. Based on conservative estimates at least 180 million birds of our study species are expected to migrate through Greece, 14% of which would not be able to resume their migration without refueling. The significance of small islands and coastal sites in the Mediterranean as obligatory refuelling sites is discussed and their conservation value for migratory birds is highlighted under the perspective of climate change.

Abstract This is submitted as an introduction to the special collection on, “Arctic Migrations in a Changing World”.

Abstract Monarch butterfly wing morphology varies substantially throughout their global range, both between resident and migratory populations and also within the migratory North American population. Here, we use a dataset comprising more than 1800 North American individuals collected between 1878-2017 to characterize the factors shaping continent-wide patterns of wing morphological variation. North American overwintering butterflies have forewings that are approximately 4.4% larger than those collected in summer breeding areas. Monarchs overwintering in Mexico have forewings that are approximately 1.8% larger than monarchs overwintering in California, conducive to the idea that migration distance is positively correlated with wing area. We find evidence for a latitudinal cline within North America, such that butterflies collected at higher latitudes have significantly larger and more elongated forewings. We also find a significant increase of approximately 4.9% in forewing area between 1878-2017, but no difference through time in wing elongation. This result is corroborated by a reanalysis of a recently published dataset of more than 600 butterflies from Mexican overwintering sites. We discuss possible reasons for this increase in wing size through time, including northward shifts in the monarch’s breeding range and changes in relative abundance of milkweed hosts, and present experimental data addressing the influence of larval host plant on adult wing morphology. Our analysis suggests that (1) migration is indeed an important selective force for monarch wing morphology; (2) wing size has increased through time in North America; (3) factors such as host plant identity must be considered to fully understand monarch wing morphological variation.

Abstract Conservation efforts directed at population declines for migratory animals must consider threats occurring at different stages often separated by vast distances. Furthermore, connectivity between populations and fidelity of individuals to specific habitats during the annual cycle are also important considerations. Avian aerial insectivores are experiencing steep population declines in North America, and those declines may be driven, in part, by conditions on the wintering grounds. Here, using geolocators (2 species; 4 individuals) and stable isotope (δ2H, δ13C and δ15N) measurements of feathers (3 species; 841 individuals), we identified approximate winter areas, and assessed migratory connectivity and among-year winter habitat fidelity for three aerial insectivores (Bank Swallow Riparia riparia, Barn Swallow Hirundo rustica and Cliff Swallow Petrochelidon pyrrhonota) that breed in northeastern North America. All three species of swallows are declining in this region. Our results, largely from the stable isotope analysis, suggest that these species likely winter throughout the Cerrado, La Plata Basin, and the Pampas, in South America. These most likely areas were similar among years (2013-2016) for Bank and Cliff Swallows, but varied for Barn Swallows (2014-2016). We found weak migratory connectivity for all three species, and, with one exception, weak habitat fidelity among years for individuals. For individual Barn Swallows captured in two or more years, we found high repeatability in δ13C values, suggesting some fidelity to similar habitats among years. The most likely wintering areas for these species coincide with large areas of South America experiencing high rates of land-use change.

Abstract The Chestnut-collared Longspur (Calcarius ornatus) is one of five grassland songbirds, endemic within North America, with populations that have declined >65% since the 1960s. These species breed and winter in the northern and southern Great Plains, respectively. Identifying migration routes, wintering sites, and the timing of their habitat use is key for understanding the relative magnitude of threats across the annual cycle and effectively targeting habitats for conservation. We tracked migratory movements of seven Chestnut-collared Longspurs with light-level geolocators deployed in Canada. Individuals wintered up to 112-1,200km apart. All followed the Central Flyway, circumvented high-elevation terrain, and traveled east of the breeding location. Unlike most songbirds, the durations of spring and fall migrations were similar; on average 42 ± 7d and 41 ± 5d during fall and spring migrations, respectively, for an approximately 2,000km migration; this highlights the need to better understand habitat requirements during migration for grassland songbirds. Using geospatial habitat data, we assessed winter distribution overlap with four other endemic grassland songbirds; wintering range overlapped 63-99%. Future studies should use more precise devices (e.g., archival GPS units), programmed for data collection dates from this study, to identify specific migratory sites for better conserving this and associated grassland species.

Abstract The migration of monarch butterflies (Danaus plexippus) in North America has a number of parallels with long-distance bird migration, including the fact that migratory populations of monarchs have larger and more elongated forewings than residents. These characteristics likely serve to optimize flight performance in monarchs, as they also do with birds. A question that has rarely been addressed thus far in birds or monarchs is if and how wing characteristics vary within a migration season. Individuals with superior flight performance should migrate quickly, and/or with minimal stopovers, and these individuals should be at the forefront of the migratory cohort. Conversely, individuals with poor flight performance and/or low endurance would be more likely to fall behind, and these would comprise the latest migrants. Here we examined how the wing morphology of migrating monarchs varies to determine if wing characteristics of early migrants differ from late migrants. We measured forewing area, elongation (length/width), and redness, which has been shown to predict flight endurance in monarchs. Based on a collection of 75 monarchs made one entire season (fall 2010), results showed that the earliest migrants (n = 20) in this cohort had significantly redder and more elongated forewings than the latest migrants (n = 17). There was also a non-significant tendency for early migrants to have larger forewing areas. These results suggest that the pace of migration in monarchs is at least partly dependent on the properties of their wings. Moreover, these data also raise a number of questions about the ultimate fate of monarchs that fall behind

The winter distribution of many migratory birds wintering in tropical Africa is poorly known. After the crossing of the Sahara Desert, some long-distance migrants typically stay in the Sahel zone for an extended period before continuing migration to their main wintering areas south of the equator. Here we show how two thrush nightingales ( Luscinia luscinia ) fitted with light-level geolocators, after a six to seven week long stay in the Sahel zone of Sudan, moved to an intermediate area in northern Kenya for a month-long stay before continuing to their final wintering areas in southern Africa. These data indicate that thrush nightingales may use three consecutive wintering sites during their stay in Africa. The migratory movements in Africa between wintering sites are well-coordinated with high precipitation in these areas, suggesting that thrush nightingales track peaks of insect abundance occurring after rains. This three-stage wintering strategy has, to our knowledge, previously not been described, and shows that long-distance migrants can have complex wintering behaviour.

Abstract This chapter presents some basic principles of locomotion and various migration strategies that result from them. As an animal accumulates fuel for migration, it actually accumulates potential migration distance, which typically has diminishing returns to added fuel mass. Depending on the mode of locomotion – running, flying, or swimming – the exact shape of this range function for migration distance may vary. Paired with assumptions about the immediate selection force acting on the migration episode (e.g., minimising energy cost, time, or mortality risk), the range curve can be used to derive optimal strategies and behaviours, including optimal fuel loads, migration step length, detours, and travel speeds. The chapter also discusses problems of drift, compensation, and altitude/depth associated with travel in a medium (air, water) that is itself in motion. Future theoretical work on optimal migration strategies will need to take into account variation in the foraging strategies of migrants and incorporate more precise estimates of individual mortality and fitness.

Abstract Animal movement is a general strategy for dealing with environmental variability, but takes on a variety of forms according to life history constraints and environmental covariates. This chapter contrasts migration with less regular nomadic movements, and invokes the niche concept for linking the diversity of movement patterns to the resource environment and the mechanisms of movement ecology. The statistical properties of the resource environment are key to understanding the merits of movement strategies in different environments, and particular emphasis is put on dimensions and scales of resource variability and predictability in space and time. While migration is frequently seen as an adaptation to avoiding severe conditions and exploiting predictable spatiotemporal variation in resource abundance, nomadism can be seen as an adaptation to variability and unpredictability rather than severity. An exploratory exposition illustrates how analysis of space-time data on resources can be helpful for explaining diversity of movement patterns.