Search

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.

Although feathers are the unifying characteristic of all birds, our understanding of the causes, mechanisms, patterns and consequences of the feather moult process lags behind that of other major avian life-history phenomena such as reproduction and long-distance migration. Migration, which evolved in many species of the temperate and arctic zones, requires high energy expenditure to endure long-distance journeys. About a third of Western-Palearctic passerines perform long-distance migrations of thousands of kilometres each year using various morphological, physiological, biomechanical, behavioural and life-history adaptations. The need to include the largely non-overlapping breeding, long-distance migration and feather moult processes within the annual cycle imposes a substantial constraint on the time over which the moult process can take place. Here, we review four feather-moult-related adaptations which, likely due to time constraints, evolved among long-distance Western-Palearctic migrants: (i) increased moult speed; (ii) increased overlap between moult and breeding or migration; (iii) decreased extent of plumage moult; and (iv) moult of part or all of the plumage during the over-wintering period in the tropics rather than in the breeding areas. We suggest that long-distance migration shaped the evolution of moult strategies and increased the diversity of these strategies among migratory passerines. In contrast to this variation, all resident passerines in the Western Palearctic moult immediately after breeding by renewing the entire plumage of adults and in some species also juveniles, while in other species juvenile moult is partial. We identify important gaps in our current understanding of the moult process that should be addressed in the future. Notably, previous studies suggested that the ancestral moult strategy is a post-breeding summer moult in the Western Palearctic breeding areas and that moult during the winter evolved due to the scheduling of long-distance migration immediately after breeding. We offer an alternative hypothesis based on the notion of southern ancestry, proposing that the ancestral moult strategy was a complete moult during the 'northern winter' in the Afro-tropical region in these species, for both adults and juveniles. An important aspect of the observed variation in moult strategies relates to their control mechanisms and we suggest that there is insufficient knowledge regarding the physiological mechanisms that are involved, and whether they are genetically fixed or shaped by environmental factors. Finally, research effort is needed on how global climate changes may influence avian annual routines by altering the scheduling of major processes such as long-distance migration and feather moult.