Dispersal costs can be classified into energetic, time, risk and opportunity costs and may be levied directly or deferred during departure, transfer and settlement. They may equally be incurred during life stages before the actual dispersal event through investments in special morphologies. Because costs will eventually determine the performance of dispersing individuals and the evolution of dispersal, we here provide an extensive review on the different cost types that occur during dispersal in a wide array of organisms, ranging from micro-organisms to plants, invertebrates and vertebrates. In general, costs of transfer have been more widely documented in actively dispersing organisms, in contrast to a greater focus on costs during departure and settlement in plants and animals with a passive transfer phase. Costs related to the development of specific dispersal attributes appear to be much more prominent than previously accepted. Because costs induce trade-offs, they give rise to covariation between dispersal and other life-history traits at different scales of organismal organisation. The consequences of (i) the presence and magnitude of different costs during different phases of the dispersal process, and (ii) their internal organisation through covariation with other life-history traits, are synthesised with respect to potential consequences for species conservation and the need for development of a new generation of spatial simulation models.

1. Light-level geolocator tags use ambient light recordings to estimate the whereabouts of an individual over the time it carried the device. Over the past decade, these tags have emerged as an important tool and have been used extensively for tracking animal migrations, most commonly small birds.

1. Currently, the deployment of tracking devices is one of the most frequently used approaches to study movement ecology of birds. Recent miniaturization of lightlevel geolocators enabled studying small bird species whose migratory patterns were widely unknown. However, geolocators may reduce vital rates in tagged birds and may bias obtained movement data.2. There is a need for a thorough assessment of the potential tag effects on small birds, as previous meta-analyses did not evaluate unpublished data and impact of multiple life-history traits, focused mainly on large species and the number of published studies tagging small birds has increased substantially.

In many migratory birds, males precede females during migration and arrival at the breeding sites. Three proximate mechanisms are proposed to explain this phenomenon of protandry: males 1) winter closer to breeding sites, 2) start spring migration earlier, and/or 3) migrate faster than females. So far, the relative contribution of these mechanisms to protandry is unknown. The present study investigated the importance of each of the 3 proximate mechanisms of protandry for a songbird migrant wintering in Africa, the northern wheatear (Oenanthe oenanthe). Two subspecies co-occur in Europe on migration, of which the leucorhoa northern wheatears breeding from Iceland to Canada have to cross the North Atlantic, whereas the nominate form breeding in Europe does not face any significant sea barrier. We show that the leucorhoa subspecies had a significantly higher degree of protandry at stopover sites across Europe than the oenanthe subspecies (−6 vs. −2 days). Leucorhoa northern wheatear's higher degree of protandry was associated with a larger age effect, in which old males preceded young males, and greater sex-specific differences in wing shape and refueling yielding higher migration speeds in males than females. In oenanthe northern wheatears, light-level geolocators revealed that males did not winter closer to the breeding sites or migrate faster than females, but initiated spring migration earlier. Our results demonstrate that the significance of the mechanisms causing protandry can differ between related taxa and highlight the importance of the advancement in male arrival date with age as a potential factor shaping the degree of protandry.

Aim Knowledge of broad‐scale biogeographical patterns of animal migration is important for understanding ecological drivers of migratory behaviours. Here, we present a flyway‐scale assessment of the spatial structure and seasonal dynamics of the Afro‐Palaearctic bird migration system and explore how phenology of the environment guides long‐distance migration. Location Europe and Africa. Time period 2009–2017. Major taxa studied Birds. Methods We compiled an individual‐based dataset comprising 23 passerine and near‐passerine species of 55 European breeding populations, in which a total of 564 individuals were tracked during migration between Europe and sub‐Saharan Africa. In addition, we used remotely sensed primary productivity data (the normalized difference vegetation index) to estimate the timing of vegetation green‐up in spring and senescence in autumn across Europe. First, we described how individual breeding and non‐breeding sites and the migratory flyways link geographically. Second, we examined how the timing of migration along the two major Afro‐Palaearctic flyways is tuned with vegetation phenology at the breeding sites. Results We found the longitudes of individual breeding and non‐breeding sites to be related in a strongly positive manner, whereas the latitudes of breeding and non‐breeding sites were related negatively. In autumn, migration commenced ahead of vegetation senescence, and the timing of migration was 5–7 days earlier along the Western flyway compared with the Eastern flyway. In spring, the time of arrival at breeding sites was c. 1.5 days later for each degree northwards and 6–7 days later along the Eastern compared with the Western flyway, reflecting the later spring green‐up at higher latitudes and more eastern longitudes. Main conclusions Migration of the Afro‐Palaearctic landbirds follows a longitudinally parallel leapfrog migration pattern, whereby migrants track vegetation green‐up in spring but depart before vegetation senescence in autumn. The degree of continentality along migration routes and at the breeding sites of the birds influences the timing of migration on a broad scale.