Birds of three worlds: moult migration to high Arctic expands a boreal-temperate flyway to a third biome

Background Knowledge on migration patterns and flyways is a key for understanding the dynamics of migratory populations and evolution of migratory behaviour. Bird migration is usually considered to be movements between breeding and wintering areas, while less attention has been paid to other long-distance movements such as moult migration. Methods We use high-resolution satellite-tracking data from 58 taiga bean geese Anser fabalis fabalis from the years 2019–2020, to study their moult migration during breeding season. We show the moulting sites, estimate the migratory connectivity between the breeding and the moulting sites, and estimate the utilization distributions during moult. We reveal migration routes and compare the length and timing of migration between moult migrants and successful breeders. Results All satellite-tracked non-breeding and unsuccessfully breeding taiga bean geese migrated annually to the island of Novaya Zemlya in the high Arctic for wing moult, meaning that a large part of the population gathers at the moulting sites outside the breeding range annually for approximately three months. Migratory connectivity between breeding and moulting sites was very low (rm = − 0.001, 95% CI − 0.1562–0.2897), indicating that individuals from different breeding grounds mix with each other on the moulting sites. Moult migrants began fall migration later in autumn than successful breeders, and their overall annual migration distance was over twofold compared to the successful breeders. Conclusions Regular moult migration makes the Arctic an equally relevant habitat for the taiga bean goose population as their boreal breeding and temperate wintering grounds, and links ecological communities in these biomes. Moult migration plays an important role in the movement patterns and spatio-temporal distribution of the population. Low migratory connectivity between breeding and moulting sites can potentially contribute to the gene flow within the population. Moult migration to the high Arctic exposes the population to the rapid impacts of global warming to Arctic ecosystems. Additionally, Novaya Zemlya holds radioactive contaminants from various sources, which might still pose a threat to moult migrants. Generally, these results show that moult migration may essentially contribute to the way we should consider bird migration and migratory flyways.

Evolution of moult-migration is directly linked to aridity of the breeding grounds in North American passerines

To avoid energy allocation conflicts, birds generally separate breeding, migration and moult during the annual cycle. North American passerines typically moult on the breeding grounds prior to autumn migration. However, some have evolved a moult-migration strategy in which they delay moult until stopping over during autumn migration. Rohwer et al . (2005) proposed the ‘push–pull hypothesis' as an explanation for the evolution of this moult strategy, but it has not been empirically tested. Poor conditions on the breeding grounds at the end of the summer would push birds to depart prior to moult, while productive stopover locations would pull them. We tested for a relationship between moult-migration and breeding grounds aridity as measured by the normalized difference vegetation index. Our results strongly support the ‘push' aspect of the push–pull hypothesis and indicate that arid breeding grounds, primarily in western North America, would drive species to evolve stopover moult-migration, although this relationship may depend upon migration distance.

Evidence of altitudinal moult-migration in a Central American hummingbird, Amazilia cyanura

For birds, moulting is an energetically costly endeavour (Murphy & King 1991), the timing and location of which may be flexible and governed by local ecological factors (Pyle et al. 2009, Rohwer et al. 2005). Some species or individuals may pause during long-distance migration, or migrate specifically to moult (Greenberg et al. 1974, Pyle et al. 2009, Rohwer et al. 2005, 2008). This strategy may be most common when food abundance reaches a nadir at the end of the breeding period, promoting movement to areas where food is more plentiful and the energetic and nutritive demands of moult may be met more suitably (Rohwer et al. 2005). This pattern is exemplified by insectivorous songbirds breeding in temperate, western North America that pause on southward migration to moult amidst the food flush that occurs following heavy rains in the Mexican monsoon region (Pyle et al. 2009, Rohwer et al. 2005, 2009), or more rarely, migrate upslope after breeding to moult in more moist, productive areas at higher elevation (Butler et al. 2002, Greenberg et al. 1974, Rohwer et al. 2008, Steele & McCormick 1995). Such altitudinal migration may be much more common in the Neotropics where many species engage in seasonal shifts in elevation. Hypotheses proposed to explain this behaviour, however, have focused on links made between migration and a principally frugivorous or nectarivorous diet (Levey & Stiles 1992, Stiles 1985, 1988) and not on moult requirements or phenology. Fruit and nectar availability may vary seasonally over an elevational gradient, and birds may migrate in order to track peak abundances (Levey & Stiles 1992, Loiselle & Blake 1991). Hummingbirds may also track arthropod (particularly spider) abundance, but this possibility remains little explored (Cotton 2007, Stiles 1980).