What makes Alpine swift ascend at twilight? Novel geolocators reveal year-round flight behaviour.

Abstract: Studying individual flight behaviour throughout the year is indispensable to understand the ecology of a bird species. Recent development in technology allows now to track flight behaviour of small long-distance bird migrants throughout its annual cycle. The specific flight behaviour of twilight ascents in birds has been documented in a few studies, but only during a short period of the year, and never quantified on the individual level. It has been suggested that twilight ascents might be a role in orientation and navigation. Previous studies had reported the behaviour only near the breeding site and during migration. We investigated year-round flight behaviour of 34 individual Alpine swifts () of four different populations in relation to twilight ascents. We recorded twilight ascents all around the year and found a twofold higher frequency in ascents during the non-breeding residence phase in Africa compared to all other phases of the year. Dawn ascents were twice as common as dusk ascents and occurred mainly when atmospheric conditions remained stable over a 24-h period. We found no conclusive support that twilight ascents are essential for recalibration of compass cues and landmarks. Data on the wing flapping intensity revealed that high activity at twilight occurred more regularly than the ascents. We therefore conclude that alpine swift generally increase flight activity-also horizontal flight-during the twilight period and we suppose that this increased flight activity, including ascents, might be part of social interactions between individuals.

Significance Statement: Year-round flight altitude tracking with a light-weight multi-sensor tag reveals that Alpine swifts ascend several hundred meters high at twilight regularly. The reason for this behaviour remains unclear and the low-light conditions at this time of the day preclude foraging as a possibility. The frequency and altitude of twilight ascents were highest during the non-breeding period, intermediate during migration and low for active breeders during the breeding phase. We discuss our findings in the context of existing hypotheses on twilight ascent and we propose an additional hypothesis which links twilight ascent with social interaction between flock members. Our study highlights how flight behaviour of individuals of a migratory bird species can be studied even during the sparsely documented non-breeding period.