Latitudinal gradients in air density create “invisible topography” at sea level affecting animal flight costs

Regional patterns in wind underpin the low-cost migratory flyways of billions of birds and insects ^1-3 , but how large-scale changes in temperature affect flight is unknown. Flight costs should increase with rising temperatures, because lift decreases as density decreases, whereas weight remains unchanged. The effects of air density on flight costs are well-established in the context of high-altitude movements and migration ^4-7 . Here, we examine the impact of air density on low-flying birds, in relation to seasonal, regional and global changes in temperature. Using multi-sensor loggers, we find that air density was the most important predictor of wingbeat frequency in red-tailed tropicbirds ( Phaethon rubricauda ) breeding year-round in Mauritius. Lower air densities in the Austral summer were associated with a small but significant increase in mean wingbeat frequency, which translated to an estimated 1-2% increase in flight costs. The variation in flight costs increased by an order of magnitude when considered in space, rather than time, with flight costs varying by ≥ 10 % across the tropicbird’s range. Changes in air density can therefore be an important determinant of flight costs even when birds are operating close to sea-level. Indeed, mapping air density at sea-level revealed that global temperature gradients cause effective altitude to vary by >2 km when considered as seasonal averages. This “invisible topography” at sea-level could have influenced the biogeography of flight morphologies and life-history traits.