Bats Use Path Integration Rather Than Acoustic Flow to Assess Flight Distance along Flyways

Navigation can be achieved using different strategies from simple beaconing to complex map-based movement [1-4]. Bats display remarkable navigation capabilities, ranging from nightly commutes of several kilometers and up to seasonal migrations over thousands of kilometers [5]. Many bats have been suggested to fly along fixed routes termed "flyways," when flying from their roost to their foraging sites [6]. Flyways commonly stretch along linear landscape elements such as tree lines, hedges, or rivers [7]. When flying along a flyway, bats must estimate the distance they have traveled in order to determine when to turn. This can be especially challenging when moving along a repetitive landscape. Some bats, like Kuhl's pipistrelles, which we studied here, have limited vision [8] and were suggested to rely on bio-sonar for navigation. These bats could therefore estimate distance using three main sensory-navigation strategies, all of which we have examined: acoustic flow, acoustic landmarks, or path integration. We trained bats to fly along a linear flyway and land on a platform. We then tested their behavior when the platform was removed under different manipulations, including changing the acoustic flow, moving the start point, and adding wind. We found that bats do not require acoustic flow, which was hypothesized to be important for their navigation [9-15], and that they can perform the task without landmarks. Our results suggest that Kuhl's pipistrelles use internal self-motion cues-also known as path integration-rather than external information to estimate flight distance for at least dozens of meters when navigating along linear flyways.