Introduction Flight and echolocation are two key behavioral innovations in the evolutionary history of bats (Fenton et al., 1995; Arita and Fenton, 1997; Teeling et al., 2000). Flight behavior provides benefits for long-distance dispersal over geographical barriers, avoidance of terrestrial predators, and niche expansion from the ground to the sky (Speakman, 2001; Jones and Teeling, 2006; Luo et al., 2019a).