Flight and echolocation behaviour of whiskered bats commuting along a hedgerow: range-dependent sonar signal design, Doppler tolerance and evidence for 'acoustic focussing'.

Echolocating bats obtain three-dimensional images of their surroundings in complete darkness by emitting sonar signals and evaluating returning echoes. When flying close to objects, bats risk collision and therefore depend on the accuracy of images--particularly in the perceived distance of obstacles, which is coded by the time delay between call and echo. Yet, during flight, such accuracy is perturbed first because bats call and receive echoes at different positions and second because echoes are modified by Doppler shifts. Certain call designs avoid both sources of ranging error, but only for a limited range of distances [the 'distance of focus' (DOF)]. Here, we show that whiskered bats (Myotis mystacinus) using broadband echolocation calls adjust call design in a range-dependent manner so that nearby obstacles are localised accurately. Such behaviour is adaptive because it reduces collision risk. The bats also reduced call duration to some extent as they approached obstacles so that most returning echoes arrived after they finished calling. This reduction in call duration during the approach to obstacles was neither the only nor the main factor that influenced DOF. Indeed, both duration and bandwidth of calls influenced DOF independently, with lower bandwidths and longer durations giving greater DOF. Our findings give a new perspective on the adaptive significance of echolocation call design in nature and have implications for sonar engineering.