Steering by echolocation: a paradigm of ecological acoustics.

1. Flights of three big brown bats (Eptesicus fuscus) landing on a hand and catching a suspended mealworm were video analysed. 2. Results were consistent with the bats using the same basic control procedure in the quite different approach tasks--namely keeping tau (r) = kr and tau (a)/tau (r) = k alpha r. Here r is the current distance to the destination; alpha is the angle between the current direction of the destination and the goal direction of final approach (beta min); tau (r) = r/r, tau (alpha) = alpha/alpha; and kr, k alpha r are constants. 3. The bats were each quite consistent on a particular task (hand or mealworm) in the values they used for the control parameters kr, k alpha r and beta min. However, different values were used in the two tasks, which reflected the different behaviour required at the destination. Flights to hand required twisting and landing upside down and approach angle beta min was closer to vertical and kr was smaller and corresponded to decelerating nearly to a stop. In contrast, the mealworms were caught in mid flight and approach angle beta min was shallower and speed of approach was about constant. 4. tau (r) might be registered acoustically by tau (echo-delay) or by tau (echo-intensity). tau (alpha) might be registered by the bat's directional hearing and gravity sense. 5. The bat's learned the tasks easily, suggesting that the control procedure they used in the experiments was part and parcel of the natural skills they had developed in the wild.