Echolocating bats prey upon small moving insects in the dark using sophisticated sonar techniques. The direction and directivity pattern of the ultrasound broadcast of these bats are important factors that affect their acoustical field of view, allowing us to investigate how the bats control their acoustic attention (pulse direction) for advanced flight maneuvers. The purpose of this study was to understand the behavioral strategies of acoustical sensing of wild Japanese house bats Pipistrellus abramus in three-dimensional (3D) space during consecutive capture flights. The results showed that when the bats successively captured multiple airborne insects in short time intervals (less than 1.5 s), they maintained not only the immediate prey but also the subsequent one simultaneously within the beam widths of the emitted pulses in both horizontal and vertical planes before capturing the immediate one. This suggests that echolocating bats maintain multiple prey within their acoustical field of view by a single sensing using a wide directional beam while approaching the immediate prey, instead of frequently shifting acoustic attention between multiple prey. We also numerically simulated the bats’ flight trajectories when approaching two prey successively to investigate the relationship between the acoustical field of view and the prey direction for effective consecutive captures. This simulation demonstrated that acoustically viewing both the immediate and the subsequent prey simultaneously increases the success rate of capturing both prey, which is considered to be one of the basic axes of efficient route planning for consecutive capture flight. The bat’s wide sonar beam can incidentally cover multiple prey while the bat forages in an area where the prey density is high. Our findings suggest that the bats then keep future targets within their acoustical field of view for effective foraging. In addition, in both the experimental results and the numerical simulations, the acoustic sensing and flights of the bats showed narrower vertical ranges than horizontal ranges. This suggests that the bats control their acoustic sensing according to different schemes in the horizontal and vertical planes according to their surroundings. These findings suggest that echolocating bats coordinate their control of the acoustical field of view and flight for consecutive captures in 3D space during natural foraging.

Three-dimensional directivity patterns of sonar sounds emitted by Japanese house bats (Pipistrellus abramus) during natural foraging were measured by a 44-channel microphone array. Just before prey capture, the terminal frequency (TF) of emitted sounds decreased, and the beam width (mean ± standard deviation) expanded from 40 ± 10° to 63 ± 9° (horizontal) and from 32 ± 10° to 52 ± 7° (vertical). P. abramus decrease the TF to simultaneously expand the beam width in both the horizontal and vertical planes, while retaining the target within the three-dimensional acoustic field of view at the final stage of capture.

In this study, 3-D flight paths and directivity pattern of the sounds emitted by Pipistrellus abramus during natural foraging were measured by a large scale microphone array system. The results show that the bats approached prey with covering the direction of them within their sonar beam. The means of horizontal and vertical beam widths were 49 deg and 46 deg, respectively. Just before capturing prey, the bats decreased the terminal frequency (TF) of the pulse. Simultaneously, the beam widths were expanded to 64 deg (horizontal) and 57 deg (vertical). We assumed a circular piston model to estimate how much the beam width was changed by decreasing the frequency of emitted pulse. It was found that the observed expansion of the beam width was smaller than those of theoretical estimations. This suggests that the bats decrease the TF of pulse for compensating their beam width narrowed by taking a large bite for the prey. We also measured echolocation calls and flight behavior of Myotis macrodactylus during natural foraging. M. macrodactylus uses FM echolocation pulse which is similar to P. abramus, but they forage for prey above the water surface. We compare echolocation strategies between two FM bats with different foraging habitat.

Aerial-feeding bats actively emit sonar sounds and capture large amounts of airborne insects a night. Microphone-array system allows us to know not only the positions where the bat emits sonar sounds (i.e., 3-D flight path) but how the bats dynamically control the acoustical field of view during searching and approaching target-prey. Here, we show echolocation strategy of bats during natural foraging revealed by the large-scale microphone-array system which covered the horizontal area of approximately 20 m × 20 m. Pipistrellus abramus was found to expand the width of their sonar beams in both horizontal and vertical planes just before the prey-capture. Since the bats emit echolocation pulses at a high rate (i.e., feeding buzz) just before capturing, the capture positions can additionally be measured. Recently, we have investigated the relationship between flight patterns, capture positions, and foraging efficiency of Myotis macrodactylus during natural foraging above the pond. Further investigation from the viewpoint of the optimal foraging would reveal echolocation and flight strategies of the bats to efficiently search and approach prey items. [This research was supported by a Grant-in-Aid for Young Scientists (B) and Scientific Research on Innovative Areas of JSPS, and the JST PRESTO program.]