Sonar broadcasts are followed by echoes at different delays from objects at different distances. When broadcasts are emitted rapidly in cluttered surroundings, echo streams from successive broadcasts overlap and cause ambiguity in matching echoes to corresponding broadcasts. To identify reactions to ambiguity in clutter, echolocating bats that emit multiple-harmonic FM sounds were trained to fly into a dense, extended array of obstacles (multiple rows of vertically hanging chains) while the sonar sounds the bat emitted were recorded with a miniature radio microphone carried by the bat. Flight paths were reconstructed from thermal-infrared video recordings. Successive rows of chains extended more than 6 m in depth, so each broadcast was followed by a series of echoes from multiple rows of chains that lasted up to 40 ms. Bats emitted sounds in pairs ("strobe groups") at short (20-40 ms) interpulse intervals (IPIs) alternating with longer IPIs (>50 ms). For many short IPIs, the stream of echoes from the first broadcast was still arriving when the second broadcast was emitted. This overlap caused ambiguity about matching echoes with broadcasts. Bats shifted frequencies of the first sound in each strobe group upward and the second sound downward by 3-6 kHz. When overlap and ambiguity ceased, frequency shifts ceased also. Frequency differences were small compared with the total broadcast band, which was 75-80 kHz wide, but the harmonic structure of echoes enhances the differences in spectrograms. Bats could use time-frequency comparisons of echoes with broadcasts to assign echoes to the corresponding broadcasts and thus avoid ambiguity.bat sonar | clutter interference | echo delay | FM sounds

SUMMARYThe echolocation sounds of Japanese CF-FM bats (Rhinolophus ferrumequinum nippon) were measured while the bats pursued a moth (Goniocraspidum pryeri) in a flight chamber. Using a 31-channel microphone array system, we investigated how CF-FM bats adjust pulse direction and beam width according to prey position. During the search and approach phases, the horizontal and vertical beam widths were ±22±5 and ±13±5deg, respectively. When bats entered the terminal phase approximately 1m from a moth, distinctive evasive flight by G. pryeri was sometimes observed. Simultaneously, the bats broadened the beam widths of some emissions in both the horizontal (44% of emitted echolocation pulses) and vertical planes (71%). The expanded beam widths were ±36±7deg (horizontal) and ±30±9deg (vertical). When moths began evasive flight, the tracking accuracy decreased compared with that during the approach phase. However, in 97% of emissions during the terminal phase, the beam width was wider than the misalignment (the angular difference between the pulse and target directions). These findings indicate that bats actively adjust their beam width to retain the moving target within a spatial echolocation window during the final capture stages.

An onboard microphone (Telemike) was developed to examine changes in the basic characteristics of echolocation sounds of small frequency-modulated echolocating bats, Pipistrellus abramus. Using a dual high-speed video camera system, spatiotemporal observations of echolocation characteristics were conducted on bats during a landing flight task in the laboratory. The Telemike allowed us to observe emitted pulses and returning echoes to which the flying bats listened during flight, and the acoustic parameters could be precisely measured without traditional problems such as the directional properties of the recording microphone and the emitted pulse, or traveling loss of the sound in the air. Pulse intensity in bats intending to land exhibited a marked decrease by 30 dB within 2 m of the target wall, and the reduction rate was approximately 6.5 dB per halving of distance. The intensity of echoes returning from the target wall indicated a nearly constant intensity (-42.6 +/- 5.5 dB weaker than the pulse emitted in search phase) within a target distance of 2 m. These findings provide direct evidence that bats adjust pulse intensity to compensate for changes in echo intensity to maintain a constant intensity of the echo returned from the approaching target at an optimal range.

To understand complex sensory-motor behavior related to object perception by echolocating bats, precise measurements are needed for echoes that bats actually listen to during flight. Recordings of echolocation broadcasts were made from flying bats with a miniature light-weight microphone and radio transmitter (Telemike) set at the position of the bat's ears and carried during flights to a landing point on a wall. Telemike recordings confirm that flying horseshoe bats (Rhinolophus ferrumequinum nippon) adjust the frequency of their sonar broadcasts to compensate for echo Doppler shifts. Returning constant frequency echoes were maintained at the bat's reference frequency +/-83 Hz during flight, indicating that the bats compensated for frequency changes with an accuracy equivalent to that at rest. The flying bats simultaneously compensate for increases in echo amplitude as target range becomes shorter. Flying bats thus receive echoes with both stabilized frequencies and stabilized amplitudes. Although it is widely understood that Doppler-shift frequency compensation facilitates detection of fluttering insects, approaches to a landing do not involve fluttering objects. Combined frequency and amplitude compensation may instead be for optimization of successive frequency modulated echoes for target range estimation to control approach and landing.

When seeing or listening to an object, we aim our attention toward it. While capturing prey, many animal species focus their visual or acoustic attention toward the prey. However, for multiple prey items, the direction and timing of attention for effective foraging remain unknown. In this study, we adopted both experimental and mathematical methodology with microphone-array measurements and mathematical modeling analysis to quantify the attention of echolocating bats that were repeatedly capturing airborne insects in the field. Here we show that bats select rational flight paths to consecutively capture multiple prey items. Microphone-array measurements showed that bats direct their sonar attention not only to the immediate prey but also to the next prey. In addition, we found that a bat's attention in terms of its flight also aims toward the next prey even when approaching the immediate prey. Numerical simulations revealed a possibility that bats shift their flight attention to control suitable flight paths for consecutive capture. When a bat only aims its flight attention toward its immediate prey, it rarely succeeds in capturing the next prey. These findings indicate that bats gain increased benefit by distributing their attention among multiple targets and planning the future flight path based on additional information of the next prey. These experimental and mathematical studies allowed us to observe the process of decision making by bats during their natural flight dynamics.bat sonar | aerial capture | microphone array | mathematical modeling | flight dynamics S electively focusing attention on a particular target allows us to effectively extract information (1-4). Animals spatially focus their attention toward prey for suitable foraging (5, 6). During prey pursuit, most animals direct their visual attention toward it [e.g., tiger beetle (7), dragonfly (8), and falcon (9)]. Specifically, for example, dragonflies maintain a prey item at a constant retinal position during approaching it so that they steer an interception flight path (interception strategy) (10). However, for multiple prey items, the direction and timing of attention for effective foraging remain unknown.Echolocating bats actively emit sonar signals to obtain surrounding information and adaptively change the characteristics of the emissions depending on the situation (11). Therefore, their attention in terms of the sonar (sonar attention) is characterized as the direction to which bats emit their sonar beams (12-14). When echolocating bats approach an airborne insect, the sonar attention directs toward the prey (12, 13). During natural foraging, the sonar attention of bats alternately shifts from their flying direction to the side (15). Additionally the wild bats are capable of capturing two prey items within the short interval of 1 s (16).We predicted that bats localize multiple prey items by distributing their attention between them and then select an efficient flight path to successively capture the multiple prey. To test this prediction, ...

Flight paths of echolocating Japanese house bats, Pipistrellus abramus, were tracked during insect hunting in a natural setting using a 32-microphone array. The array surrounded the foraging area, locating each bat, and determined the directional aim of the sonar beam. Successive interceptions, indicated by feeding "buzzes" and post-buzz pauses, occurred singly at intervals from over 20 s down to multiple interceptions at 2-3 s intervals. Bats flew on looping, curved paths. Turning radius tightened as rate of interceptions increased, keeping the bat in a smaller area of higher capture density. Broadcast beams shifted direction during search, often alternating between the direction of flight and another direction where, moments later, the next interception would occur. Broadcasts also shifted direction between the current target and the next target. Bats time-share biosonar attention between objects by alternating acoustic gaze. During search, most interpulse intervals (IPIs) were 70-120 ms, but bats interspersed long IPIs up to 200 ms when the rate of interception was low and flight paths followed broad curves. Mathematical modeling of search paths demonstrated that circular flight-paths with occasional long IPIs would be more effective for target search than either random, correlated random, or linear flights.

We examined the intra-individual variation in resting frequency of the constant-frequency component of the second harmonic of the pulse (F (rest)) over 4 years in a laboratory colony of the Taiwanese leaf-nosed bat (Hipposideros terasensis). Patterns of change in F (rest) were observed when individuals were added to or removed from the colony so that we investigated whether F (rest) was affected by neighboring colony members. F (rest) of each bat continually showed a long-term gradual change throughout the year, and all bats in the colony increased or decreased their F (rest) in the same direction as a group non-seasonally. The greatest short-term changes were observed when new bats with a relatively low F (rest) joined the colony and F (rest) of new bats converged with those of the original colony members around 8 -16 days after their introduction. Conversely, a single individual showed sudden short-term decrease in F (rest) after its isolation from other colony members. These findings strongly indicate that F (rest) is flexible according to the presence of neighboring conspecific bats. We suggest that the audio-vocal feedback for conspecific pulses appears to be involved in the short- or long-term intra-individual variation in F (rest) other than factors previously thought such as age or season.

Biosonar behavior was examined in Taiwanese leaf-nosed bats (Hipposideros terasensis; CF-FM bats) during flight. Echolocation sounds were recorded using a telemetry microphone mounted on the bat's head. Flight speed and three-dimensional trajectory of the bat were reconstructed from images taken with a dual high-speed video camera system. Bats were observed to change the intensity and emission rate of pulses depending on the distance from the landing site. Frequencies of the dominant second harmonic constant frequency component (CF2) of calls estimated from the bats' flight speed agreed strongly with observed values. Taiwanese leaf-nosed bats changed CF2 frequencies depending on flight speed, which caused the CF2 frequencies of the Doppler-shifted echoes to remain constant. Pulse frequencies were also estimated using echoes returning directly ahead of the bat and from its sides for two different flight conditions: landing and U-turn. Bats in flight may periodically alter their attended angles from the front to the side when emitting echolocation pulses.