Echolocating greater horseshoe bats (Rhinolophus ferrumequinum) emit their biosonar pulses nasally, through nostrils surrounded by fleshy appendages (‘noseleaves’) that diffract the outgoing ultrasonic waves. Movements of one noseleaf part, the lancet, were measured in live bats using two synchronized high speed video cameras with 3D stereo reconstruction, and synchronized with pulse emissions recorded by an ultrasonic microphone. During individual broadcasts, the lancet briefly flicks forward (flexion) and is then restored to its original position. This forward motion lasts tens of milliseconds and increases the curvature of the affected noseleaf surfaces. Approximately 90% of the maximum displacements occurred within the duration of individual pulses, with 70% occurring towards the end. Similar lancet motions were not observed between individual pulses in a sequence of broadcasts. Velocities of the lancet motion were too small to induce Doppler shifts of a biologically-meaningful magnitude, but the maximum displacements were significant in comparison with the overall size of the lancet and the ultrasonic wavelengths. Three finite element models were made from micro-CT scans of the noseleaf post mortem to investigate the acoustic effects of lancet displacement. The broadcast beam shapes were found to be altered substantially by the observed small lancet movements. These findings demonstrate that—in addition to the previously described motions of the anterior leaf and the pinna—horseshoe bat biosonar has a third degree of freedom for fast changes that can happen on the time scale of the emitted pulses or the returning echoes and could provide a dynamic mechanism for the encoding of sensory information.
Since many bat species thrive in densely vegetated habitats, echoes from foliage are likely to be of prime importance to the animals’ sensory ecology, be it as clutter that masks prey echoes or as sources of information about the environment. To better understand the characteristics of foliage echoes, a new model for the process that generates these signals has been developed. This model takes leaf size and orientation into account by representing the leaves as circular disks of varying diameter. The two added leaf parameters are of potential importance to the sensory ecology of bats, e.g., with respect to landmark recognition and flight guidance along vegetation contours. The full model is specified by a total of three parameters: leaf density, average leaf size, and average leaf orientation. It assumes that all leaf parameters are independently and identically distributed. Leaf positions were drawn from a uniform probability density function, sizes and orientations each from a Gaussian probability function. The model was found to reproduce the first-order amplitude statistics of measured example echoes and showed time-variant echo properties that depended on foliage parameters. Parameter estimation experiments using lasso regression have demonstrated that a single foliage parameter can be estimated with high accuracy if the other two parameters are known a priori. If only one parameter is known a priori, the other two can still be estimated, but with a reduced accuracy. Lasso regression did not support simultaneous estimation of all three parameters. Nevertheless, these results demonstrate that foliage echoes contain accessible information on foliage type and orientation that could play a role in supporting sensory tasks such as landmark identification and contour following in echolocating bats.
Horseshoe bats emit biosonar pulses through the nostrils and diffract the outgoing ultrasonic pulses with baffles, so-called "noseleaves," that surround the nostrils. The noseleaves have complex static geometries and can furthermore undergo dynamic shape changes during emission of the biosonar pulses. The posterior noseleaf part, the lancet, has been shown to carry out anterior-posterior flicking motions during biosonar emissions with average lancet tip displacements of about 1 mm. Here, the acoustic effects of the interplay between the lancet furrows and shape change (lancet rotation) on the emission beam were investigated using the animated digital models obtained from the noseleaves of greater horseshoe bats (Rhinolophus ferrumequinum). It was found that forward lancet rotations increase the amount of sound energy allocated to secondary amplitude maxima (sidelobes) in the beampattern, but only in the presence of the furrows. The interaction between static and dynamic features can be readily quantified by roughness (standard deviation about local mean) of the amplitude distribution of the beampatterns. This effect goes beyond the static impact of the furrows on the width of the mainlobe. It could allow the bats to send out their pulses through a sequence of qualitatively different beampatterns.
Echolocating bats use ultrasonic pulses to collect information about their environments. Some of this information is encoded at the baffle structures-noseleaves (emission) and pinnae (reception)-that act as interfaces between the bats' biosonar systems and the external world. The baffle beam patterns encode the direction-dependent sensory information as a function of frequency and hence represent a view of the environment. To generate diverse views of the environment, the bats can vary beam patterns by changes to (1) the wavelengths of the pulses or (2) the baffle geometries. Here we compare the variability in sensory information encoded by just the use of frequency or baffle shape dynamics in horseshoe bats. For this, we use digital and physical prototypes of both noseleaf and pinnae. The beam patterns for all prototypes were either measured or numerically predicted. Entropy was used as a measure to compare variability as a measure of sensory information encoding capacity. It was found that new information was acquired as a result of shape dynamics. Furthermore, the overall variability available for information encoding was similar in the case of frequency or shape dynamics. Thus, shape dynamics allows the horseshoe bats to generate diverse views of the environment in the absence of broadband biosonar signals.
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