Foliage is very common feature in the habitats of echolocation bats and thus its echoes constitute the major input of bats' sensory systems. Acquiring useful information from vegetation echoes facilitates the bats significantly in the navigation and foraging behaviors. To better understand the foliage echoes, in this dissertation, a computer model was constructed to simulate foliage echoes with following simplifications: approximating leaves as circular disks, leaving out shading effects between leaves, and distributing leaves uniformly in the space. Then one tree can be described with three parameters in the model, leaf radius, orientation, and leaf density, where the first two determine the beampattern of each leaf. Compared with echoes collected from real trees, the simulation echoes are qualitatively accurate, i.e., they match in waveforms and also first-order statistics. Since the ground truth is known in the model, the three parameters were estimated with lasso model by selecting 40 features from each echo. The results have shown that estimation of one parameter with the other two known is usually successful with coefficient of determination close to one, and the classification still has reasonable accuracy when the number of known parameter is reduced to one. Besides, the three simplifications were examined with both experimental and simulation approaches. To assess the acoustic impact of leaf geometry on individual leaves, experiments were carried out by ensonifying leaves from both a single and different species. How the leaves' impulse responses change according to their equivalent radii was investigated. The simulation model of disks fits the experiments done with real leaves within one species and across species reasonably well. Shading effect is found to exist locally when two disks were 25 cm apart and were both in pulse direction. In addition, the inhomogeneous distribution of leaves was introduced by using the branching patterns of L-system. The evaluation of inhomogeneity in echoes produced with two distributions shows that there is always inhomogeneity in echoes, and L-system model does bring more inhomogeneity but not to the same extent as changes in the relative orientation between sonar beam and foliage do. / Ph. D. / Echolocating bats use ultrasonic waves to navigate and forage at night in the forest. They constantly emit pulses and analyze the returning echoes to perceive the surroundings. Foliage echoes are common and important input of their sensory systems, yet what accessible information is contained in foliage echoes for bats is not fully understood. Hence, this dissertation has built an efficient computer model to compute vegetation echoes. To simplify the problem, leaves with various shapes were approximated as circular disks. Besides, every leaf was assumed to be “visible” to sonar, in other words, even if one leaf was shaded by another in the pulse direction, it can still interact with sonar as if the front leaf didn’t exist. Then the leaves were uniformly distributed in the space. With the simplifications above, foliage can be described with three parameters, mean leaf radius, orientation, and leaf density. By varying the three parameters to match features of different trees, a large amount of echoes can be calculated efficiently. Compared with measured echoes from real trees, the simulation echoes are similar with them in terms of waveforms and probability density functions. If producing echoes with two parameters fixed and the third randomly chosen from certain range, the random parameter can be estimated with a linear model, lasso regression model, by extracting features from the echoes as inputs. The estimation is accurate. But if varying one of the two known parameters, the accuracy of estimation is largely reduced. Besides, the three simplifications were examined if they have impact on the simulation results. Impulse responses from leaf specimens were measured with a bio-mimetic sonar head in the anechoic chamber where noise and unwanted reverberations are largely weakened. Experiments were also carried out for two disks of same size by aligning them in the direction of sound emission to quantify shading effect, which shows that shading effect exists locally. Then branching patterns were introduced to the simulation model using L-system that consists of a set of rules to determine how branches grow. The results demonstrates that the simplifications do affect the model accuracy but the influence may be compensated.

Bats are known for their highly capable biosonar systems which make them be able to navigate and forage in dense vegetation. Their biosonar system consists of one emitter (nose or mouth) and two receivers (ears). Some bat species, e.g. in the rhinolophid and hipposiderid families, have complicated pinna motion patterns. It has been shown that these pinna motion patterns fall into two distinct categories: rigid motions and non-rigid motions. In the current work, the pinna of Pratt's leaf-nosed bat (Hipposideros pratti) was used as a biological model system to understand how a sensor could benefit from variability. Hence, the variability in the rigid pinna motions and in the non-rigid pinna motions has been investigated by tracking a dense set of landmarks on the pinna surface with stereo vision. Axis-angle representations have shown that the rigid pinna motions exhibited a large continuous variation with rotation axes covering 180 degrees in azimuth and elevation. Distributions of clusters of the landmarks on the pinna surface have shown that the non-rigid pinna motions fall into at least two subgroups. Besides, the acoustic impact of the rigid pinna motions have been investigated using a biomimetic pinna. Normalized mutual information between the acoustic inputs with different rotation axes has shown that different rotation axes can provide at least 50% new sensory information. These results demonstrate that the variability in the pinna motions is an interesting concept for sensor, and how the bats approach that needs to be further investigated. / Master of Science / Sensors have been developed for a long time, and they can be used to detect the environments and then deliver the required sensing information. There are many different types of sensors, such as vision-based sensors (infrared camera and laser scanner) and sound-based sensors (sonar and radar). Ultrasonic transducers are one of the sound-based sensors, and they are more stable and reliable in environments where smoke or steam is present. Similar to human-made ultrasonic transducers, bats have developed highly capable biosonar systems that consist of one ultrasonic emitter (nose or mouth) and two ultrasonic receivers (ears), and these biosonar systems enable them to fly and hunt in cluttered environments. Some bats, e.g. rhinolophid and hipposiderid bats, have dynamic noseleaves (elaborate baffle shapes surrounding the nostrils) and pinna (outer ear), and these could enhance the sensing abilities of bats. Hence, the purpose of this thesis has been to investigate this variability to improve the human-made sensors by focusing on the dynamic pinna of the bats. It has been shown that bats have two distinct categories of pinna motions: rigid motions which change only the orientation of the pinna, and non-rigid motions which change also the shape of the pinna. However, the variability within the rigid and non-rigid pinna motions has received little attention. Therefore, the present work has investigated the variability in the rigid pinna motions and in the non-rigid pinna motions. Landmark points were placed on the pinna of certain bats and the pinna motions were tracked by high-speed video cameras. The rigid pinna motions exhibit a large continuous variation in where the pinna is orientated during rotation. Distributions of clusters of the landmarks on the pinna have shown that the non-rigid pinna motions fall into at least two subgroups. The acoustic impacts of the rigid pinna motions have been studied by a biomimetic pinna which reproduced the observed range of the rigid pinna motions. Ultrasonic signals mimicking the bats were emitted to be received by the biomimetic pinna. Based on these signals, it has been shown that different rotation axes and even small changes can provide over 50% new sensory information. These findings give engineers a potential way to improve the human-made sensors.

Bats are a paragon of evolutionary success. They rely on parsimonious sensory inputs provided by echolocation, yet are able to master lives in complex environments. The outer ears (pinnae) of bats are intricately shaped receiver baffles that encode sensory information through a diffraction process. In some bat species with particularly sophisticated biosonar systems, such as horseshoe bats (Rhinolophidae), the pinnae are characterized by static as well as dynamic geometrical features. Furthermore, bats from these species can deform their pinnae while the returning ultrasonic waves impinge on them. Hence, these dynamic pinna geometries could be a substrate for novel, dynamic sensory encoding paradigms. In this dissertation, two aspects of this dynamic sensing process were investigated: (i) Do local shape features impact the acoustic effects during dynamic deformation of the bat pinna? and (ii) do these shape deformations provide a substrate for the dynamic encoding of sensory information? For this, a family of simplified biomimetic prototypes has been designed based on obliquely truncated cones manufactured from sheets of isobutyl rubber. These prototypes were augmented with biomimetic local shape features as well as with a parsimonious deformation mechanism based on a single linear actuator. An automated setup for the acoustic characterization of the time-variant prototype shapes has been devised and used to characterize the acoustic responses of the prototypes as a function of direction. It was found that the effects of local shape features did interact with each other and with the deformation of the overall shape. The impact of the local features was larger for bent than for upright shape configurations. Although the tested devices were much simpler than actual bat pinnae, they were able to reproduce numerical beampattern predictions that have been obtained for deforming horseshoe bat pinnae in a qualitative fashion. The dynamically deformable biomimetic pinna shapes were estimated to increase the sensory encoding capacity of the device by unit[80]{%} information when compared to static baffles. To arrive at this estimate, spectral clustering was used to break up the direction- and deformation-depended device transfer function into a discrete signal alphabet. For this alphabet, we could estimate the joint signal entropy across a bending cycle as a measure for sensory coding capacity. The results presented in this thesis suggest that bat biosonar posses unique dynamic sensing abilities which have no equivalent in man-made technologies. Sensing paradigms derived from bat biosonar could hence inspire new deformable wave-diffracting structures for the advancement in sensor technology. / Ph. D.

Research investigating the behavioral and physiological responses of bats to echoes typically includes analysis of acoustic signals from microphones and/or microphone arrays, using time difference of arrival (TDOA) between array elements or the microphones to locate flying bats (azimuth and elevation). This has provided insight into transmission adaptations with respect to target distance, clutter, and interference. Microphones recording transmitted signals and echoes near a stationary bat provide sound pressure as a function of time but no directional information. This dissertation introduces spatial audio techniques to bat biosonar studies as a complementary method to the current TDOA based acoustical study methods. This work proposes a couple of feasible methods based on spatial audio techniques, that both track bats in flight and pinpoint the directions of echoes received by a bat. A spatial audio/soundfield microphone array is introduced to measure sounds in the sonar frequency range (20-80 kHz) of the big brown bat (Eptesicus fuscus). The custom-built ultrasonic tetrahedral soundfield microphone consists of four capacitive microphones that were calibrated to match magnitude and phase responses using a transfer function approach. Ambisonics, a signal processing technique used in three-dimensional (3D) audio applications, is used for the basic processing and reproduction of the signals measured by the soundfield microphone. Ambisonics provides syntheses and decompositions of a signal containing its directional properties, using the relationship between the spherical harmonics and the directional properties. As the first proposed method, a spatial audio decoding technique called HARPEx (High Angular Resolution Planewave Expansion) was used to build a system providing angle and elevation estimates. HARPEx can estimate the direction of arrivals (DOA) for up to two simultaneous sources since it decomposes a signal into two dominant planewaves. Experiments proved that the estimation system based on HARPEx provides accurate DOA estimates of static or moving sources. It also reconstructed a smooth flight-path of a bat by accurately estimating its direction at each snapshot of pulse measurements in time. The performance of the system was also assessed using statistical analyses of simulations. A signal model was built to generate microphone capsule responses to a virtual source emitting an LFM signal (3 ms, two harmonics: 40-22 kHz and 80-44 kHz) at an angle of 30° in the simulations. Medians and RMSEs (root-mean-square error) of 10,000 simulations for each case represent the accuracy and precision of the estimations, respectively. Results show lower d (distance between a capsule and the soundfield microphone center) or/and higher SNR (signal-to-noise ratio) are required to achieve higher estimator performance. The Cramer-Rao lower bounds (CRLB) of the estimator are also computed with various d and SNR conditions. The CRLB which is for TDOA based methods does not cover the effects of different incident angles to the capsules and signal delays between the capsules due to a non-zero d, on the estimation system. This shows the CRLB is not a proper tool to assess the estimator performance. For the second proposed method, the matched-filter technique is used instead of HARPEx to build another estimation system. The signal processing algorithm based on Ambisonics and the matched-filter approach reproduces a measured signal in various directions, and computes matched-filter responses of the reproduced signals in time-series. The matched-filter result points a target(s) by the highest filter response. This is a sonar-like estimation system that provides information of the target (range, direction, and velocity) using sonar fundamentals. Experiments using a loudspeaker (emitter) and an artificial or natural target (either stationary or moving) show the system provides accurate estimates of the target's direction and range. Simulations of imitating a situation where a bat emits a pulse and receives an echo from a target (30°) were also performed. The echo sound level is determined using the sonar equation. The system processed the virtual bat pulse and echo, and accurately estimated the direction, range, and velocity of the target. The simulation results also appear to recommend an echo level over -3 dB for accurate and precise estimations (below 15% RMSE for all parameters). This work proposes two methods to track bats in flight or/and pinpoint the directions of targets using spatial audio techniques. The suggested methods provide accurate estimates of the direction, range, or/and velocity of a bat based on its pulses or of a target based on echoes. This demonstrates these methods can be used as key tools to reconstruct bat biosonar. They would be also an independent tool or a complementary option to TDOA based methods, for bat echolocation studies. The developed methods are believed to be also useful in improving man-made sonar technology. / Doctor of Philosophy / While bats are one of the most intriguing creatures to the general population, they are also a popular subject of study in various disciplines. Their extraordinary ability to navigate and forage irrespective of clutter using echolocation has gotten attention from many scientists and engineers. Research investigating bats typically includes analysis of acoustic signals from microphones and/or microphone arrays. Using time difference of arrival (TDOA) between the array elements or the microphones is probably the most popular method to locate flying bats (azimuth and elevation). Microphone responses to transmitted signals and echoes near a bat provide sound pressure but no directional information. This dissertation proposes a complementary way to the current TDOA methods, that delivers directional information by introducing spatial audio techniques. This work shows a couple of feasible methods based on spatial audio techniques, that can both track bats in flight and pinpoint the directions of echoes received by a bat. An ultrasonic tetrahedral soundfield microphone is introduced as a measurement tool for sounds in the sonar frequency range (20-80 kHz) of the big brown bat (Eptesicus fuscus). Ambisonics, a signal processing technique used in three-dimensional (3D) audio applications, is used for the basic processing of the signals measured by the soundfield microphone. Ambisonics also reproduces a measured signal containing its directional properties. As the first method, a spatial audio decoding technique called HARPEx (High Angular Resolution Planewave Expansion) was used to build a system providing angle and elevation estimates. HARPEx can estimate the direction of arrivals (DOA) for up to two simultaneous sound sources. Experiments proved that the estimation system based on HARPEx provides accurate DOA estimates of static or moving sources. The performance of the system was also assessed using statistical analyses of simulations. Medians and RMSEs (root-mean-square error) of 10,000 simulations for each simulation case represent the accuracy and precision of the estimations, respectively. Results show shorter distance between a capsule and the soundfield microphone center, or/and higher SNR (signal-to-noise ratio) are required to achieve higher performance. For the second method, the matched-filter technique is used to build another estimation system. This is a sonar-like estimation system that provides information of the target (range, direction, and velocity) using matched-filter responses and sonar fundamentals. Experiments using a loudspeaker (emitter) and an artificial or natural target (either stationary or moving) show the system provides accurate estimates of the target's direction and range. Simulations imitating a situation where a bat emits a pulse and receives an echo from a target (30°) were also performed. The system processed the virtual bat pulse and echo, and accurately estimated the direction, range, and velocity of the target. The suggested methods provide accurate estimates of the direction, range, or/and velocity of a bat based on its pulses or of a target based on echoes. This demonstrates these methods can be used as key tools to reconstruct bat biosonar. They would be also an independent tool or a complementary option to TDOA based methods, for bat echolocation studies. The developed methods are also believed to be useful in improving sonar technology.