Echolocation Pulses Research Articles

Cerebellar activity predicts vocalization in fruit bats

Echolocating bats exhibit remarkable auditory behaviors, enabled by adaptations both within and outside their auditory system. Yet research on echolocating bats has focused mostly on brain areas that belong to the classic ascending auditory pathway. This study provides direct evidence linking the cerebellum, an evolutionarily ancient and non-classic auditory structure, to vocalization and hearing. We report that in the fruit-eating bat Carollia perspicillata, external sounds can evoke cerebellar responses with latencies below 20ms. Such fast responses are indicative of early inputs to the bat cerebellum. After establishing fruit-eating bats as a good model to study cerebellar auditory responses, we searched for a neural correlate of vocal production within the cerebellum. We investigated spike trains and field potentials occurring before and after vocalization and found that the type of sound produced (echolocation pulses or communication calls) can be decoded from pre-vocal and post-vocal neural signals, with prediction accuracies that reach above 85%. The latter provides a direct correlate of vocalization in an ancient motor-coordination structure that lies outside of the classic ascending auditory pathway. Taken together, our findings provide evidence of specializations for vocalization and hearing in the cerebellum of an auditory specialist.

Buzzfindr: Automating the detection of feeding buzzes in bat echolocation recordings.

Quantification of bat communities and habitat heavily rely on non-invasive acoustic bat surveys the scope of which has greatly amplified with advances in remote monitoring technologies. Despite the unprecedented amount of acoustic data being collected, analysis of these data is often limited to simple species classification which provides little information on habitat function. Feeding buzzes, the rapid sequences of echolocation pulses emitted by bats during the terminal phase of prey capture, have historically been used to evaluate foraging habitat quality. Automated identification of feeding buzzes in recordings could benefit conservation by helping identify critical foraging habitat. I tested if detection of feeding buzzes in recordings could be automated with bat recordings from Ontario, Canada. Data were obtained using three different recording devices. The signal detection method involved sequentially scanning narrow frequency bands with the "Bioacoustics" R package signal detection algorithm, and extracting temporal and signal strength parameters from detections. Buzzes were best characterized by the standard deviation of the time between consecutive pulses, the average pulse duration, and the average pulse signal-to-noise ratio. Classification accuracy was highest with artificial neural networks and random forest algorithms. I compared each model's receiver operating characteristic curves and random forest provided better control over the false-positive rate so it was retained as the final model. When tested on a new dataset, buzzfindr's overall accuracy was 93.4% (95% CI: 91.5%- 94.9%). Overall accuracy was not affected by recording device type or species frequency group. Automated detection of feeding buzzes will facilitate their integration in the analytical workflow of acoustic bat studies to improve inferences on habitat use and quality.

Sex Differences in Cochlear Transcriptomes in Horseshoe Bats.

Sexual dimorphism of calls is common in animals, whereas studies on the molecular basis underlying this phenotypic variation are still scarce. In this study, we used comparative transcriptomics of cochlea to investigate the sex-related difference in gene expression and alternative splicing in four Rhinolophus taxa. Based on 31 cochlear transcriptomes, we performed differential gene expression (DGE) and alternative splicing (AS) analyses between the sexes in each taxon. Consistent with the degree of difference in the echolocation pulse frequency between the sexes across the four taxa, we identified the largest number of differentially expressed genes (DEGs) and alternatively spliced genes (ASGs) in R. sinicus. However, we also detected multiple DEGs and ASGs in taxa without sexual differences in echolocation pulse frequency, suggesting that these genes might be related to other parameters of echolocation pulse rather than the frequency component. Some DEGs and ASGs are related to hearing loss or deafness genes in human or mice and they can be considered to be candidates associated with the sexual differences of echolocation pulse in bats. We also detected more than the expected overlap of DEGs and ASGs in two taxa. Overall, our current study supports the important roles of both DGE and AS in generating or maintaining sexual differences in animals.

S-method based on modified group delay for time-frequency analysis

A noise robust time-frequency representation based on the S-method (SM) and modified group delay (MGD) (S-MGD) is proposed in this article. The SM is free from cross-terms as it is the sum of individual Wigner distributions of different signal components and provides high-energy concentration both in time and frequency. The MGD reduces the Gibbs ripple and spurious spectral peaks because of noise, without using any frequency smoothing, and thus maintains the best frequency resolution as that of rectangular window. Hence,a combination of SM and MGD provides the desirable features of both the methods. The proposed method is tested for different signals like time varying random process, frequency shift keying, synthesized car engine sound/pressure signals using a solo combustion and a real-time echolocation pulse emitted by a big brown bat (Eptesicus fuscus) which indicate that the new S-MGD has improved noise immunity and frequency resolution compared to the method that uses frequency smoothing to suppress noise and Gibbs effects.

Development of the hyolaryngeal architecture in horseshoe bats: insights into the evolution of the pulse generation for laryngeal echolocation

BackgroundThe hyolaryngeal apparatus generates biosonar pulses in the laryngeally echolocating bats. The cartilage and muscles comprising the hyolarynx of laryngeally echolocating bats are morphologically modified compared to those of non-bat mammals, as represented by the hypertrophied intrinsic laryngeal muscle. Despite its crucial contribution to laryngeal echolocation, how the development of the hyolarynx in bats differs from that of other mammals is poorly documented. The genus Rhinolophus is one of the most sophisticated laryngeal echolocators, with the highest pulse frequency in bats. The present study provides the first detailed description of the three-dimensional anatomy and development of the skeleton, cartilage, muscle, and innervation patterns of the hyolaryngeal apparatus in two species of rhinolophid bats using micro-computed tomography images and serial tissue sections and compares them with those of laboratory mice. Furthermore, we measured the peak frequency of the echolocation pulse in active juvenile and adult individuals to correspond to echolocation pulses with hyolaryngeal morphology at each postnatal stage.ResultsWe found that the sagittal crests of the cricoid cartilage separated the dorsal cricoarytenoid muscle in horseshoe bats, indicating that this unique morphology may be required to reinforce the repeated closure movement of the glottis during biosonar pulse emission. We also found that the cricothyroid muscle is ventrally hypertrophied throughout ontogeny, and that the cranial laryngeal nerve has a novel branch supplying the hypertrophied region of this muscle. Our bioacoustic analyses revealed that the peak frequency shows negative allometry against skull growth, and that the volumetric growth of all laryngeal cartilages is correlated with the pulse peak frequency.ConclusionsThe unique patterns of muscle and innervation revealed in this study appear to have been obtained concomitantly with the acquisition of tracheal chambers in rhinolophids and hipposiderids, improving sound intensity during laryngeal echolocation. In addition, significant protrusion of the sagittal crest of the cricoid cartilage and the separated dorsal cricoarytenoid muscle may contribute to the sophisticated biosonar in this laryngeally echolocating lineage. Furthermore, our bioacoustic data suggested that the mineralization of these cartilages underpins the ontogeny of echolocation pulse generation. The results of the present study provide crucial insights into how the anatomy and development of the hyolaryngeal apparatus shape the acoustic diversity in bats.

Open-source machine learning BANTER acoustic classification of beaked whale echolocation pulses

Passive acoustic monitoring is increasingly used for assessing populations of marine mammals; however, analysis of large datasets is limited by our ability to easily classify sounds detected. Classification of beaked whale acoustic events, in particular, requires evaluation of multiple lines of evidence by expert analysts. Here we present a highly automated approach to acoustic detection and classification using supervised machine learning and open source software methods. Data from four large scale surveys of beaked whales (northwestern North Atlantic, southwestern North Atlantic, Hawaii, and eastern North Pacific) were analyzed using PAMGuard (acoustic detection), PAMpal (acoustic analysis) and BANTER (hierarchical random forest classifier). Overall classification accuracy ranged from 88% for the southwestern North Atlantic data to 97% for the northwestern North Atlantic. Results for many species could likely be improved with increased sample sizes, consideration of alternative automated detectors, and addition of relevant environmental features. These methods provide a highly automated approach to acoustic detection and classification using open source methods that can be readily adopted for other species and geographic regions.

Deviance Detection to Natural Stimuli in Population Responses of the Brainstem of Bats.

Deviance detection describes an increase of neural response strength caused by a stimulus with a low probability of occurrence. This ubiquitous phenomenon has been reported for humans and multiple other species, from subthalamic areas to the auditory cortex. Cortical deviance detection has been well characterized by a range of studies using a variety of different stimuli, from artificial to natural, with and without a behavioral relevance. This allowed the identification of a broad variety of regularity deviations that are detected by the cortex. Moreover, subcortical deviance detection has been studied with simple stimuli that are not meaningful to the subject. Here, we aim to bridge this gap by using noninvasively recorded auditory brainstem responses (ABRs) to investigate deviance detection at population level in the lower stations of the auditory system of a highly vocal species: the bat Carollia perspicillata (of either sex). Our present approach uses behaviorally relevant vocalization stimuli that are similar to the animals' natural soundscape. We show that deviance detection in ABRs is significantly stronger for echolocation pulses than for social communication calls or artificial sounds, indicating that subthalamic deviance detection depends on the behavioral meaning of a stimulus. Additionally, complex physical sound features like frequency- and amplitude modulation affected the strength of deviance detection in the ABR. In summary, our results suggest that the brain can detect different types of deviants already in the brainstem, showing that subthalamic brain structures exhibit more advanced forms of deviance detection than previously known.

Description of the echolocation pulses of insectivorous bats with new records for Southwest Colombia

Despite the high diversity of bats in neotropics, traditional methods such as mist nets, harp traps and roost detection have limitations in capturing that diversity in a landscape, with most detected species restricted to those that forage in the undergrowth or enclosed spaces. Therefore, acoustic records become a tool that complements and enhances the efforts to get more complete bat inventories while avoiding alterations in usual foraging activities and disruption in their life cycles. This study describes the acoustic parameters (spectral and temporal variables) of the echolocation pulses of insectivorous bats to characterise different species of bats in Southwest Colombia acoustically. We recorded echolocation calls between December 2017 and May 2020 in the Andean and Pacific regions of the Department of Nariño. We analysed 81 sequences of echolocation calls from eight bat species belonging to three families: Vespertilionidae, Molossidae and Emballonuridae. We perform recordings on free-flying bats with identity corroboration by capture for recording in flight rooms and examination in the hand. Myotis riparius and Lasiurus blossevillii were recorded for the first time in the Nariño Department. M. albescens, M. keaysi, M. riparius and L. blossevillii (Vespertilionidae) had pulses of frequency modulated (FM) with a quasi-constant frequency (QCF) ending; Molossus molossus, Tadarida brasiliensis and Promops centralis (Molossidae) had pulses with constant frequency (CF) and QCF; and Saccopteryx bilineata (Emballonuridae) had pulses with QCF. This study contributes to the efforts to facilitate the identification of insectivorous bats of the Neotropics using the acoustic monitoring approaches, represents a reference to compare the acoustic studies in Southwestern Colombia and contributes to increasing our knowledge of the bat diversity in the region.

Artificial light affects foraging behavior of a synanthropic bat.

Artificial light at night has been considered an emerging threat to global biodiversity. However, the impacts of artificial light on foraging behavior in most wild animals remain largely unclear. Here, we aimed to assess whether artificial light affects foraging behavior in Asian parti-colored bats (Vespertilio sinensis). We manipulated the spectra of light-emitting diode (LED) lighting in a laboratory. Using video and audio recording, we monitored foraging onset, total foraging time, food consumption, freezing behavior (temporary cessation of body movement), and echolocation vocalizations in triads of bats under each lighting condition. Analyses showed that the foraging activities of experimental bats were reduced under LED light. Green, yellow, and red light had greater negative effects on bats' foraging onset, total foraging time, and food consumption than white and blue light. LED light of different spectra induced increased freezing time and echolocation vocalizations in captive bats, except for the white light. The peak wavelength of light emission correlated positively with freezing time, estimated echolocation pulse rate (the number of echolocation pulses per minute), and foraging onset, but negatively with total foraging time and food consumption. These results demonstrate that artificial light disturbs foraging behavior in Asian parti-colored bats. Our findings have implications for understanding the influencing mechanism of light pollution on bat foraging.

Geographic distribution of the Cross Seamount beaked whale based on acoustic detections

AbstractBeaked whales produce frequency‐modulated echolocation pulses that appear to be species‐specific, allowing passive acoustic monitoring to play a role in understanding spatio‐temporal patterns. The Cross Seamount beaked whale is known only from its unique echolocation signal (BWC) with no confirmed species identification. This beaked whale spans the Pacific Ocean from the Mariana Archipelago to Baja California, Mexico, south to the equator, but only as far north as latitude 29°N. Within these warm waters, 92% of BWC detections occurred at night, 6% during crepuscular periods, and only 2% during daylight hours. Detections of BWC signals on drifting recorders with a vertical hydrophone array at 150 m depth demonstrated that foraging often occurred shallow in the water column (<150 m). No other species of beaked whale to date has been documented foraging in waters this shallow. Given their nocturnal, shallow foraging dives, this species appears to prefer prey that may be available in the water column only during those hours. The foraging behavior of Cross Seamount beaked whales appears to be unique among all beaked whales, and these findings contribute additional ecological and acoustic information which can help guide future efforts to identify this cryptic whale.

An Efficient Neural Network Design Incorporating Autoencoders for the Classification of Bat Echolocation Sounds.

Bats are widely distributed around the world, have adapted to many different environments and are highly sensitive to changes in their habitat, which makes them essential bioindicators of environmental changes. Passive acoustic monitoring over long durations, like months or years, accumulates large amounts of data, turning the manual identification process into a time-consuming task for human experts. Automated acoustic monitoring of bat activity is therefore an effective and necessary approach for bat conservation, especially in wind energy applications, where flying animals like bats and birds have high fatality rates. In this work, we provide a neural-network-based approach for bat echolocation pulse detection with subsequent genus classification and species classification under real-world conditions, including various types of noise. Our supervised model is supported by an unsupervised learning pipeline that uses autoencoders to compress linear spectrograms into latent feature vectors that are fed into a UMAP clustering algorithm. This pipeline offers additional insights into the data properties, aiding in model interpretation. We compare data collected from two locations over two consecutive years sampled at four heights (10 m, 35 m, 65 m and 95 m). With sufficient data for each labeled bat class, our model is able to comprehend the full echolocation soundscape of a species or genus while still being computationally efficient and simple by design. Measured classification F1 scores in a previously unknown test set range from 92.3% to 99.7% for species and from 94.6% to 99.4% for genera.

Spectrally non-overlapping background noise disturbs echolocation via acoustic masking in the CF-FM bat, Hipposideros pratti

The environment noise may disturb animal behavior and echolocation via three potential mechanisms: acoustic masking, reduced attention and noise avoidance. Compared with the mechanisms of reduced attention and noise avoidance, acoustic masking is thought to occur only when the signal and background noise overlap spectrally and temporally. In this study, we investigated the effects of spectrally non-overlapping noise on echolocation pulses and electrophysiological responses of a constant frequency-frequency modulation (CF-FM) bat, Hipposideros pratti. We found that H. pratti called at higher intensities while keeping the CFs of their echolocation pulses consistent. Electrophysiological tests indicated that the noise could decrease auditory sensitivity and sharp intensity tuning, suggesting that spectrally non-overlapping noise imparts an acoustic masking effect. Because anthropogenic noises are usually concentrated at low frequencies and are spectrally non-overlapping with the bat's echolocation pulses, our results provide further evidence of negative consequences of anthropogenic noise. On this basis, we sound a warning against noise in the foraging habitats of echolocating bats.

Mutual mother-pup acoustic identification in Asian particolored bats.

In many vertebrates, vocal communication is crucial in parent-offspring interactions, and parents are often able to discriminate between the calls of their own and others' offspring. There are many reports on the unidirectional recognition of isolation calls of pups by maternal bats, but few studies on the ability of bat pups to recognize maternal acoustic signals. In this study, we investigated whether the echolocation pulses of female Asian particolored bats (Vespertilio sinensis) and isolation calls of pups differ statistically among individuals. We used two-choice playback experiments to test whether the mothers and pups of V.sinensis can recognize each other by acoustic signals. Both the echolocation pulses of mother bats and the isolation calls of pups contained sufficient individual characteristics. Playback experiments showed that mothers were able to recognize isolation calls of pups, and most pups greater than 12 days old were able to distinguish echolocation pulses of their own mother from those of other mothers. This is the first use of two-choice acoustic signal playback experiments to confirm that pups can recognize their mothers by echolocation calls. The results provide behavioral evidence for bidirectional recognition of acoustic signals between mothers and infants in frequency-modulated type bats.

Dive characteristics of Cross Seamount beaked whales from long‐term passive acoustic monitoring at the Pacific Missile Range Facility, Kauaʻi

AbstractBeaked whale foraging pulses were detected on the Pacific Missile Range Facility (PMRF) off Kauaʻi, Hawaiʻi, via long term passive acoustic monitoring. The unidentified pulses do not match foraging pulses of known species on the range but are similar to the unidentified beaked whale first detected at Cross Seamount, Hawaiʻi. Although there has not been a visual confirmation of the unidentified beaked whale species, analysis of data collected from 2007 to 2019 has identified beaked whale foraging dive characteristics from echolocation pulses. From the 13 years of data, the most distinct patterns were that all foraging dives occurred at night and the nighttime foraging dive rate was 0.11 group vocal periods (GVP) per hour, with most detections on shallow hydrophones (625–1,000 m deep) over steep bathymetric slopes. Data collected during U.S. Navy training events were used to compare dive behavior during mid‐frequency active sonar (MFAS) activity against baseline periods; it was determined that the whales reduced GVPs during sonar and these remained low for at least 3 days after the training events. These results are the first long‐term record of acoustic signals from the Cross Seamount beaked whale and provide important insights into their habitat use and occurrence patterns.

Echolocation-related reversal of information flow in a cortical vocalization network

The mammalian frontal and auditory cortices are important for vocal behavior. Here, using local-field potential recordings, we demonstrate that the timing and spatial patterns of oscillations in the fronto-auditory network of vocalizing bats (Carollia perspicillata) predict the purpose of vocalization: echolocation or communication. Transfer entropy analyses revealed predominant top-down (frontal-to-auditory cortex) information flow during spontaneous activity and pre-vocal periods. The dynamics of information flow depend on the behavioral role of the vocalization and on the timing relative to vocal onset. We observed the emergence of predominant bottom-up (auditory-to-frontal) information transfer during the post-vocal period specific to echolocation pulse emission, leading to self-directed acoustic feedback. Electrical stimulation of frontal areas selectively enhanced responses to sounds in auditory cortex. These results reveal unique changes in information flow across sensory and frontal cortices, potentially driven by the purpose of the vocalization in a highly vocal mammalian model.

Airport noise disturbs foraging behavior of Japanese pipistrelle bats.

The expansion of anthropogenic noise poses an emerging threat to the survival and reproductive success of various organisms. Previous investigations have focused on the detrimental effects of anthropogenic noise on the foraging behavior in some terrestrial and aquatic animals. Nevertheless, the role of airport noise in impairing foraging activities of most wild animals has been neglected. Here, we aimed to assess whether foraging behavior in free‐living Japanese pipistrelle bats (Pipistrellus abramus) can be disturbed by airport noise. We used audio recording to monitor foraging activities of bats at 11 sites around the runway of a municipal airport. We quantified noise level and spectra, aircraft activity, habitat type, nightly temperature, wind speed, and moon phase for each site. The analysis revealed that noise level and aircraft activity were significant negative predictors for the number of bat passes and feeding buzzes around the runway, even after controlling for the effects of other environmental factors. There was no marked spectral overlap between bat echolocation pulses and airport noise in the presence and absence of low‐flying aircraft. The spectro‐temporal parameters of echolocation vocalizations emitted by bats were dependent on noise level, aircraft activity, and habitat type. These results provide correlative evidence that airport noise can reduce foraging activities of wild pipistrelle bats. Our findings add to the current knowledge of adverse impacts of airport noise on foraging bats in artificial ecosystems and provide a basis for further research on the mechanisms behind noise pollution near airports.

Effect of anthropogenic noise on the echolocation pulses of the bats Molossus sinaloae and Mormoops megalophylla

Anthropogenic noise interferes with the acoustic signals of various wildlife species. For bats that use echolocation, noise can mask the information received in the echo. The effect of anthropogenic noise on the time and frequency components of echolocation pulses emitted by the aerial insectivorous bats Molossus sinaloae and Mormoops megalophylla in urban and natural habitats were evaluated. We hypothesized that the frequency components of pulses would increase in response to masking, while time components would not change significantly. To this end, acoustic recordings of both species were made in the two types of habitats using ultrasonic detectors; simultaneously, the intensity of the ambient noise was measured. Frequency (kHz) and time (ms) were analyzed for each echolocation pulse using the BatSound 4.2 software. Consistent with our hypothesis, the results showed that under background noise of 75 dB in an urban environment, M. sinaloae increased the low and high frequencies of its echolocation pulses by 5.8 kHz on average. For M. megalophylla, no increase in pulse frequencies was observed. Contrary to our expectation, the time components of pulses for M. sinaloae were modified, being of shorter duration in urban sites. Increasing the maximum amplitude-frequency by M. sinaloae may be a response to the Lombard effect, i. e., the increase in vocal amplitude in response to increased background noise. It is important to carry out studies focused on understanding the modification of echolocation pulses, mainly for species living in urban environments.

Detection distances in desert dwelling, high duty cycle echolocators: A test of the foraging habitat hypothesis.

High Duty Cycle (HDC) echolocating bats use high frequency echolocation pulses that are clutter resistant, but their high frequencies give them limited range. Despite their unique ability to reject background clutter while simultaneously detecting fluttering prey, the frequency of their echolocation pulses has a strong correlation with level of environmental clutter, lower frequency pulses of HDC bats being associated with more open environments. The Foraging Habitat Hypothesis (FHH) proposes that the ecological significance of these lower frequency pulses in HDC bats in open environments is that they allow longer prey detection distances. To test the FHH, we compared the frequencies, Source Levels (SLs) and detection distances of Rhinolophus capensis, a HDC bat that has been shown to vary its call frequency in relation to habitat structure. As a further test of the FHH we investigated the SLs and detection distances of Rhinolophus damarensis (a heterospecific species that occurs in the same open desert environment as R. capensis but echolocates at a higher dominant pulse frequency). In the open desert, R. capensis emitted both lower frequency and higher SL pulses giving them longer detection distances than R. capensis in the cluttered fynbos. SL contributed more to differences in detection distances in both R. capensis and R. damarensis than frequency. In a few instances, R. damarensis achieved similar detection distances to desert–inhabiting R. capensis by emitting much higher SLs despite their average SLs being lower. These results suggest that lower frequency echolocation pulses are not a prerequisite for open desert living but may increase detection distance while avoiding energetic costs required for high SLs.

Horseshoe bats use not changes in echo delay but Doppler shift to perceive approaching objects

Echolocating bats use echo delay for target ranging and Doppler shift information for relative velocity recognition. However, how they perceive moving objects remains unclear. To investigate this question, we played back echolocation pulses in real-time as virtual echoes to Japanese horseshoe bats (Rhinolophus ferrumequinum nippon) on a perch in a flight room. Since echoes coming back from an approaching object are theoretically characterized by both changes in echo delay and the presence of Doppler shift, we reproduced an artificial approaching object by encoding these two acoustic parameters in the virtual echoes. As a result, only Doppler shift evoked bats flight reaction, showing that they use only Doppler shift and not change in echo delay to perceive approaching objects. Also, we played back only constant frequency (CF) component and confirmed that they use the CF component to detect Doppler shift. Furthermore, as a response to the Doppler shift in the perceived echo, bats increased the bandwidth of the terminal component of their pulse. Surprisingly, this response occurred in the very first pulse after Doppler shift, which indicates bats can adapt their echolocation pulse characteristics to changing situations within a pulse. [This work was supported by JSPS KAKENHI Grant Nos. 18H03786 and 16H06542.]

Amplitude discrimination is predictably affected by echo frequency filtering in wideband echolocating bats.

Big brown bats echolocate using wideband frequency-modulated (FM) ultrasonic pulses, perceiving target range from echo delay and target size from echo amplitude. Echolocation pulses contain two prominent down-sweeping harmonics (FM1, ∼55-22 kHz; FM2, ∼100-55 kHz), which are affected differently by propagation to the target and back to the bat. Previous work demonstrates that big brown bats utilize the low frequencies in FM1 for target ranging, while FM2 only contributes if FM1 is also present. The present experiments test the hypothesis that the bat's ability to discriminate echo amplitude is also affected by selectively attenuating FM1 or FM2 in target or nontarget echoes. Bats were trained to perform an amplitude discrimination task with virtual echo targets located 83 cm away. Echo delay was fixed and echo amplitude was varied, while either FM1 or FM2 was attenuated by highpass or lowpass filtering. Bats' performance decreased when lower frequencies were attenuated in target echoes and when higher frequencies were attenuated in nontarget echoes. Performance was reversed in the opposite filtering conditions. The bat's ability to distinguish between virtual targets varying in amplitude at the same simulated range indicates a high level of focused attention for perceptual isolation of target from non-target echoes.