Age and interpulse interval relation from newborn to adult sperm whale (Physeter macrocephalus) off Mauritius.

Sperm whales (Physeter macrocephalus) have been studied for decades, but the development of their clicks during the animal growth is not yet well known. The click they emit during socialization and echolocation contains information about the length of their acoustic organs and, therefore the length of the body through the interpulse interval (IPI). This paper provides the first IPI/age relationship for juvenile male and female sperm whales (Physeter macrocephalus) based on field recordings of individuals whose age is largely known. Across 9 years, audiovisual recordings of a Mauritian sperm whale social unit were carried out. Adult female and juvenile sperm whales were identified and aged. The dataset made from those recordings is publicly available. The interpulse interval was measured for individuals whose ages ranged from 7 days to around 38 years. The growth of the acoustic organ of juveniles showed an early inter-individual variability as well as sexual dimorphism. Usual growth models were also fitted, predicting a mean I P I ∞ of 3.5 ms for adults and a physical maturity reached at around 30 years old. The use of passive acoustic monitoring (PAM) is one of the main tools used to study sperm whales. This IPI-age relationship may aid demographic studies on sperm whales by enabling PAM to assess the ages of recorded sperm whales.

Parameter estimation of the hyperbolic frequency-modulated bat calls using hyperbolic scale transform.

Echolocating bats are known to vary their waveforms at the phases of searching, approaching, and capturing the prey. It is meaningful to estimate the parameters of the calls for bat species identification and the technological improvements of the synthetic systems, such as radar and sonar. The type of bat calls is species-related, and many calls can be modeled as hyperbolic frequency- modulated (HFM) signals. To obtain the parameters of the HFM-modeled bat calls, a reversible integral transform, i.e., hyperbolic scale transform (HST), is proposed to transform a call into two-dimensional peaks in the "delay-scale" domain, based on which harmonic separation and parameter estimation are realized. Compared with the methods based on time-frequency analysis, the HST-based method does not need to extract the instantaneous frequency of the bat calls, only searching for peaks. The verification results show that the HST is suitable for analyzing the HFM-modeled bat calls containing multiple harmonics with a large energy difference, and the estimated parameters imply that the use of the waveforms from the searching phase to the capturing phase is beneficial to reduce the ranging bias, and the trends in parameters may be useful for bat species identification.

Echolocating bats show species-specific variation in susceptibility to acoustic forward masking.

Echolocating bats rely on precise auditory temporal processing to detect echoes generated by calls that may be emitted at rates reaching 150-200 Hz. High call rates can introduce forward masking perceptual effects that interfere with echo detection; however, bats may have evolved specializations to prevent repetition suppression of auditory responses and facilitate detection of sounds separated by brief intervals. Recovery of the auditory brainstem response (ABR) was assessed in two species that differ in the temporal characteristics of their echolocation behaviors: Eptesicus fuscus, which uses high call rates to capture prey, and Carollia perspicillata, which uses lower call rates to avoid obstacles and forage for fruit. We observed significant species differences in the effects of forward masking on ABR wave 1, in which E. fuscus maintained comparable ABR wave 1 amplitudes when stimulated at intervals of <3 ms, whereas post-stimulus recovery in C. perspicillata required 12 ms. When the intensity of the second stimulus was reduced by 20-30 dB relative to the first, however, C. perspicillata showed greater recovery of wave 1 amplitudes. The results demonstrate that species differences in temporal resolution are established at early levels of the auditory pathway and that these differences reflect auditory processing requirements of species-specific echolocation behaviors.

A social foraging trade-off in echolocating bats reveals that they benefit from some conspecifics but are impaired when many are around.

Social foraging is very common in the animal kingdom. Numerous studies have documented collective foraging in various species and many reported the attraction of various species to foraging conspecifics. It is nonetheless difficult to quantify the benefits and costs of collective foraging, especially in the wild. We examined the benefits and costs of social foraging using on-board microphones mounted on freely foraging Molossus nigricans bats. This allowed us to quantify the bats' attacks on prey and to assess their success as a function of conspecific density. We found that the bats spent most of their time foraging at low conspecific densities, during which their attacks were most successful in terms of prey items captured per time unit. Notably, their capture rate dropped when conspecific density became either too high or too low. Our findings thus demonstrate a clear social foraging trade-off in which the presence of a few conspecifics probably improves foraging success, whereas the presence of too many impairs it.

Auditory cortical regions show resting-state functional connectivity with the default mode-like network in echolocating bats.

Echolocating bats are among the most social and vocal of all mammals. These animals are ideal subjects for functional MRI (fMRI) studies of auditory social communication given their relatively hypertrophic limbic and auditory neural structures and their reduced ability to hear MRI gradient noise. Yet, no resting-state networks relevant to social cognition (e.g., default mode-like networks or DMLNs) have been identified in bats since there are few, if any, fMRI studies in the chiropteran order. Here, we acquired fMRI data at 7 Tesla from nine lightly anesthetized pale spear-nosed bats (Phyllostomus discolor). We applied independent components analysis (ICA) to reveal resting-state networks and measured neural activity elicited by noise ripples (on: 10 ms; off: 10 ms) that span this species' ultrasonic hearing range (20 to 130 kHz). Resting-state networks pervaded auditory, parietal, and occipital cortices, along with the hippocampus, cerebellum, basal ganglia, and auditory brainstem. Two midline networks formed an apparent DMLN. Additionally, we found four predominantly auditory/parietal cortical networks, of which two were left-lateralized and two right-lateralized. Regions within four auditory/parietal cortical networks are known to respond to social calls. Along with the auditory brainstem, regions within these four cortical networks responded to ultrasonic noise ripples. Iterative analyses revealed consistent, significant functional connectivity between the left, but not right, auditory/parietal cortical networks and DMLN nodes, especially the anterior-most cingulate cortex. Thus, a resting-state network implicated in social cognition displays more distributed functional connectivity across left, relative to right, hemispheric cortical substrates of audition and communication in this highly social and vocal species.

Bats integrate multiple echolocation and flight tactics to track prey.

The ability of "target tracking," such as keeping a target object in sight, is crucial for various activities. However, most sensing systems experience a certain degree of delay due to information processing, which challenges accurate target tracking. The long history of studies on animal behavior has revealed several tactics for it, although a systematic understanding of how individual tactics are combined into a strategy has not been reached. This study demonstrates a multifaceted tracking strategy in animals, which mitigates the adverse delay effects with small implementation costs. Using an active-sensing bat to measure their sensing state while chasing natural prey, we found that bats use a tracking strategy by combining multiple echolocation and flight tactics. The three echolocation tactics, namely the predictive control of sensing direction accompanied by adjusting the sensing rate and angular range, produce a direct compensation effect. Simultaneously, the flight tactic, the counter maneuver, assists echolocation by stabilizing the target direction. Our simulation results demonstrate that these combined tactics improve tracking accuracy over a wide range of delay constraints. In addition, a concise rule based on the angular velocity between bats and targets explains how bats control these tactics, suggesting that bats successfully reduce the burden of multitasking management. Our findings reveal the sophisticated strategy in animals' tracking systems and provide insights into understanding and developing efficiently integrated strategies in target tracking across various disciplines.

Changes in primary visual and auditory cortex of blind and sighted adults following 10 weeks of click-based echolocation training.

Recent work suggests that the adult human brain is very adaptable when it comes to sensory processing. In this context, it has also been suggested that structural "blueprints" may fundamentally constrain neuroplastic change, e.g. in response to sensory deprivation. Here, we trained 12 blind participants and 14 sighted participants in echolocation over a 10-week period, and used MRI in a pre-post design to measure functional and structural brain changes. We found that blind participants and sighted participants together showed a training-induced increase in activation in left and right V1 in response to echoes, a finding difficult to reconcile with the view that sensory cortex is strictly organized by modality. Further, blind participants and sighted participants showed a training induced increase in activation in right A1 in response to sounds per se (i.e. not echo-specific), and this was accompanied by an increase in gray matter density in right A1 in blind participants and in adjacent acoustic areas in sighted participants. The similarity in functional results between sighted participants and blind participants is consistent with the idea that reorganization may be governed by similar principles in the two groups, yet our structural analyses also showed differences between the groups suggesting that a more nuanced view may be required.

Clutter resilience via auditory stream segregation in echolocating greater mouse-eared bats.

Bats use echolocation to navigate and hunt in darkness, and must in that process segregate target echoes from unwanted clutter echoes. Bats may do this by approaching a target at steep angles relative to the plane of the background, utilizing their directional transmission and receiving systems to minimize clutter from background objects, but it remains unknown how bats negotiate clutter that cannot be spatially avoided. Here, we tested the hypothesis that when movement no longer offers spatial release, echolocating bats mitigate clutter by calling at lower source levels and longer call intervals to ease auditory streaming. We trained five greater mouse-eared bats (Myotis myotis) to land on a spherical loudspeaker with two microphones attached. We used a phantom-echo setup, where the loudspeaker/target transmitted phantom clutter echoes by playing back the bats' own calls at time delays of 1, 3 and 5 ms with a virtual target strength 7 dB higher than the physical target. We show that the bats successfully landed on the target, irrespective of the clutter echo delays. Rather than decreasing their source levels, the bats used similar source level distributions in clutter and control trials. Similarly, the bats did not increase their call intervals, but instead used the same distribution of call intervals across control and clutter trials. These observations reject our hypothesis, leading us to conclude that bats display great resilience to clutter via short auditory integration times and acute auditory stream segregation rather than via biosonar adjustments.

Automated echolocation classifiers vary in accuracy for northeastern U.S. bat species.

Acoustic surveys of bat echolocation calls are an important management tool for determining presence and probable absence of threatened and endangered bat species. In the northeastern United States, software programs such as Bat Call Identification (BCID), Kaleidoscope Pro (KPro), and Sonobat can automatically classify ultrasonic detector sound files, yet the programs' accuracy in correctly classifying calls to species has not been independently assessed. We used 1,500 full-spectrum reference calls with known identities for nine northeastern United States bat species to test the accuracy of these programs using calculations of Positive Predictive Value (PPV), Negative Predictive Value (NPV), Sensitivity (SN), Specificity (SP), Overall Accuracy, and No Information Rate. We found that BCID performed less accurately than other programs, likely because it only operates on zero-crossing data and may be less accurate for recordings converted from full-spectrum to zero-crossing. NPV and SP values were high across all species categories for SonoBat and KPro, indicating these programs' success at avoiding false positives. However, PPV and SN values were relatively low, particularly for individual Myotis species, indicating these programs are prone to false negatives. SonoBat and KPro performed better when distinguishing Myotis species from non-Myotis species. We expect less accuracy from these programs for acoustic recordings collected under normal working conditions, and caution that a bat acoustic expert should verify automatically classified files when making species-specific regulatory or conservation decisions.

Detection and validation of common noctule bats (Nyctalus noctula) with a pulse radar and acoustic monitoring in the proximity of an onshore wind turbine.

This paper presents the results of bats detected with marine radar and their validation with acoustic detectors in the vicinity of a wind turbine with a hub height of 120 m. Bat detectors are widely used by researchers, even though the common acoustic detectors can cover only a relatively small volume. In contrast, radar technology can overcome this shortcoming by offering a large detection volume, fully covering the rotor-swept areas of modern wind turbines. Our study focused on the common noctule bats (Nyctalus noctula). The measurement setup consisted of a portable X-band pulse radar with a modified radar antenna, a clutter shielding fence, and an acoustic bat detector installed in the wind turbine's nacelle. The radar's detection range was evaluated using an analytical simulation model. We developed a methodology based on a strict set of criteria for selecting suitable radar data, acoustic data and identified bat tracks. By applying this methodology, the study data was limited to time intervals with an average duration of 48 s, which is equal to approximately 20 radar images. For these time intervals, 323 bat tracks were identified. The most common bat speed was extracted to be between 9 and 10 m/s, matching the values found in the literature. Of the 323 identified bat tracks passed within 80 m of the acoustic detector, 32% had the potential to be associated with bat calls due to their timing, directionality, and distance to the acoustic bat detector. The remaining 68% passed within the studied radar detection volume but out of the detection volume of the acoustic bat detector. A comparison of recorded radar echoes with the expected simulated values indicated that the in-flight radar cross-section of recorded common noctule bats was mostly between 1.0 and 5.0 cm2, which is consistent with the values found in the literature for similar sized wildlife.

Complexin-1 enhances ultrasound neurotransmission in the mammalian auditory pathway.

Unlike megabats, which rely on well-developed vision, microbats use ultrasonic echolocation to navigate and locate prey. To study ultrasound perception, here we compared the auditory cortices of microbats and megabats by constructing reference genomes and single-nucleus atlases for four species. We found that parvalbumin (PV)+ neurons exhibited evident cross-species differences and could respond to ultrasound signals, whereas their silencing severely affected ultrasound perception in the mouse auditory cortex. Moreover, megabat PV+ neurons expressed low levels of complexins (CPLX1-CPLX4), which can facilitate neurotransmitter release, while microbat PV+ neurons highly expressed CPLX1, which improves neurotransmission efficiency. Further perturbation of Cplx1 in PV+ neurons impaired ultrasound perception in the mouse auditory cortex. In addition, CPLX1 functioned in other parts of the auditory pathway in microbats but not megabats and exhibited convergent evolution between echolocating microbats and whales. Altogether, we conclude that CPLX1 expression throughout the entire auditory pathway can enhance mammalian ultrasound neurotransmission.

Tiger beetles produce anti-bat ultrasound and are probable Batesian moth mimics.

Echolocating bats and their eared insect prey are in an acoustic evolutionary war. Moths produce anti-bat sounds that startle bat predators, signal noxiousness, mimic unpalatable models and jam bat sonar. Tiger beetles (Cicindelidae) also purportedly produce ultrasound in response to bat attacks. Here we tested 19 tiger beetle species from seven genera and showed that they produce anti-bat signals to playback of authentic bat echolocation. The dominant frequency of beetle sounds substantially overlaps the sonar calls of sympatric bats. As tiger beetles are known to produce defensive chemicals such as benzaldehyde and hydrogen cyanide, we hypothesized that tiger beetle sounds are acoustically advertising their unpalatability. We presented captive big brown bats (Eptesicus fuscus) with seven different tiger beetle species and found that 90 out of 94 beetles were completely consumed, indicating that these tiger beetle species are not aposematically signalling. Instead, we show that the primary temporal and spectral characteristics of beetle warning sounds overlap with sympatric unpalatable tiger moth (Arctinae) sounds and that tiger beetles are probably Batesian mimics of noxious moth models. We predict that many insect taxa produce anti-bat sounds and that the acoustic mimicry rings of the night sky are hyperdiverse.

Bat2Web: A Framework for Real-Time Classification of Bat Species Echolocation Signals Using Audio Sensor Data.

Bats play a pivotal role in maintaining ecological balance, and studying their behaviors offers vital insights into environmental health and aids in conservation efforts. Determining the presence of various bat species in an environment is essential for many bat studies. Specialized audio sensors can be used to record bat echolocation calls that can then be used to identify bat species. However, the complexity of bat calls presents a significant challenge, necessitating expert analysis and extensive time for accurate interpretation. Recent advances in neural networks can help identify bat species automatically from their echolocation calls. Such neural networks can be integrated into a complete end-to-end system that leverages recent internet of things (IoT) technologies with long-range, low-powered communication protocols to implement automated acoustical monitoring. This paper presents the design and implementation of such a system that uses a tiny neural network for interpreting sensor data derived from bat echolocation signals. A highly compact convolutional neural network (CNN) model was developed that demonstrated excellent performance in bat species identification, achieving an F1-score of 0.9578 and an accuracy rate of 97.5%. The neural network was deployed, and its performance was evaluated on various alternative edge devices, including the NVIDIA Jetson Nano and Google Coral.

Superfast Lombard response in free-flying, echolocating bats.

Acoustic cues are crucial to communication, navigation, and foraging in many animals, which hence face the problem of detecting and discriminating these cues in fluctuating noise levels from natural or anthropogenic sources. Such auditory dynamics are perhaps most extreme for echolocating bats that navigate and hunt prey on the wing in darkness by listening for weak echo returns from their powerful calls in complex, self-generated umwelts.1,2 Due to high absorption of ultrasound in air and fast flight speeds, bats operate with short prey detection ranges and dynamic sensory volumes,3 leading us to hypothesize that bats employ superfast vocal-motor adjustments to rapidly changing sensory scenes. To test this hypothesis, we investigated the onset and offset times and magnitude of the Lombard response in free-flying echolocating greater mouse-eared bats exposed to onsets of intense constant or duty-cycled masking noise during a landing task. We found that the bats invoked a bandwidth-dependent Lombard response of 0.1-0.2 dB per dB increase in noise, with very short delay and relapse times of 20 ms in response to onsets and termination of duty-cycled noise. In concert with the absence call time-locking to noise-free periods, these results show that free-flying bats exhibit a superfast, but hard-wired, vocal-motor response to increased noise levels. We posit that this reflex is mediated by simple closed-loop audio-motor feedback circuits that operate independently of wingbeat and respiration cycles to allow for rapid adjustments to the highly dynamic auditory scenes encountered by these small predators.

Echolocating Bats Have Evolved Decreased Susceptibility to Noise-Induced Temporary Hearing Losses.

Glenis Long championed the application of quantitative psychophysical methods to understand comparative hearing abilities across species. She contributed the first psychophysical studies of absolute and masked hearing sensitivities in an auditory specialist, the echolocating horseshoe bat. Her data demonstrated that this bat has hyperacute frequency discrimination in the 83-kHz range of its echolocation broadcast. This specialization facilitates the bat's use of Doppler shift compensation to separate echoes of fluttering insects from concurrent echoes of non-moving objects. In this review, we discuss another specialization for hearing in a species of echolocating bat that contributes to perception of echoes within a complex auditory scene. Psychophysical and behavioral studies with big brown bats show that exposures to long duration, intense wideband or narrowband ultrasonic noise do not induce significant increases in their thresholds to echoes and do not impair their ability to orient through a naturalistic sonar scene containing multiple distracting echoes. Thresholds of auditory brainstem responses also remain low after intense noise exposures. These data indicate that big brown bats are not susceptible to temporary threshold shifts as measured in comparable paradigms used with other mammals, at least within the range of stimulus parameters that have been tested so far. We hypothesize that echolocating bats have evolved a decreased susceptibility to noise-induced hearing losses as a specialization for echolocation in noisy environments.

Accurate species classification of Arctic toothed whale echolocation clicks using one-third octave ratios.

Passive acoustic monitoring has been an effective tool to study cetaceans in remote regions of the Arctic. Here, we advance methods to acoustically identify the only two Arctic toothed whales, the beluga (Delphinapterus leucas) and narwhal (Monodon monoceros), using echolocation clicks. Long-term acoustic recordings collected from moorings in Northwest Greenland were analyzed. Beluga and narwhal echolocation signals were distinguishable using spectrograms where beluga clicks had most energy >30 kHz and narwhal clicks had a sharp lower frequency limit near 20 kHz. Changes in one-third octave levels (TOL) between two pairs of one-third octave bands were compared from over one million click spectra. Narwhal clicks had a steep increase between the 16 and 25 kHz TOL bands that was absent in beluga click spectra. Conversely, beluga clicks had a steep increase between the 25 and 40 kHz TOL bands that was absent in narwhal click spectra. Random Forest classification models built using the 16 to 25 kHz and 25 to 40 kHz TOL ratios accurately predicted the species identity of 100% of acoustic events. Our findings support the use of echolocation TOL ratios in future automated click classifiers for acoustic monitoring of Arctic toothed whales and potentially for other odontocete species.

Click detection rate variability of central North Pacific sperm whales from passive acoustic towed arrays.

Passive acoustic monitoring (PAM) is an optimal method for detecting and monitoring cetaceans as they frequently produce sound while underwater. Cue counting, counting acoustic cues of deep-diving cetaceans instead of animals, is an alternative method for density estimation, but requires an average cue production rate to convert cue density to animal density. Limited information about click rates exists for sperm whales in the central North Pacific Ocean. In the absence of acoustic tag data, we used towed hydrophone array data to calculate the first sperm whale click rates from this region and examined their variability based on click type, location, distance of whales from the array, and group size estimated by visual observers. Our findings show click type to be the most important variable, with groups that include codas yielding the highest click rates. We also found a positive relationship between group size and click detection rates that may be useful for acoustic predictions of group size in future studies. Echolocation clicks detected using PAM methods are often the only indicator of deep-diving cetacean presence. Understanding the factors affecting their click rates provides important information for acoustic density estimation.

Correlated evolution between body size and echolocation in bats (order Chiroptera).

BACKGROUND: Body size and echolocation call frequencies are related in bats. However, it is unclear if this allometry applies to the entire clade. Differences have been suggested between nasal and oral emitting bats, as well as between some taxonomic families. Additionally, the scaling of other echolocation parameters, such as bandwidth and call duration, needs further testing. Moreover, it would be also interesting to test whether changes in body size have been coupled with changes in these echolocation parameters throughout bat evolution. Here, we test the scaling of peak frequency, bandwidth, and call duration with body mass using phylogenetically informed analyses for 314 bat species. We specifically tested whether all these scaling patterns differ between nasal and oral emitting bats. Then, we applied recently developed Bayesian statistical techniques based on large-scale simulations to test for the existence of correlated evolution between body mass and echolocation. RESULTS: Our results showed that echolocation peak frequencies, bandwidth, and duration follow significant allometric patterns in both nasal and oral emitting bats. Changes in these traits seem to have been coupled across the laryngeal echolocation bats diversification. Scaling and correlated evolution analyses revealed that body mass is more related to peak frequency and call duration than to bandwidth. We exposed two non-exclusive kinds of mechanisms to explain the link between size and each of the echolocation parameters. CONCLUSIONS: The incorporation of Bayesian statistics based on large-scale simulations could be helpful for answering macroevolutionary patterns related to the coevolution of traits in bats and other taxonomic groups.

Molecular adaptations underlying high-frequency hearing in the brain of CF bats species.

BACKGROUND: The majority of bat species have developed remarkable echolocation ability, especially for the laryngeally echolocating bats along with high-frequency hearing. Adaptive evolution has been widely detected for the cochleae in the laryngeally echolocating bats, however, limited understanding for the brain which is the central to echolocation signal processing in the auditory perception system, the laryngeally echolocating bats brain may also undergo adaptive changes. RESULT: In order to uncover the molecular adaptations related with high-frequency hearing in the brain of laryngeally echolocating bats, the genes expressed in the brain of Rhinolophus ferrumequinum (CF bat) and Myotis pilosus (FM bat) were both detected and also compared. A total of 346,891 genes were detected and the signal transduction mechanisms were annotated by the most abundant genes, followed by the transcription. In hence, there were 3,088 DEGs were found between the two bat brains, with 1,426 highly expressed in the brain of R. ferrumequinum, which were significantly enriched in the neuron and neurodevelopmental processes. Moreover, we found a key candidate hearing gene, ADCY1, playing an important role in the R. ferrumequinum brain and undergoing adaptive evolution in CF bats. CONCLUSIONS: Our study provides a new insight to the molecular bases of high-frequency hearing in two laryngeally echolocating bats brain and revealed different nervous system activities during auditory perception in the brain of CF bats.

Bioinspiration from bats and new paradigms for autonomy in natural environments.

Achieving autonomous operation in complex natural environment remains an unsolved challenge. Conventional engineering approaches to this problem have focused on collecting large amounts of sensory data that are used to create detailed digital models of the environment. However, this only postpones solving the challenge of identifying the relevant sensory information and linking it to action control to the domain of the digital world model. Furthermore, it imposes high demands in terms of computing power and introduces large processing latencies that hamper autonomous real-time performance. Certain species of bats that are able to navigate and hunt their prey in dense vegetation could be a biological model system for an alternative approach to addressing the fundamental issues associated with autonomy in complex natural environments. Bats navigating in dense vegetation rely on clutter echoes, i.e. signals that consist of unresolved contributions from many scatters. Yet, the animals are able to extract the relevant information from these input signals with brains that are often less than 1 g in mass. Pilot results indicate that information relevant to location identification and passageway finding can be directly obtained from clutter echoes, opening up the possibility that the bats' skill can be replicated in man-made autonomous systems.