Rapid jamming avoidance in biosonar

The sonar systems of bats and dolphins are in many ways superior to man-made sonar and radar systems, and considerable effort has been devoted to understanding the signal-processing strategies underlying these capabilities. A major feature determining the efficiency of sonar systems is the sensitivity to noise and jamming signals. Previous studies indicated that echolocating bats may adjust their signal structure to avoid jamming ('jamming avoidance response'; JAR). However, these studies relied on behavioural correlations and not controlled experiments. Here, we provide the first experimental evidence for JAR in bats. We presented bats (Tadarida brasiliensis) with 'playback stimuli' consisting of recorded echolocation calls at one of six frequencies. The bats exhibited a JAR by shifting their call frequency away from the presented playback frequency. When the approaching bats were challenged by an abrupt change in the playback stimulus, they responded by shifting their call frequencies upwards, away from the playback. Interestingly, even bats initially calling below the playback's frequency shifted their frequencies upwards, 'jumping' over the playback frequency. These spectral shifts in the bats' calls occurred often within less than 200 ms, in the first echolocation call emitted after the stimulus switch-suggesting that rapid jamming avoidance is important for the bat.     

Phase sensitivity in bat sonar revisited

An echolocating bat produces echoes consisting of the convolution of echolocation call and the impulse response (IR) of the ensonified object. A crucial question in animal sonar is whether bats are able to extract this IR from the echo. The bat inner ear generates a frequency representation of call and echo and IR extraction in the frequency domain requires accurate analysis of both magnitude and phase information. Previous studies investigating the phase sensitivity of bats using a jitter paradigm reported a temporal acuity down to 10 ns, suggesting perfect sonar phase representation. In a phantom-target playback experiment, we investigate the perceptual phase sensitivity of the bat Phyllostomus discolor using a novel approach: instead of manipulating IR phase by changing IR delay (jitter paradigm), we randomized IR phase and thus lengthened the IR over time, leaving the magnitude spectrum unchanged. Our results show that phase sensitivity, as reflected in the analysis of signal duration, appears to be much lower than phase sensitivity, as reflected in the analysis of signal onset. The current data indicate that different temporal aspects of sonar processing are encoded with very different temporal resolution and thus an overall claim of “phase sensitivity” as such cannot be maintained.     

Click-based echolocation in bats: not so primitive after all

Echolocating bats of the genus Rousettus produce click sonar signals, using their tongue (lingual echolocation). These signals are often considered rudimentary and are believed to enable only crude performance. However, the main argument supporting this belief, namely the click’s reported long duration, was recently shown to be an artifact. In fact, the sonar clicks of Rousettus bats are extremely short, ~50–100 μs, similar to dolphin vocalizations. Here, we present a comparison between the sonar systems of the ‘model species’ of laryngeal echolocation, the big brown bat (Eptesicus fuscus), and that of lingual echolocation, the Egyptian fruit bat (Rousettus aegyptiacus). We show experimentally that in tasks, such as accurate landing or detection of medium-sized objects, click-based echolocation enables performance similar to laryngeal echolocators. Further, we describe a sophisticated behavioral strategy for biosonar beam steering in clicking bats. Finally, theoretical analyses of the signal design—focusing on their autocorrelations and wideband ambiguity functions—predict that in some aspects, such as target ranging and Doppler-tolerance, click-based echolocation might outperform laryngeal echolocation. Therefore, we suggest that click-based echolocation in bats should be regarded as a viable echolocation strategy, which is in fact similar to the biosonar used by most echolocating animals, including whales and dolphins.     

Calls in the Forest: A Comparative Approach to How Bats Find Tree Cavities

Although tree cavities are a particularly critical resource for forest bats, how bats search for and find new roosts is still poorly known. Building on a recent study on the sensory basis of roost finding in the noctule (Ruczynski et al. 2007), here we take a comparative approach to how bats find roosts. We tested the hypothesis that species' flight abilities and echolocation call characteristics play important roles in how well and by which cues bats find new tree roosts. We used the very manoeuvrable, faintly echolocating brown long-eared bat ( Plecotus auritus ) and the less manoeuvrable, louder Daubenton's bat ( Myotis daubentonii ) as study species. The species are sympatric in European temperate forests and both roost in tree cavities. We trained bats in short-term captivity to find entrances to tree cavities and experimentally manipulated the sensory cues available to them. In both species, cue type influenced the search time for successful cavity detection. Visual, olfactory and temperature cues did not improve the bats' performance over the performance by echolocation alone. Eavesdropping on conspecific echolocation calls played back from inside the cavity decreased search time in Daubenton's bat ( M. daubentonii ), underlining the double function of echolocation signals – orientation and communication. This was not so in the brown long-eared bat ( P. auritus ) that has low call amplitudes. The highly manoeuvrable P. auritus found cavities typically from flight and the less manoeuvrable M. daubentonii found more entrances during crawling. Comparison with the noctule data from Ruczyński et al. (2007) indicates that manoeuvrability predicts the mode of cavity search. It further highlights the importance of call amplitude for eavesdropping and cavity detection in bats.     

Detection of sonar signals in the presence of pulses of masking noise by the echolocating bat,Eptesicus fuscus

Summary Two big brown bats (Eptesicus fuscus) were trained to report the presence or absence of a virtual sonar target. The bats' sensitivity to transient masking was investigated by adding 5 ms pulses of white noise delayed from 0 to 16 ms relative to the target echo. When signal and masker occurred simultaneously, the bats required a signal energy to noise spectrum level ratio of 35 dB for 50% probability of detection. When the masker was delayed by 2 ms or more there was no significant masking and echo energy could be reduced by 30 dB for the same probability of detection. The average duration of the most energetic sonar signal of each trial was measured to be 1.7 ms and 2.4 ms for the two bats, but a simple relation between detection performance and pulse duration was not found.In a different experiment the masking noise pulses coincided with the echo, and the duration of the masker was varied from 2 to 37.5 ms. The duration of the masker had little or no effect on the probability of detection.The findings are consistent with an aural integration time constant of about 2 ms, which is comparable to the duration of the cries. This is an order of magnitude less than found in backward masking experiments with humans and may be an adaptation to the special constraints of echolocation. The short time of sensitivity to masking may indicate that the broad band clicks of arctiid moths produced as a countermeasure to bat predation are unlikely to function by masking the echo of the moth.Abbreviations SPL sound pressure level - SD standard deviation - SE standard error - BW bandwidth     

Female big brown bats, Eptesicus fuscus, recognize sex from a caller's echolocation signals

The echolocation calls of bats function in prey capture and navigation but are not commonly regarded as communicative signals. However, because bats' echolocation calls show patterns of variability, they may transmit information about a bat, such as its age, individual identity or sex. For echolocation calls to function in this manner, variation in calls must be reliably linked to the characteristics of the bat, as has been shown in a number of studies. However, few studies have asked whether bats respond to this variation. We tested whether female big brown bats can identify the sex of an unfamiliar bat from playbacks of its echolocation calls. Playback consisted of a 30-s preplayback period, a 60-s playback period of either male or female echolocation calls, and a 30-s postplayback period. In the playback and postplayback periods the vocalization rates of female bats changed significantly relative to the preplayback period depending on the sex of the playback stimulus, indicating that they could determine sex from the echolocation calls. These findings support the possibility that echolocation calls play a role in communication in big brown bats.     

The role of echolocation in the hunting of terrestrial prey – new evidence for an underestimated strategy in the gleaning bat, Megaderma lyra

The observation that gleaning bats detect prey by its noises, together with difficulties in recording their faint sonar calls, have led some authors to conclude that gleaning bats may not use echolocation in certain hunting situations. In particular, it is conjectured that echolocation plays no role in the classification and tracking of prey. In the present study, we show that the gleaning bat, Megaderma lyra, is able to find silent and motionless prey on the ground. The significance of sonar for catching a variety of terrestrial prey is established in a standardized situation. Sonar calls were found to be emitted during all stages, i.e. approach, hovering above the prey, and return to the roost, of every hunting flight. The harmonic pattern of the calls differed significantly between these stages, calls with three or more prominent components prevailing during hovering. Bats identified prey and rejected dummies while hovering above them. During this stage, increased call rates and reduced call durations were found. Echolocation activity during, and the duration of, the hovering phase depended on prey type, in particular on prey movement. The prey-dependent shifts in sonar activity, the broadband call structure with an emphasis on higher harmonics, and a systematic shift of the calls' peak frequencies during hovering, are discussed as adaptations to identifying prey by sonar.     

Neural coding of echo-envelope disparities in echolocating bats

The effective use of echolocation requires not only measuring the delay between the emitted call and returning echo to estimate the distance of an ensonified object. To locate an object in azimuth and elevation, the bat’s auditory system must analyze the returning echoes in terms of their binaural properties, i.e., the echoes’ interaural intensity and time differences (IIDs and ITDs). The effectiveness of IIDs for echolocation is undisputed, but when bats ensonify complex objects, the temporal structure of echoes may facilitate the analysis of the echo envelope in terms of envelope ITDs. Using extracellular recordings from the auditory midbrain of the bat, Phyllostomus discolor, we found a population of neurons that are sensitive to envelope ITDs of echoes of their sonar calls. Moreover, the envelope-ITD sensitivity improved with increasing temporal fluctuations in the echo envelopes, a sonar parameter related to the spatial statistics of complex natural reflectors like vegetation. The data show that in bats envelope ITDs may be used not only to locate external, prey-generated rustling sounds but also in the context of echolocation. Specifically, the temporal fluctuations in the echo envelope, which are created when the sonar emission is reflected from a complex natural target, support ITD-mediated echolocation.     

Can two streams of auditory information be processed simultaneously? Evidence from the gleaning bat Antrozous pallidus

A tenet of auditory scene analysis is that we can fully process only one stream of auditory information at a time. We tested this assumption in a gleaning bat, the pallid bat (Antrozous pallidus) because this bat uses echolocation for general orientation, and relies heavily on prey-generated sounds to detect and locate its prey. It may therefore encounter situations in which the echolocation and passive listening streams temporally overlap. Pallid bats were trained to a dual task in which they had to negotiate a wire array, using echolocation, and land on one of 15 speakers emitting a brief noise burst in order to obtain a food reward. They were forced to process both streams within a narrow 300 to 500 ms time window by having the noise burst triggered by the bats initial echolocation pulses as it approached the wire array. Relative to single task controls, echolocation and passive sound localization performance was slightly, but significantly, degraded. The bats also increased echolocation interpulse intervals during the dual task, as though attempting to reduce temporal overlap between the signals. These results suggest that the bats, like humans, have difficulty in processing more than one stream of information at a time.     

Echolocating bats cry out loud to detect their prey

Echolocating bats have successfully exploited a broad range of habitats and prey. Much research has demonstrated how time-frequency structure of echolocation calls of different species is adapted to acoustic constraints of habitats and foraging behaviors. However, the intensity of bat calls has been largely neglected although intensity is a key factor determining echolocation range and interactions with other bats and prey. Differences in detection range, in turn, are thought to constitute a mechanism promoting resource partitioning among bats, which might be particularly important for the species-rich bat assemblages in the tropics. Here we present data on emitted intensities for 11 species from 5 families of insectivorous bats from Panamá hunting in open or background cluttered space or over water. We recorded all bats in their natural habitat in the field using a multi-microphone array coupled with photographic methods to assess the bats' position in space to estimate emitted call intensities. All species emitted intense search signals. Output intensity was reduced when closing in on background by 4-7 dB per halving of distance. Source levels of open space and edge space foragers (Emballonuridae, Mormoopidae, Molossidae, and Vespertilionidae) ranged between 122-134 dB SPL. The two Noctilionidae species hunting over water emitted the loudest signals recorded so far for any bat with average source levels of ca. 137 dB SPL and maximum levels above 140 dB SPL. In spite of this ten-fold variation in emitted intensity, estimates indicated, surprisingly, that detection distances for prey varied far less; bats emitting the highest intensities also emitted the highest frequencies, which are severely attenuated in air. Thus, our results suggest that bats within a local assemblage compensate for frequency dependent attenuation by adjusting the emitted intensity to achieve comparable detection distances for prey across species. We conclude that for bats with similar hunting habits, prey detection range represents a unifying constraint on the emitted intensity largely independent of call shape, body size, and close phylogenetic relationships.     

Active head rolls enhance sonar-based auditory localization performance

Animals utilize a variety of active sensing mechanisms to perceive the world around them. Echolocating bats are an excellent model for the study of active auditory localization. The big brown bat (Eptesicus fuscus), for instance, employs active head roll movements during sonar prey tracking. The function of head rolls in sound source localization is not well understood. Here, we propose an echolocation model with multi-axis head rotation to investigate the effect of active head roll movements on sound localization performance. The model autonomously learns to align the bat’s head direction towards the target. We show that a model with active head roll movements better localizes targets than a model without head rolls. Furthermore, we demonstrate that active head rolls also reduce the time required for localization in elevation. Finally, our model offers key insights to sound localization cues used by echolocating bats employing active head movements during echolocation.     

Prey-Correlated Spectral Changes in Echolocation Sounds of the Indian False Vampire Megaderma lyra

False Vampires ( Megaderma lyra ) are gleaning bats which emit brief (1 ms) and faint echolocation signals consisting of four harmonics of a shallow frequency downward modulated fundamental (27–19 kHz). The complete signal spans a frequency range from 100 to 19 kHz. In sound recordings from three experimental animals we show that Megaderma lyra shifts the dominant frequency in the echolocation signals in relation to the type of prey offered and to flight style. During roaming flights the mean peak frequency was 63.2 ± 9 kHz (third harmonic). In prey catching flights, peak frequencies were shifted into the fourth harmonic. In flights towards a dish of crawling mealworms, mean peak frequency was raised to 91.2 ± 3.3 kHz. When the bats flew towards living mice the dominant frequency was further increased to 99.8 ± 5.2 kHz, and the second and third harmonic were at least 10 dB fainter or no longer recordable. The additional frequency shift when flying towards mice was not only a consequence of the dominance of the fourth harmonic but also of an additional rise of the fundamental harmonic by nearly 2 kHz. These prey-correlated frequency shifts in echolocation calls showed little variation between the three experimental animals and were reproducible over time. They occurred at or even before take-off of the bats. This is the first report of target-correlated transient adaptations in echolocation calls of any bat species.     

Different Auditory Feedback Control for Echolocation and Communication in Horseshoe Bats

Auditory feedback from the animal''s own voice is essential during bat echolocation: to optimize signal detection, bats continuously adjust various call parameters in response to changing echo signals. Auditory feedback seems also necessary for controlling many bat communication calls, although it remains unclear how auditory feedback control differs in echolocation and communication. We tackled this question by analyzing echolocation and communication in greater horseshoe bats, whose echolocation pulses are dominated by a constant frequency component that matches the frequency range they hear best. To maintain echoes within this “auditory fovea”, horseshoe bats constantly adjust their echolocation call frequency depending on the frequency of the returning echo signal. This Doppler-shift compensation (DSC) behavior represents one of the most precise forms of sensory-motor feedback known. We examined the variability of echolocation pulses emitted at rest (resting frequencies, RFs) and one type of communication signal which resembles an echolocation pulse but is much shorter (short constant frequency communication calls, SCFs) and produced only during social interactions. We found that while RFs varied from day to day, corroborating earlier studies in other constant frequency bats, SCF-frequencies remained unchanged. In addition, RFs overlapped for some bats whereas SCF-frequencies were always distinctly different. This indicates that auditory feedback during echolocation changed with varying RFs but remained constant or may have been absent during emission of SCF calls for communication. This fundamentally different feedback mechanism for echolocation and communication may have enabled these bats to use SCF calls for individual recognition whereas they adjusted RF calls to accommodate the daily shifts of their auditory fovea.     

Using echolocation monitoring to model bat occupancy and inform mitigations at wind energy facilities

Fatalities of migratory bats, many of which use low frequency (<35 kHz; LowF) echolocation calls, have become a primary environmental concern associated with wind energy development. Accordingly, strategies to improve compatibility between wind energy development and conservation of bat populations are needed. We combined results of continuous echolocation and meteorological monitoring at multiple stations to model conditions that explained presence of LowF bats at a wind energy facility in southern California. We used a site occupancy approach to model nightly LowF bat presence while accounting for variation in detection probability among echolocation detectors and heights. However, we transposed the spatial and temporal axes of the conventional detection history matrix such that occupancy represented proportion of nights, rather than monitoring points, on which LowF bats were detected. Detectors at 22 m and 52 m above ground had greater detection probabilities for LowF bats than detectors at 2 m above ground. Occupancy of LowF bats was associated with lower nightly wind speeds and higher nightly temperatures, mirroring results from other wind energy facilities. Nevertheless, we found that building separate models for each season and considering solutions with multiple covariates resulted in better fitting models. We suggest that use of multiple environmental variables to predict bat presence could improve efficiency of turbine operational mitigations (e.g., changes to cut-in speeds) over those based solely on wind speed. Increased mitigation efficiencies could lead to greater use of mitigations at wind energy facilities with benefits to bat populations. © 2011 The Wildlife Society.     

The transfer function of a target limits the jitter detection threshold with signals of echolocating FM-bats

The delay jitter discrimination threshold in bats is a disputed subject. Some investigators have obtained results indicating that bats are able to discriminate alternations in delay down to 10 ns, which appears incredible for purely physical reasons. Using actual bat echolocation sequences recorded during an easy detection task to measure simulated delay jitter, it is shown here that jitter detection thresholds in the order of some tens of nanoseconds are actually physically realizable. However, if the transfer function of the target simulating apparatus is not perfect, the lowest thresholds are in the order of hundreds of nanoseconds and variable between individual bats. This phenomenon is shown to arise as a consequence of the variation in signal parameters from call to call. When the transfer function from a real jitter experiment was artificially applied to the echoes, the jitter detection thresholds again were several hundred nanoseconds. This is the first study to point out a limiting role of the transfer function of a system faced with variations in echolocation signal parameters, something that should be considered in evaluating all sonar systems with variable signal structure.     

‘Compromise’ in Echolocation Calls between Different Colonies of the Intermediate Leaf-Nosed Bat (Hipposideros larvatus)

Each animal population has its own acoustic signature which facilitates identification, communication and reproduction. The sonar signals of bats can convey social information, such as species identity and contextual information. The goal of this study was to determine whether bats adjust their echolocation call structures to mutually recognize and communicate when they encounter the bats from different colonies. We used the intermediate leaf-nosed bats (Hipposideros larvatus) as a case study to investigate the variations of echolocation calls when bats from one colony were introduced singly into the home cage of a new colony or two bats from different colonies were cohabitated together for one month. Our experiments showed that the single bat individual altered its peak frequency of echolocation calls to approach the call of new colony members and two bats from different colonies adjusted their call frequencies toward each other to a similar frequency after being chronically cohabitated. These results indicate that the ‘compromise’ in echolocation calls might be used to ensure effective mutual communication among bats.     

A comparison of two bat detectors: which is most likely to detect New Zealand’s Chalinolobus tuberculatus?

ABSTRACT

The presence of bat species is commonly determined by placing acoustic bat detectors that record bat echolocation calls in the habitat they are likely to use. Detection rates are affected by variables including type of detection unit used. We compared detection rates of long-tailed bat (Chalinolobus tuberculatus) echolocation calls between two types of automated bat detectors: Wildlife Acoustics SMZC Zero Crossing Bat Recorders (ZC), and Frequency Compression Automated Bat Monitoring units (FC) produced by New Zealand’s Department of Conservation. Units were placed in locations where bats were known to be present, but not all detected bats. The median number of bat passes recorded by FC units over 10 nights was 20 compared with a median of 3 bat passes for ZC units. ZC units also detected bats over significantly fewer nights than FC units. These results suggest FC units are more sensitive and therefore better to use where long-tailed bats are expected to be at low abundance or only present infrequently. Because of inconsistencies in detection rates, we recommend the use of only one model of the detector within a monitoring project. Our data also suggests that surveys should take place over long periods to maximise likelihood of detecting bats, if present.     

Echolocation calls produced by Kuhl's pipistrelles in different flight situations

Flexibility in the echolocation call structure of bats can improve their performances, because, in some situations, some signal designs are better than others. Hence, at least some bats should adjust their echolocation calls according to the setting in which they are operating but also to the specific task at hand, that is their behavioral intention. We studied variation in the echolocation calls of Pipistrellus kuhlii emitted during four flight situations that were similar in setting but differed in behavioral context: emergence from a roost, commuting to and from foraging sites, foraging and returning to a roost. Echolocation calls produced by P. kuhlii differed significantly according to the flight situation. Call types differed most distinctly between foraging and commuting. We also found a high variance in the emergence calls we recorded, perhaps reflecting pre- and post-takeoff calls. Discriminant function analysis on calls emitted while foraging, commuting or returning to the roost classified the calls to the correct group 73.3% of the time. The differences between bats' echolocation calls in different flight situations might indicate an intrinsic change in the bat's behavior. Recognizing these differences could be crucial when using call variables to identify bat species.