Discrimination performance and echolocation signal integration requirements for target detection and distance determination in the CF/FM bat,Noctilio albiventris

Summary Bats of the speciesNoctilio albiventris were trained to detect the presence of a target or to discriminate differences in target distance by means of echolocation. During the discrimination trials, the bats emitted pairs of pulses at a rate of 7–10/s. The first was an 8 ms constant frequency (CF) signal at about 75 kHz. This was followed after about 28 ms by a short-constant frequency/ frequency modulated (short-CF/FM) signal composed of a 6 ms CF component at about 75 kHz terminating in a 2 ms FM component sweeping downward to about 57 kHz. There was no apparent difference in the pulse structure or emission pattern used for any of the tasks. The orientation sounds of bats flying in the laboratory and hunting prey under natural conditions follow the same general pattern but differ in interesting ways.The bats were able to discriminate a difference in target distance of 13 mm between two simultaneously presented targets and of 30 mm between single sequentially presented targets around an absolute distance of 35 cm, using a criterion of 75% correct responses.The bats were unable to detect the presence of the target or to discriminate distance in the presence of continuous white noise of 54 dB or higher SPL. Under conditions of continuous white noise, the bats increased their pulse repetition rate and the relative proportion of CF/FM pulses.The bats required a minimum of 1–2 successive CF/FM pulse-echo pairs for target detection and 2–3 to discriminate a 5 cm difference in distance. When the distance discrimination tasks were made more difficult by reducing the difference in distance between the two targets the bats needed to integrate information from a greater number of successive CF/FM pulse-echo pairs to make the discrimination.Abbreviations CF constant frequency - FM frequency modulation     

The role of prey-generated sounds,vision, and echolocation in prey localization by the African batCardioderma cor (Megadermatidae)

Summary Cardioderma cor responded with head movements and flight toward speakers broadcasting calls of frogs and crickets which contained only sonic frequencies. Unlike the frog-eating bat,Trachops cirrhosus, they did not make contact with the speakers. Prey movements that generated sonic and ultrasonic sounds were both sufficient and necessary for the bats to localize and capture prey. Prey dragged across a glass sheet with a thin layer of water did not generate sounds and bats did not attempt to capture these prey, even with the availability of visual and echolocation cues. There was no evidence for the use of visual cues while hunting; bats did not localize prey more readily in light than darkness. Prey were presented such that their movements initially generated sounds, but then the prey moved onto the water layer of the glass sheet and sounds were eliminated. The bats emitted echolocation signals while hunting in this situation; however, the information from these signals was not utilized. The bats landed at the site that prey last made sound. These results demonstrate the importance of passive hearing for prey localization in this bat, and further suggest that when preygenerated sounds and echolocation signals offer conflicting information the bat's behavior is guided by the former.     

Passive sound localization of prey by the pallid bat (Antrozous p. pallidus)

Summary The pallid bat (Antrozous p. pallidus) uses passive sound localization to capture terrestrial prey. This study of captive pallid bats examined the roles of echolocation and passive sound localization in prey capture, and focused on their spectral requirements for accurate passive sound localization.Crickets were used as prey throughout these studies. All tests were conducted in dim, red light in an effort to preclude the use of vision. Hunting performance did not differ significantly in red light and total darkness, nor did it differ when visual contrast between the terrestrial prey and the substrate was varied, demonstrating that the bats did not use vision to locate prey.Our bats apparently used echolocation for general orientation, but not to locate prey. They did not increase their pulse emission rate prior to prey capture, suggesting that they were not actively scanning prey. Instead, they required prey-generated sounds for localization. The bats attended to the sound of walking crickets for localization, and also attacked small, inanimate objects dragged across the floor. Stationary and/or anesthetized crickets were ignored, as were crickets walking on substrates that greatly attenuated walking sounds. Cricket communication sounds were not used in prey localization; the bats never captured stationary, calling crickets.The accuracy of their passive sound localization was tested with an open-loop passive sound localization task that required them to land upon an anesthetized cricket tossed on the floor. The impact of a cricket produced a single 10–20 ms duration sound, yet with this information, the bats were able to land within 7.6 cm of the cricket from a maximum distance of 4.9 m. This performance suggests a sound localization accuracy of approximately ±1° in the horizontal and vertical dimensions of auditory space. The lower frequency limit for accurate sound localization was between 3–8 kHz. A physiological survey of frequency representation in the pallid bat inferior colliculus suggests that this lower frequency limit is around 5 kHz.     

Echolocation behaviours of the Japanese pipistrelle batPipistrellus abramus during foraging flight

The acoustic structure of echolocation pulses emitted by Japanese pipistrellePipistrellus abramus (Temminck, 1840) bats during different phases of aerial hawking is described here for the first time. Behavioural observations of the foraging flight in conjunction with acoustical analysis of echolocation pulses indicated a flight path consisting of four distinct phases following the reconnaissance or search phase. Short (∼4.68 ms) and relatively broadband frequencymodulated (FM) pulses (∼23.55 kHz bandwidth) were emitted at a repetition rate of 15 Hz during presumed target approach. Presumed insect capture consisted of an early and a late buzz phase. Both buzz types were emitted at high repetition rates (111 Hz in early to 222 Hz in late) and consisted of very short, broadband FM pulses (1.26 ms in early to 0.3 ms in late). There was also a characteristically sharp drop in both the peak and terminal frequencies of each echolocation pulse during the transition from early to late buzz. No pulses were recorded during the final phase of foraging referred to as a “post-buzz pause”. Thus the foraging behaviour of this species consisted of five sequential phases involving four broad types of echolocation pulses.     

FIELD RECORDINGS OF ECHOLOCATION AND SOCIAL SIGNALS FROM THE GLEANING BAT MYOTIS SEPTENTRIONALIS

ABSTRACT

We recorded echolocation and ultrasonic social signals of the bat Myotis septentrionalis. The bats foraged for insects resting on or fluttering about an outdoor screen to which they were attracted by a ‘backlight’. The bats used nearly linearly modulated echolocation signals of high frequency (117 to 49 kHz, see Tables) with a weak second harmonic. The orientational signals from patrolling bats were about 2.4 ms in duration and occurred at a repetition rate of about 18 Hz (see Figure 3). The signals used by bats as they approached the screen were of shorter duration (0.72 ms) and occurred at higher rates (33.8 Hz) (Table 2 and Figure 4). We registered one feeding ‘buzz’ (Figure 5). We recorded social signals when two bats patrolled the hunting area. The social signals were characterized by their longer durations (6 ms, see Table 1), lower frequencies (70 to 30 kHz), and curvilinear sweeps (Figures 7 and 8). We calculated the source levels of orientational and social signals using the differences in arrival times at three microphones in a linear array (Figures 1 and 2). The source levels were on average 102 dB peSPL at 10 cm (Table 1). We could not calculate source levels of the signals used by bats as they approached the screen at close range, but these signals were much weaker (about 65 dB peSPL at the microphone).     

Echolocation and obstacle avoidance in the hipposiderid batAsellia tridens

Summary The echolocation sounds of the hipposiderid batAsellia tridens consist of a constant frequency (cf) component followed by a frequency modulated (fm) terminal downward sweep of 19–21 kHz. The cf-part constitutes about 7/10 of the entire signal. In individual roosting animals the frequencies of the cf-part of consecutive sounds (resting frequency) is kept very constant but varies from bat to bat. In 18Asellia tridens resting frequencies between 111–124 kHz have been measured.The sound duration in roosting and free flying bats is between 7–10 ms. In the approach and terminal phase of bats landing on a perch or flying through obstacles, the sound duration is reduced and the repetition rate increased the nearer the bat approaches the target. At the end of the terminal phase sound durations of a minimum of 3 ms have been measured. Flying bats lower their emission frequency in order to compensate for Doppler shifts caused by the flight movement. The echofrequency is therefore kept constant about 150–200 Hz above the resting frequency.In flights through obstacles consisting of vertically stretched wires with different diameters, the bats were able to avoid wires down to a diameter of 0.065 mm whereas at 0.05 mm the percentage of flights without collisions is far below the chance level. The results demonstrate that the echolocation behavior of the hipposiderid batAsellia tridens does not differ fundamentally from that of rhinolophid bats. As a result, a new suggestion for categorization of bats producing cf-fm orientation sounds is put forward.Abbreviations cf constant frequency component - fm frequency modulated component - P probability of collision-free flights through an obstacle of ertically tretched wires - I interval between wires - D minimal diameter of a bat with folded wings; , angle at which a bat approaches an obstacle - f _A frequency of the cf-component of the emitted sound - f _E frequency of the cf-component of the echo - f _M frequency of the cf-component of the sounds recorded with the microphone - c speed of sound Supported by the Deutsche Forschungsgemeinschaft grant no. Schn 138/6-9We thank W. Hollerbach for technical assistance.     

THE CLICK-SOUNDS OF NARWHALS (MONODON MONOCEROS) IN INGLEFIELD BAY, NORTHWEST GREENLAND

We studied the sounds of narwhals ( Monodon monoceros ) foraging in the open waters in Northwest Greenland. We used a linear, vertical array of three hydrophones (depth 10 m, 30 m, 100 m) with a fourth hydrophone (depth 30 m) about 20 m from the vertical array. A smaller fifth hydrophone (depth 2 m) allowed for registering frequencies up to 125 kHz (± 2 dB) when signals were recorded at 762 mm/set on an instrumentation tape recorder. Clicks were the prevalent signals, but we heard whistles occasionally. We separated the clicks into two classes: click trains that had rates of 3-10 clicks/sec and click bursts having rates of 110-150 clicks/sec. The spectra of train clicks had maximum amplitudes at 48 ± 10 kHz and a duration of 29 ± 6 psec. The spectra of burst clicks had maximum amplitudes at 19 ± 1 kHz and a duration of 40 ± 3 psec. By analogy with other dolphin species, narwhals presumably use the clicks for echolocation during orientation and for locating prey. The narwhal click patterns resemble those of insectivorous bats. Click trains might correspond to bat searching signals and click bursts to the bat's terminal "buzz", emitted just before prey capture.     

Echolocation behavior of the Japanese horseshoe bat in pursuit of fluttering prey

Echolocation sounds of Rhinolophus ferrumequinum nippon as they approached a fluttering moth (Goniocraspidum pryeri) were investigated using an on-board telemetry microphone (Telemike). In 40?% of the successful moth-capture flights, the moth exhibited distinctive evasive flight behavior, but the bat pursued the moth by following its flight path. When the distance to the moth was approximately 3-4?m, the bats increased the duration of the pulses to 65-95?ms, which is 2-3 times longer than those during landing flight (30-40?ms). The mean of 5.8 long pulses were emitted before the final buzz phase of moth capture, without strengthening the sound pressure level. The mean duration of long pulses (79.9?±?7.9?ms) corresponded to three times the fluttering period of G. pryeri (26.5?×?3?=?79.5?ms). These findings indicate that the bats adjust the pulse duration to increase the number of temporal repetitions of fluttering information rather than to produce more intense sonar sounds to receive fine insect echoes. The bats exhibited Doppler-shift compensation for echoes returning from large static objects ahead, but not for echoes from target moths, even though the bats were focused on capturing the moths. Furthermore, the echoes of the Telemike recordings from target moths showed spectral glints of approximately 1-1.5?kHz caused by the fluttering of the moths but not amplitude glints because of the highly acoustical attenuation of ultrasound in the air, suggesting that spectral information may be more robust than amplitude information in echoes during moth capturing flight.     

Frequency alternation and an offbeat rhythm indicate foraging behavior in the echolocating bat, <Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>

The greater sac-winged bat, Saccopteryx bilineata (Emballonuridae), uses two distinct echolocation call sequences: a ‘monotonous’ sequence, where bats emit ~48 kHz calls at a relatively stable rate, and a frequency-alternating sequence, where bats emit calls at ~45 kHz (low-note call) and ~48 kHz (high-note call). The frequencies of these low–high-note pairs remain stable within sequences. In Panama, we recorded echolocation calls from S. bilineata with a multi-microphone array at two sites: one a known roosting site, the other a known foraging site. Our results indicate that this species (1) only produces monotonous sequences in non-foraging contexts and, at times, directly after emitting a feeding buzz and (2) produces frequency-alternating sequences when actively foraging. These latter sequences are also characterized by an unusual, offbeat emission rhythm. We found significant positive relationships between (1) call intensity and call duration and (2) call intensity and distance from clutter. However, these relationships were weaker than those reported for bats from other families. We speculate on how call frequency alternation and an offbeat emission rhythm might reflect a novel strategy for prey detection at the edge of complex habitat in this ancient family of bats.     

Complex sound analysis in the lesser bulldog bat: evidence for a mechanism for processing frequency elements of frequency modulated signals over restricted time intervals

A stereotypical approach phase vocalization response of the lesser bulldog bat, Noctilio albiventris, to artificial echoes simulating a virtual approaching object was used to assess the ability of the bat to analyze and extract distance information from the artificial echoes. The performance of the bat was not significantly different when presented with naturally structured CF/FM echoes containing FM elements that sweep continuously from about 75-55 kHz in 4 ms or with CF/FM echoes containing FM components constructed from a series of 98 pure tone frequency steps, each with a duration of 0.04 ms. The performance of the bat remained unchanged when the duration of the tone steps was increased up to 0.08 ms but declined sharply to a level that was significantly below that seen with a naturally structured echo when the steps were 0.09 ms or longer. The performance of the bat depended on the duration of the individual tone steps, which could not exceed a specific upper limit of about 0.08 ms. The study suggests that the bats have adaptations for processing individual narrow band segments of FM signals over specific time intervals.Abbreviations CF constant frequency - FM frequency modulation     

Complex sound analysis in the FM bat Eptesicus fuscus,correlated with structural parameters of frequency modulated signals

Big brown bats, Eptesicus fuscus, were presented with artificial frequency modulated (FM) echoes that simulated an object becoming progressively closer to the bat. A stereotyped approach phase behavioral response of the bat to the virtual approaching target was used to determine the ability of the bat to analyze FM signals for target distance information. The degree to which the bats responded with approach phase behavior to a virtual approaching target was similar when they were presented with either a naturally structured artificial FM echo or an artificial FM echo constructed from a series of brief pure tone steps. The ability of the bats to respond to an FM signal structured from a sequence of pure tone elements depended on the number of pure tone steps in the series; the bats required the presentation of tone-step FM signals containing about 83 or greater pure tone elements. Moreover, the duration of the individual tone steps of the tone-step FM signals could not exceed a specific upper limit of about 0.05 ms. Finally, it appears that the bats were able to independently resolve individual tone steps within the tone-step FM signals that were separated by about 450 Hz or more.Abbreviations CF constant frequency - FM frequency modulation     

Adaptive mechanisms underlying the bat biosonar behavior

For survival, bats of the suborder Microchiropetra emit intense ultrasonic pulses and analyze the weak returning echoes to extract the direction, distance, velocity, size, and shape of the prey. Although these bats and other mammals share the common layout of the auditory pathway and sound coding mechanism, they have highly developed auditory systems to process biologically relevant pulses at the expense of a reduced visual system. During this active biosonar behavior, they progressively shorten the pulse duration, decrease the amplitude and pulse-echo gap as they search, approach and finally intercept the prey. Presumably, these changes in multiple pulse parameters throughout the entire course of hunting enable them to extract maximal information about localized prey from the returning echoes. To hunt successfully, the auditory system of these bats must be less sensitive to intense emitted pulses but highly sensitive to weak returning echoes. They also need to recognize and differentiate the echoes of their emitted pulses from echoes of pulses emitted by other conspecifics. Past studies have shown the following mechanical and neural adaptive mechanisms underlying the successful bat biosonar behavior: (1) Forward orienting and highly mobile pinnae for effective scanning, signal reception, sound pressure transformation and mobile auditory sensitivity; (2) Avoiding and detecting moving targets more successfully than stationary ones; (3) Coordinated activity of highly developed laryngeal and middle ear muscles during pulse emission and reception; (4) Mechanical and neural attenuation of intense emitted pulses to prepare for better reception of weak returning echoes; (5) Increasing pulse repetition rate to improve multiple-parametric selectivity to echoes; (6) Dynamic variation of duration selectivity and recovery cycle of auditory neurons with hunting phase for better echo analysis; (7) Maximal multiple-parametric selectivity to expected echoes returning within a time window after pulse emission; (8) Pulse-echo delaysensitive neurons in higher auditory centers for echo ranging; (9) Corticofugal modulation to improve on-going multiple-parametric signal processing and reorganize signal representation, and (10) A large area of the superior colliculus, pontine nuclei and cerebellum that is sensitive to sound for sensori-motor integration. All these adaptive mechanisms facilitate the bat to effectively extract prey features for successful hunting.     

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.     

Target ranging and the role of time-frequency structure of synthetic echoes in big brown bats,Eptesicus fuscus

Summary Echolocating bats judge the distance to a target on basis of the delay between the emitted cry and the returning echo. In a phantom echo set-up it was investigated how changes in the time-frequency structure of synthetic echoes affect ranging accuracy of big brown bats, Eptesicus fuscus.A one channel phantom target simulator and a Y/N paradigm was used. Five Eptesicus fuscus were trained to discriminate between phantom targets with different virtual distances (delays). The phantom echo was stored in a memory and broadcast from a loudspeaker after a certain delay following the bat's triggering of the system via a trigger microphone. The ranging accuracy was compared using 5 different signals with equal energy as phantom echoes: a standard cry (a natural bat cry), two kinds of noise signals, a high pass, and a low pass filtered version of the standard cry.The standard cry was recorded from one of the bats while judging the distance to a real target. The duration was 1.1 ms, the first harmonic swept down from 55 to 25 kHz and there was energy also in the second and third harmonic. Both noise signals had the same duration, power spectrum, and energy as the standard cry. One noise signal was stored in a memory and hence was exactly the same each time the bat triggered the system. The other variable noise signal was produced by storing the envelope of the standard cry and multiplying on-line with band pass filtered noise. The time-frequency structure (e.g. rise time) of this noise signal changed from triggering to triggering. The filtered signals were produced by either 40 kHz high pass or 40 kHz low pass filtering of the standard cry.The range difference thresholds for the 5 bats were around 1–2 cm (51–119 us) using the standard cry as echo. The range difference threshold with both noise signals was 7–8 cm (around 450 s delay difference). The 40 kHz high pass filtered cry increased the threshold to approximately twice the threshold with the standard cry. With the 40 kHz low pass filtered cry the threshold was increased 2.5–3 times relative to the threshold with the standard cry. A single bat was tested with a signal filtered with a 55 kHz low pass filter leaving the whole first harmonic. The threshold was the same as that with the standard signal.The reduced ranging accuracy with the filtered signals indicates that the full band width of the first harmonic is utilised for ranging by the bats. The substantial reduction in accuracy with the noise signals indicates that not only the full band width but also the orderly time-frequency structure (the FM sweep) of the cry is important for ranging in echolocating bats.Abbreviations FM frequency modulated - CF constant frequency - peSPL peak equivalent sound pressure level - SD standard deviation - SE standard error of mean - EPROM erasable programmable read only memory - FFT fast Fourier transform - S/N signal-to-noise ratio     

The adaptive value of increasing pulse repetition rate during hunting by echolocating bats

During hunting, bats of suborder Microchiropetra emit intense ultrasonic pulses and analyze the weak returning echoes with their highly developed auditory system to extract the information about insects or obstacles. These bats progressively shorten the duration, lower the frequency, decrease the intensity and increase the repetition rate of emitted pulses as they search, approach, and finally intercept insects or negotiate obstacles. This dynamic variation in multiple parameters of emitted pulses predicts that analysis of an echo parameter by the bat would be inevitably affected by other co-varying echo parameters. The progressive increase in the pulse repetition rate throughout the entire course of hunting would presumably enable the bat to extract maximal information from the increasing number of echoes about the rapid changes in the target or obstacle position for successful hunting. However, the increase in pulse repetition rate may make it difficult to produce intense short pulse at high repetition rate at the end of long-held breath. The increase in pulse repetition rate may also make it difficult to produce high frequency pulse due to the inability of the bat laryngeal muscles to reach its full extent of each contraction and relaxation cycle at a high repetition rate. In addition, the increase in pulse repetition rate increases the minimum threshold (i.e. decrease auditory sensitivity) and the response latency of auditory neurons. In spite of these seemingly physiological disadvantages in pulse emission and auditory sensitivity, these bats do progressively increase pulse repetition rate throughout a target approaching sequence. Then, what is the adaptive value of increasing pulse repetition rate during echolocation? What are the underlying mechanisms for obtaining maximal information about the target features during increasing pulse repetition rate? This article reviews the electrophysiological studies of the effect of pulse repetition rate on multiple-parametric selectivity of neurons in the central nucleus of the inferior colliculus of the big brown bat, Eptesicus fuscus using single repetitive sound pulses and temporally patterned trains of sound pulses. These studies show that increasing pulse repetition rate improves multiple-parametric selectivity of inferior collicular neurons. Conceivably, this improvement of multiple-parametric selectivity of collicular neurons with increasing pulse repetition rate may serve as the underlying mechanisms for obtaining maximal information about the prey features for successful hunting by bats.     

The tiny difference between foraging and communication buzzes uttered by the Mexican free-tailed bat, <Emphasis Type="Italic">Tadarida brasiliensis</Emphasis>

Echolocating insectivorous bats consummate prey captures using a distinct vocal motor pattern commonly known as the terminal or feeding buzz, which is widely considered a fixed motor pattern executed independently of auditory feedback influences. The Mexican free-tailed bat, Tadarida brasiliensis, offers an opportunity to explore the role of sensory feedback in buzzing because they emit similar buzzes both in flight during foraging and while stationary as communication sounds. Here we compared the spectral and temporal patterns of foraging and communication buzzes to address whether or not auditory feedback may influence buzz patterns. We found that while foraging buzzes uttered in open space were composed of generic FM calls, communication buzzes were composed of an adapted CF–FM call similar to the call type used by T. brasiliensis when navigating in confined spaces. This provides the first evidence that some bats can make significant context-dependent changes in the spectral parameters of calls within their buzz. We also found that inter-pulse intervals, but not call durations, were different within the two buzz types. These observations indicate that though a common pattern generator hierarchically organizes all buzzes, T. brasiliensis retains a significant capacity to adapt the spectral and temporal patterns of elements within its buzzes.     

Sensory ecology of predator–prey interactions: responses of the AN2 interneuron in the field cricket, <Emphasis Type="Italic">Teleogryllus oceanicus</Emphasis> to the echolocation calls of sympatric bats

We observed the responses of the AN2 interneuron in the Pacific field cricket, Teleogryllus oceanicus, a cell implicated in eliciting avoidance flight away from bats, to acoustic stimuli representing the echolocation calls of bats as well as field recordings of search and gleaning attack calls of six species of insectivorous sympatric bats (West Australia, Australia: Tadarida australis, Chalinolobus goudii, Nyctophilus geoffroyi; Queensland, Australia: Vespadelus pumilus, Myotis adversus; Kauai, Hawaii: Lasiurus cinereus). The broad frequency sensitivity of the AN2 cell indicates that T. oceanicus has evolved to detect a wide range of echolocation call frequencies. The reduced sensitivity of this cell at frequencies higher than 70 kHz suggests that some bats (e.g., the gleaning species, N. geoffroyi) may circumvent this insects auditory defences by using frequency-mismatched (allotonic) calls. The calls of the freetail bat, T. australis evoked the strongest response in the AN2 cell but, ironically, this may allow this bat to prey upon T. oceanicus as previous studies report that under certain conditions, flying crickets exhibit ambiguous directional responses towards frequencies similar to those emitted by this bat. Short duration calls (1–2 ms) are sufficient to evoke AN2 responses with instantaneous spike periods capable of causing defensive flight behaviours; most bats tested emit calls of durations greater than this. The short calls of N. geoffroyi produced during gleaning attacks may reduce this species acoustic conspicuousness to this cricket.     

Keeping up with bats: dynamic auditory tuning in a moth

Many night-flying insects evolved ultrasound sensitive ears in response to acoustic predation by echolocating bats . Noctuid moths are most sensitive to frequencies at 20-40 kHz , the lower range of bat ultrasound . This may disadvantage the moth because noctuid-hunting bats in particular echolocate at higher frequencies shortly before prey capture and thus improve their echolocation and reduce their acoustic conspicuousness . Yet, moth hearing is not simple; the ear's nonlinear dynamic response shifts its mechanical sensitivity up to high frequencies. Dependent on incident sound intensity, the moth's ear mechanically tunes up and anticipates the high frequencies used by hunting bats. Surprisingly, this tuning is hysteretic, keeping the ear tuned up for the bat's possible return. A mathematical model is constructed for predicting a linear relationship between the ear's mechanical stiffness and sound intensity. This nonlinear mechanical response is a parametric amplitude dependence that may constitute a feature common to other sensory systems. Adding another twist to the coevolutionary arms race between moths and bats, these results reveal unexpected sophistication in one of the simplest ears known and a novel perspective for interpreting bat echolocation calls.