Variations in the echolocation calls of the European free-tailed bat

The echolocation calls of Tadarida teniotis were studied in an outdoor flight enclosure (captive individuals) and in the wild using single microphones or an array of four microphones. Calls were characterized by measures of 10 call variables. Comparison of individual calls recorded on four microphones arrayed in a tetrahedron with 1 m between each microphone revealed that all calls were not equally detectable by all microphones but that there were no significant differences in call features obtained from calls recorded on all four microphones. A comparison of 47 calls recorded by all four microphones showed no significant differences in the features of the four recordings of each call. Analysis of calls of five individuals flying individually in an outdoor flight cage revealed significant individual differences in call features. In the field, T. teniotis used long, narrowband search-phase calls, usually without harmonics. Analysis of 1876 search-phase echolocation calls of T. teniotis recorded in the field in Israel and Greece in 2002, 2005 and 2006 showed significant year-to-year and site-to-site differences in some call features. When flying in the presence of conspecifics, T. teniotis changed their echolocation calls. We found a range of different buzzes in the wild, and based on their structure we attempted to classify them as feeding and social buzzes. The features of individual calls comprising buzzes differed significantly among buzzes, and yet there were no consistent differences between what we classified as feeding and social buzzes.     

Moth hearing in response to bat echolocation calls manipulated independently in time and frequency

We measured the auditory responses of the noctuid moth Noctua pronuba to bat echolocation calls which were manipulated independently in time and frequency. Such manipulations are important in understanding how insect hearing influences the evolution of echolocation call characteristics. We manipulated the calls of three bat species (Rhinolophus hipposideros, Myotis nattereri and Pipistrellus pipistrellus) that use different echolocation call features by doubling their duration or reducing their frequency, and measured the auditory thresholds from the A1 cells of the moths. Knowing the auditory responses of the moth we tested three predictions. (i) The ranking of the audibility of unmanipulated calls to the moths should be predictable from their temporal and/or frequency structure. This was supported. (ii) Doubling the duration of the calls should increase their audibility by ca. 3 dB for all species. Their audibility did indeed increase by 2.1-3.5 dB. (iii) Reducing the frequency of the calls would increase their audibility for all species. Reducing the frequency had small effects for the two bat species which used short duration (2.7-3.6 ms) calls. However, the relatively long-duration (50 ms), largely constant-frequency calls of R. hipposideros increased in audibility by 21.6 dB when their frequency was halved. Time and frequency changes influence the audibility of calls to tympanate moths in different ways according to call design. Large changes in frequency and time had relatively small changes on the audibility of calls for short, largely broadband calls. Channelling energy into the second harmonic of the call substantially decreased the audibility of calls for bats which use long-duration, constant-frequency components in echolocation calls. We discuss our findings in the contexts of the evolution of both bat echolocation call design and the potential responses of insects which hear ultrasound.     

Extremely high frequency sensitivity in a ‘simple’ ear

An evolutionary war is being played out between the bat, which uses ultrasonic calls to locate insect prey, and the moth, which uses microscale ears to listen for the approaching bat. While the highest known frequency of bat echolocation calls is 212 kHz, the upper limit of moth hearing is considered much lower. Here, we show that the greater wax moth, Galleria mellonella, is capable of hearing ultrasonic frequencies approaching 300 kHz; the highest frequency sensitivity of any animal. With auditory frequency sensitivity that is unprecedented in the animal kingdom, the greater wax moth is ready and armed for any echolocation call adaptations made by the bat in the on-going bat–moth evolutionary war.     

Support for the allotonic frequency hypothesis in an insectivorous bat community

The allotonic frequency hypothesis proposes that certain insectivorous bat species can prey upon moths that can hear bat echolocation calls by using echolocation frequencies outside the sensitivity range of moth ears. The hypothesis predicts that the peak frequencies of bat echolocation calls are correlated with the incidence of moths in the diets of these bats. The aim of this study was to test this prediction on a bat community dominated by bats using low duty cycle echolocation calls, i.e. aerial foraging, insectivorous species using frequency modulated calls. The community consisted of nine species, two molossids, Sauromys petrophillus and Tadarida aegyptiaca, five vespertilionids, Eptesicus capensis, Eptesicus hottentotus, Miniopteris schreibersii, Myotis tricolor, and Myotis lesueuri, one rhinolophid, Rhinolophus clivosus, and one nycterid, Nycteris thebaica. The insect fauna in the habitat used by the bat community was suited to the testing of the allotonic frequency hypothesis because more than 90% of the moths comprising the insect fauna were tympanate. These included Pyralidae (3.8%), Geometridae (44.9%), Notodontidae (3.8%), Arctiidae (4.6%), Lymantriidae (0.8%) and Noctuidae (32.4%). As predicted, peak echolocation frequency was correlated with the incidence of moths in the diets of these nine species (r=0.98, df=7, P<0.01). Furthermore, multivariate analysis revealed that echolocation frequency (t=9.91, n=129, P<0.001) was a better predictor of diet than forearm length (t=5.51, n=129, P<0.001) or wing area (t=-3.41, n=129, P<0.001). This suggests that the selection pressure exerted by moth hearing might have acted directly on call frequency and secondarily on body size and wing morphology, as part of the same adaptive complex. It is unlikely that dietary differences were due to temporal and spatial differences in the availability of prey because the pattern of differences in skull morphology of the nine species supported our dietary analyses. The skull morphology of a bat represents a historical record of the kind of diet it has become adapted to over its evolutionary history. These results suggest that prey defences may mediate other factors structuring bat communities, e.g. competition. Competition may be reduced for those species of bats that can circumvent prey defences.     

Scanning Behavior in Echolocating Common Pipistrelle Bats (Pipistrellus pipistrellus)

Echolocating bats construct an auditory world sequentially by analyzing successive pulse-echo pairs. Many other mammals rely upon a visual world, acquired by sequential foveal fixations connected by visual gaze saccades. We investigated the scanning behavior of bats and compared it to visual scanning. We assumed that each pulse-echo pair evaluation corresponds to a foveal fixation and that sonar beam movements between pulses can be seen as acoustic gaze saccades. We used a two-dimensional 16 microphone array to determine the sonar beam direction of succeeding pulses and to characterize the three dimensional scanning behavior in the common pipistrelle bat (Pipistrellus pipistrellus) flying in the field. We also used variations of signal amplitude of single microphone recordings as indicator for scanning behavior in open space. We analyzed 33 flight sequences containing more than 700 echolocation calls to determine bat positions, source levels, and beam aiming. When searching for prey and orienting in space, bats moved their sonar beam in all directions, often alternately back and forth. They also produced sequences with irregular or no scanning movements. When approaching the array, the scanning movements were much smaller and the beam was moved over the array in small steps. Differences in the scanning pattern at various recording sites indicated that the scanning behavior depended on the echolocation task that was being performed. The scanning angles varied over a wide range and were often larger than the maximum angle measurable by our array. We found that echolocating bats use a “saccade and fixate” strategy similar to vision. Through the use of scanning movements, bats are capable of finding and exploring targets in a wide search cone centered along flight direction.     

马铁菊头蝠幼蝠能够辨别种群间回声定位声波差异

蝙蝠的回声定位声波普遍存在地理变化,然而尚不清楚幼蝠在扩散前能否辨别不同种群间的声波差异。本文采用"双选择声学回放实验",研究马铁菊头蝠(Rhinolophus ferrumequinum)幼蝠(1月龄)对回声定位声波地理变化的辨别能力及行为反应。采用4个行为参数衡量幼蝠对本种群(CC)与外种群(JN)回声定位声波的行为反应:每一回放室飞入的次数、飞行时间、停留次数、探测时间。Mann-Whitney U tests表明,马铁菊头蝠1月龄个体对回放声波的行为反应,除每一回放室停留的次数外,其他3个参数均存在显著差异(P0.05),且在本种群声波回放室(CC)的飞行次数和时间以及探测时间均高于外种群声波回放室(JN)。对4个行为参数进行主成分分析,Mann-Whitney U tests表明,回放本种群声波(CC)与外种群声波(JN)之间,第一主成分得分(PC1)存在显著差异(P0.01)。配对T检验表明,幼蝠对本种群声波反应更加强烈(PC1:t10=5.25,P0.001;PC2:t10=2.34,P0.05)。本研究说明马铁菊头蝠幼蝠能够辨别不同种群间回声定位声波的差异。     

在不同状态和生境复杂度中大蹄蝠回声定位叫声的可塑性

蝙蝠回声定位声波的可塑性对其适应不同状态、生境以及捕食和社会交流具有重要的作用。为进一步研究大蹄蝠的回声定位声波在不同状态和生境下的可塑性,通过室内行为实验,对大蹄蝠在4 种不同状态(室内飞行、静息、布袋内和手持)和4 种不同生境复杂度(室外、室内0 棵树、室内1 棵树、室内5 棵树)条件下飞行的回声定位声波特征进行研究。结果表明:大蹄蝠的回声定位声波为CF - FM 型,通常连续发出2 - 4 个脉冲组成一个脉冲组。对比大蹄蝠在4 种不同状态下的回声定位叫声发现,主频按静息、布袋内、手持、飞行的顺序依次降低,后端FM 频宽则按手持、布袋内、飞行和静息的顺序依次变短;而脉冲间隔和脉冲时程则均按静 息、飞行、布袋内、手持的顺序依次增加。对比大蹄蝠在4 种不同生境复杂度中飞行的回声定位叫声发现,主频按室外、室内0 棵树、室内1 棵树、室内5 棵树依次提高,而脉冲时程及脉冲间隔则依次缩短;室外放飞条件下的后端FM 频宽比室内飞行的短。研究结果说明,大蹄蝠在不同状态、不同生境复杂度条件下的回声定位叫声具有明显的可塑性和生境适应性。     

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.     

Echolocation range and wingbeat period match in aerial-hawking bats

Aerial-hawking bats searching the sky for prey face the problem that flight and echolocation exert independent and possibly conflicting influences on call intervals. These bats can only exploit their full echolocation range unambiguously if they emit their next call when all echoes from the preceding call would have arrived. However, not every call interval is equally available. The need to reduce the high energetic costs of echolocation forces aerial-hawking bats to couple call emission to their wingbeat. We compared the wingbeat periods of 11 aerial-hawking bat species with the delays of the last-expected echoes. Acoustic flight-path tracking was employed to measure the source levels (SLs) of echolocation calls in the field. SLs were very high, extending the known range to 133 dB peak equivalent sound pressure level. We calculated the maximum detection distances for insects, larger flying objects and background targets. Wingbeat periods were derived from call intervals. Small and medium-sized bats in fact matched their maximum detection range for insects and larger flying targets to their wingbeat period. The tendency to skip calls correlated with the species' detection range for background targets. We argue that a species' call frequency is at such a pitch that the resulting detection range matches their wingbeat period.     

Flight performance, echolocation and foraging behaviour in noctule bats Nyctalus noctula

The noctule Nyctulus noctula (Schreber, 1774) is a relatively large (c. 25 g) insectivorous bat which catches insects on the wing (by aerial hawking). Emergence at a maternity roost was earliest relative to sunset when females were lactating, and bats may then have risked predation by flying at higher light levels during a period of high energy demand. Flight performance was quantified by using stereophotogrammetry. At feeding sites bats flew at 6.0 ± 2.1 m/s. This was faster than predicted minimum power speed (V _mp), and either between V _mp and maximum range speed (V _mr), or close to their predicted V _mr, depending on which aerodynamic model of flight power requirements was used. The echolocation behaviour of noctules is flexible. Long duration, low frequency calls (c. 20 kHz) with little frequency modulation were emitted while cruising, but at foraging sites the calls became more frequency-modulated. As the noctule is traditionally thought of as using low frequency echolocation, it was expected to receive weak echoes from small targets and therefore to specialize in eating large insect prey. Although the bats ate mainly beetles, large numbers of small dipterans were also eaten. The noctule is probably able to detect such small items because, when foraging, its calls become broadband and sweep from high frequencies. Higher harmonics are also present, and these may assist in the detection of small prey. In noctules, as in many bats, there appears to be a 1:1 link between wingbeat and call production during the search phase of foraging.     

普通长翼蝠食性结构及其回声定位与体型特征

在普通长翼蝠(Miniopterus fuliginosus)的捕食区内用灯诱法和网捕法调查潜在食物(昆虫)种类; 用粪便分析法鉴定普通长翼蝠的食物组成,发现其主要捕食体型较大的鳞翅目和鞘翅目昆虫,体积百分比分别为55%和38%.普通长翼蝠具有相对狭长的翼,翼展比为6.94 ± 0.13;翼载为(9.85 ± 0.83)N/m2,相对较大.飞行状态下普通长翼蝠的回声定位叫声为调频下扫型,声脉冲时程为(1.45 ± 0.06)ms,脉冲间隔为(63.08 ± 21.55)ms,主频较低,为(44.50 ± 2.26)kHz.研究表明,普通长翼蝠的形态特征和回声定位特征与其捕食行为有着密切的联系.     

南蝠回声定位叫声的分析

蝙蝠科是翼手目中种类最繁多、分布最广泛、进化最成功的科之一 ,全球共有 42属 35 5种(Ｎｏｗａｋ ,1991)。该类群的大多数物种都以超声波回声定位来进行捕食 ,其回声定位行为的多样性以及捕食策略的多样性 ,一直都是动物生态学中的研究热点。南蝠 (Ｉａｉｏ)属蝙蝠科南蝠属 ,为单型种 ,主要分布于我国 (罗蓉等 ,1993)。它是蝙蝠科中体形最大者 ,以前对其生态学方面的研究非常少 ,而对其回声定位的研究则未见报道。南蝠捕食时的叫声与飞行及悬挂状态下的叫声的基本特征一致 (声谱图及谐波等 ) ,仅在叫声次数上有一定差异。因此本文将录制南…     

The Divergence of Echolocation Frequency in Horseshoe Bats: Moth Hearing,Body Size or Habitat?

A phylogenetic approach was used to test three hypotheses regarding the evolution of diversity in the echolocation frequencies used by horseshoe bats (family Rhinolophidae, genus Rhinolophus): 1) Allotonic Frequency Hypothesis (high frequency echolocation in the Rhinolophidae resulted from coevolution with moth hearing); 2) Allometry Hypothesis (echolocation frequency is negatively scaled with body size and evolutionary changes in echolocation frequencies are correlated with changes in body size in the Rhinolophidae); and 3) Foraging Habitat Hypothesis (evolution of echolocation frequency is associated with changes in habitat type). Both discrete and continuous character sets were used for ancestral state reconstructions and for investigating patterns of evolution between frequency and body size, and frequency and habitat type. Contrary to the prediction of the Allotonic Frequency Hypothesis, echolocation frequency in the Rhinolophidae did not increase over time, which would be expected if moth hearing and bat echolocation frequency coevolved. The number of extant species that exhibit calls within moth hearing ranges was not significantly different from the number of species that echolocate outside of moth hearing range. There was also no correlation between changes in frequency and changes in habitat type as predicted by the Foraging Habitat Hypothesis. Instead, the evolution of echolocation frequency within the Rhinolophidae was correlated with changes in body size as predicted by the Allometry Hypothesis.     

Bat echolocation calls facilitate social communication

Bat echolocation is primarily used for orientation and foraging but also holds great potential for social communication. The communicative function of echolocation calls is still largely unstudied, especially in the wild. Eavesdropping on vocal signatures encoding social information in echolocation calls has not, to our knowledge, been studied in free-living bats so far. We analysed echolocation calls of the polygynous bat Saccopteryx bilineata and found pronounced vocal signatures encoding sex and individual identity. We showed experimentally that free-living males discriminate approaching male and female conspecifics solely based on their echolocation calls. Males always produced aggressive vocalizations when hearing male echolocation calls and courtship vocalizations when hearing female echolocation calls; hence, they responded with complex social vocalizations in the appropriate social context. Our study demonstrates that social information encoded in bat echolocation calls plays a crucial and hitherto underestimated role for eavesdropping conspecifics and thus facilitates social communication in a highly mobile nocturnal mammal.     

‘No cost of echolocation for flying bats’ revisited

Echolocation is energetically costly for resting bats, but previous experiments suggested echolocation to come at no costs for flying bats. Yet, previous studies did not investigate the relationship between echolocation, flight speed, aerial manoeuvres and metabolism. We re-evaluated the 'no-cost' hypothesis, by quantifying the echolocation pulse rate, the number of aerial manoeuvres (landings and U-turns), and the costs of transport in the 5-g insectivorous bat Rhogeessa io (Vespertilionidae). On average, bats (n = 15) travelled at 1.76 ± 0.36 m s?1 and performed 11.2 ± 6.1 U-turns and 2.8 ± 2.9 ground landings when flying in an octagonal flight cage. Bats made more U-turns with decreasing wing loading (body weight divided by wing area). At flight, bats emitted 19.7 ± 2.7 echolocation pulses s?1 (range 15.3-25.8 pulses s?1), and metabolic rate averaged 2.84 ± 0.95 ml CO? min?1, which was more than 16 times higher than at rest. Bats did not echolocate while not engaged in flight. Costs of transport were not related to the rate of echolocation pulse emission or the number of U-turns, but increased with increasing number of landings; probably as a consequence of slower travel speed when staying briefly on ground. Metabolic power of flight was lower than predicted for R. io under the assumption that energetic costs of echolocation call production is additive to the aerodynamic costs of flight. Results of our experiment are consistent with the notion that echolocation does not add large energetic costs to the aerodynamic power requirements of flight in bats.     

Understanding signal design during the pursuit of aerial insects by echolocating bats: tools and applications

Bats are among the few predators that can exploit the large quantities of aerial insects active at night. They do this by using echolocation to detect, localize, and classify targets in the dark. Echolocation calls are shaped by natural selection to match ecological challenges. For example, bats flying in open habitats typically emit calls of long duration, with long pulse intervals, shallow frequency modulation, and containing low frequencies-all these are adaptations for long-range detection. As obstacles or prey are approached, call structure changes in predictable ways for several reasons: calls become shorter, thereby reducing overlap between pulse and echo, and calls change in shape in ways that minimize localization errors. At the same time, such changes are believed to support recognition of objects. Echolocation and flight are closely synchronized: we have monitored both features simultaneously by using stereo photogrammetry and videogrammetry, and by acoustic tracking of flight paths. These methods have allowed us to quantify the intensity of signals used by free-living bats, and illustrate systematic changes in signal design in relation to obstacle proximity. We show how signals emitted by aerial feeding bats can be among the most intense airborne sounds in nature. Wideband ambiguity functions developed in the processing of signals produce two-dimensional functions showing trade-offs between resolution of time and velocity, and illustrate costs and benefits associated with Doppler sensitivity and range resolution in echolocation. Remarkably, bats that emit broadband calls can adjust signal design so that Doppler-related overestimation of range compensates for underestimation of range caused by the bat's movement in flight. We show the potential of our methods for understanding interactions between echolocating bats and those prey that have evolved ears that detect bat calls.     

Adaptive auditory risk assessment in the dogbane tiger moth when pursued by bats

Moths and butterflies flying in search of mates risk detection by numerous aerial predators; under the cover of night, the greatest threat will often be from insectivorous bats. During such encounters, the toxic dogbane tiger moth, Cycnia tenera uses the received intensity, duration and emission pattern of the bat''s echolocation calls to determine when, and how many, defensive ultrasonic clicks to produce in return. These clicks, which constitute an acoustic startle response, act as warning signals against bats in flight. Using an integrated test of stimulus generalization and dishabituation, here we show that C. tenera is able to discriminate between the echolocation calls characteristic of a bat that has only just detected it versus those of a bat actively in pursuit of it. We also show that C. tenera habituates more profoundly to the former stimulus train (‘early attack’) than to the latter (‘late attack’), even though it was initially equally responsive to both stimuli. Matched sensory and behavioural data indicate that reduced responsiveness reflects habituation and is not merely attributable to sensory adaptation or motor fatigue. In search of mates in the face of bats, C. tenera''s ability to discriminate between attacking bats representing different levels of risk, and to habituate less so to those most dangerous, should function as an adaptive cost–benefit trade-off mechanism in nature.     

Directional sensitivity of echolocation in the horseshoe bat,Rhinolophus ferrumequinum

Summary The directionality of sound emission by a horseshoe bat (Rhinolophus ferrumequinum) has been determined for the constant frequency component of its orientation sounds. The bat was fixed in the center of an acoustic perimeter and the SPL of the orientation sounds measured with a scanning microphone at different angles compared with the SPL measured by another microphone located in the direction perpendicular to the plane of the horseshoe-like structure of the nose-leaf. The maximum SPL was always found in this direction which also corresponds to the flight direction of a bat in horizontal flight. Above and lateral to this direction the SPL decreases steadily with -6 dB-points at 24 above and 23 lateral. Below the flight direction we found a prominent side lobe with a -6 dB-point at 64 .When the present data are combined with measurements of the behavioral directionality of hearing at the same frequency (Grinnell and Schnitzler, 1977), the directionality diagram of the entire echolocation system is very narrow and points in the flight direction. The prominent downward side lobe of emission does not conspicuously increase echolocation effectiveness in the direction of the ground, since hearing sensitivity is falling off so steeply in that direction. However, without this downward beam of emission, signals from below the bat would be that much less effective.Interference with the structure of the nose-leaf by covering the upper part with vaseline or plugging the left nostril destroyed the smoothness of the normal sound field and demonstrated that this complex organ is a highly functional structure optimized in the course of evolution.With differences in mood or attention, the emitted pulses varied by as much as 20 dB (80–100 dB). The emission directionality pattern also varied. In most cases, as orientation sounds increased in SPL, the acoustic beam became smaller.Supported by Deutsche Forschungsgemeinschaft, grant No. Schn 138/1-6, Stiftung Volkswagenwerk, grant No. 111 858, and the Alexander von Humboldt StiftungWe thank W. Hollerbach and C. Nitsche for technical assistance.     

Gleaning bat echolocation calls do not elicit antipredator behaviour in the Pacific field cricket, <Emphasis Type="Italic">Teleogryllus oceanicus</Emphasis> (Orthoptera: Gryllidae)

Bats that glean prey (capture them from surfaces) produce relatively inconspicuous echolocation calls compared to aerially foraging bats and could therefore be difficult predators to detect, even for insects with ultrasound sensitive ears. In the cricket Teleogryllus oceanicus, an auditory interneuron (AN2) responsive to ultrasound is known to elicit turning behaviour, but only when the cricket is in flight. Turning would not save a cricket from a gleaning bat so we tested the hypothesis that AN2 elicits more appropriate antipredator behaviours when crickets are on the ground. The echolocation calls of Nyctophilus geoffroyi, a sympatric gleaning bat, were broadcast to singing male and walking female T. oceanicus. Males did not cease singing and females did not pause walking more than usual in response to the bat calls up to intensities of 82 dB peSPL. Extracellular recordings from the cervical connective revealed that the echolocation calls elicited AN2 action potentials at high firing rates, indicating that the crickets could hear these stimuli. AN2 appears to elicit antipredator behaviour only in flight, and we discuss possible reasons for this context-dependent function.     

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.