A gating mechanism for sound pattern recognition is correlated with the temporal structure of echolocation sounds in the rufous horseshoe bat

Summary The rufous horseshoe bat, Rhinolophus rouxi, was trained to discriminate differences in target distance. Loud free running artificial pulses, simulating the bat's natural long-CF/FM echolocation sounds, interfered with the ability of the bat to discriminate target distance. Interference occurred when the duration of the CF component of the CF/FM artificial pulse was between 2 and 70 ms. A brief (2.0 ms) CF signal 2–68 ms before an isolated FM signal was as effective as a continuous CF component of the same duration. When coupled with the bat's own emissions, a 2 ms FM sweep alone was effective in interfering when it came 42 to 69 ms after the onset of the bat's pulse. The coupled FM artificial pulses did not interfere when they began during the bat's own emissions.It appears that the onset of the CF component activates a gating mechanism that establishes a time window during which FM component signals must occur for proper neural processing. A comparison with a similar gating mechanism in Noctillo albiventris, which emits short-CF/FM echolocation sounds, reveals that the temporal parameters of the time window of the gating mechanism are species specific and specified by the temporal structure of the echolocation sound pattern of each species.Abbreviations FM frequency modulated - CF constant frequency     

Discrimination of fluttering targets by the FM-batPipistrellus stenopterus?

Summary Five bats of the speciesPipistrellus stenopterus were trained in a two-alternative forced-choice procedure to discriminate between two fluttering targets. The positive target simulated an insect with a 50 Hz wingbeat rate. The negative target was varied between 0 and 48 Hz.The bats were able to discriminate a target with 41 Hz from a target with 50 Hz with 75% correct choices. In the discrimination task, they typically emitted echolocation calls of 2–4 ms duration sweeping from 60 kHz to 30 kHz. The duty cycle (i.e. fraction of time filled with echolocation sounds) increased when the targets fluttered, but was always lower than 3%.The performance ofP. stenopterus in discriminating fluttering targets is comparable to that of bats emitting longer sounds with constant-frequency (CF) components and a higher duty cycle. The FM-sounds ofP. stenopterus are short compared with the period of the fluttering targets, and therefore make it difficult for the animal to measure the time interval between two acoustic glints. Other cues may be prominent, such as the frequency modulation by Doppler shifts from the moving blades.     

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     

Frequency tracking and Doppler shift compensation in response to an artificial CF/FM echolocation sound in the CF/FM bat,Noctilio albiventris

Summary Bats of the speciesNoctilio albiventris emit short-constant frequency/frequency modulated (short-CF/FM) pulses with a CF component frequency at about 75 kHz. Bats sitting on a stationary platform were trained to discriminate target distance by means of echolocation. Loud, free-running artificial pulses, simulating the bat's natural CF/FM echolocation sounds or with systematic modifications in the frequency of the sounds, were presented to the bats during the discrimination trials. When the CF component of the artificial CF/FM sound was between 72 and 77 kHz, the bats shifted the frequency of the CF component of their own echolocation sounds toward that of the artificial pulse, tracking the frequency of the artificial CF component.Bats flying within a large laboratory flight cage were also presented with artificial pulses. Bats in flight lower the frequency of their emitted pulses to compensate for Doppler shifts caused by their own flight speed and systematically shift the frequency of their emitted CF component so that the echo CF frequency returns close to that of the CF component of the artificial CF/FM pulse, over the frequency range where tracking occurs.Abbreviations CF constant frequency - FM frequency modulation     

Discrimination of insect wingbeat-frequencies by the batRhinolophus ferrumequinum

Summary Five Greater Horseshoe bats,Rhinolophus ferrumequinum, were trained in a two-alternative forced-choice procedure to discriminate between artificial echoes of insects fluttering at different wingbeat rates. The stimuli were electronically produced phantom targets simulating fluttering insects with various wingbeat frequencies (Figs. 3, 4). Difference thresholds for wingbeat rates of 50 Hz and 100 Hz were determined. For an S+ of 50 Hz the difference threshold values lay between 2.8 and 4.6 Hz for individual bats; with an S+ of 100 Hz they increased to between 9.8 and 12.0 Hz (Figs. 5, 6, Table 1).Three bats, previously trained to discriminate between a S+ of 50 Hz and a S– with a lower wingbeat rate, were tested with higher frequency stimuli. When they had to decide between their old S+ of 50 Hz and either a 60 or 70 Hz echo two bats continued to select the 50 Hz stimulus while the third bat now preferred the faster fluttering insects (Table 2).During the discrimination task the echolocation behavior of the bats was monitored. When the phantom targets were presented all bats increased their duty-cycle of sound emission from about 40% to sometimes near 70%. They did so by either emitting longer echolocation calls or by increasing the sound repetition rate (Figs. 7, 8).The results show that Greater Horseshoe bats can determine the wingbeat rate of flying insects with an accuracy between 6 and 12%. Possible cues for flutter rate determination by cf-fm bats from natural and artificial insect echoes are discussed.Abbreviations DC duty-cycle - PD pulse duration - PI pulse interval - cf constantfrequency - fm frequency modulation     

Fluttering target detection in Hipposiderid bats

Summary Two species of Hipposiderid bats,Hiposideros speoris andH. lankadiva, which both emit short CF-FM echolocation calls, were trained in a two-alternative forced-choice procedure to discriminate between an oscillating target and a motionless one. Two different targets were used: (1) the membrane of a low-frequency loudspeaker, producing sinusoidal frequency- and amplitude modulations and (2) a small rotating propeller, which produced short acoustical glints. In both casesH. lankadiva learned to discriminate between the oscillating and the motionless target. When the loudspeaker was used, thresholds for minimal modulation depths at different oscillation frequencies were determined. At loudspeaker membrane oscillation frequencies of 10 to 100 Hz the 75% correct thresholds lay between 90 and 300 Hz (Fig. 3).H. speoris could not be trained to react to the moving membrane, even at very high oscillation amplitudes. When the rotating propeller was the positive target, however,H. speoris learned very quickly to discriminate it from a motionless one. By decreasing the rotation speed it was possible to measure the minimal detectable glint-frequency for each bat. It lay at 67, 44, and just under 15 glints/s for the three specimens (Fig. 4). During the discrimination task both bat species increased their duty-cycle just prior to a decision by emitting long sequences of echolocation calls with short inter-pulse intervals. The duration of individual pulses remained relatively constant (Figs. 6 and 7). Possible mechanisms for discrimination of the oscillating targets are discussed and the importance of glints in the echoes for fluttering target detection is emphasized.Abbreviations A₀ amplitude of loudspeaker membrane oscillation - AM amplitude modulation - CF constant frequency - DC duty-cycle - FM frequency modulation - f _max maximal frequency modulation depth in echo - PD pulse duration - PI pulse interval - f _osc oscillation frequency     

Echolocation by the long-eared bat,Plecotus phyllotis

Summary The echolocating bat,Plecotus phyllotis (Vespertilionidae), uses long-CF/FM and FM sonar sounds in different situations. The CF component in long-CF/FM sounds occurs at 27 kHz and has a duration of 20 to 200 ms. The FM component sweeps down from 24 to 12 kHz, with a prominent second harmonic from 40 to 22 kHz. This second harmonic sweep is interrupted at 28 to 25 kHz, providing a notch in the spectrum of the FM component at the CF frequency. This notch probably permits isolation of CF and FM components in echoes for separate processing, thus avoiding mutual interference with the different kinds of target information the two components convey. The FM component is also used without the CF component as a sonar sound. Two other FM orientation sounds are used when the bat is in a confined space such as a room. One contains only the second and fourth harmonics of the 24 to 12 kHz fundamental sweep, while the other contains only the fifth harmonic. This bat's repertoire of sonar sounds closely matches the hearing capacities of the genus.We thank P.H. Dolkart and W.A. Lavender, of Washington University, and the Nevada State Parks Department for their assistance. This research was supported by Grant # BMS-72-02351-A01 from the National Science Foundation.     

Echolocation by free-tailed bats (Tadarida)

Summary The echolocation of bats in the genusTadarida is highly adaptive to different acoustic conditions. These bats use different types of sonar signals with a diversity usually observed in comparisons across families of bats.Tadarida brasiliensis andT. macrotis search for airborne prey in open, uncluttered spaces using narrow-band, short CF signals with no FM components. They add broadband FM components while dropping the CF components when approaching or capturing prey. Only one harmonic is present in these insect-pursuit signals. When flying in cluttered situations or echolocating in a laboratory room,T. brasiliensis uses multiple-harmonic FM signals. Stationary bats tend to use linear frequency sweeps and moving bats tend to use curvilinear frequency sweeps or linear period sweeps. When emerging from a roost they initially use a short-CF/FM signal, changing to an FM signal as they fly away. The acuity of perception of target range inT. brasiliensis is about 1.0 to 1.5 cm and is determined by the bandwidth of the target-ranging sonar signals as represented by their autocorrelation functions. Many less adaptable species of bats use signals corresponding to part of the sonar repertoire ofTadarida. The functions of short CF or narrowband signals for detection and FM or broadband signals for resolution and acoustic imaging identified from comparisons among such species are confirmed by observations of echolocation byTadarida. The differences observed in echolocation among many species and families of bats appear to be evolutionary adaptations to some of the same features of the acoustic environment to whichTadarida responds behaviorally.Abbreviations CF frequency modulated - FM constant frequency - LPM linear period modulation - LFM linear-frequency modulation We thank Prof. T.T. Sandel, Prof. D.R. Griffin, Dr. George Pollak, and P.H. Dolkart for their advice and assistance. This research was supported by Grant No. BMS 72-02351-A01 from the National Science Foundation and by Biomedical Research Support Grant No. RR-07054 from the Division of Research Resources, National Institutes of Health.     

Harmonic and frequency structure used for echolocation sound pattern recognition and distance information processing in the rufous horseshoe bat

Summary The rufous horseshoe bat, Rhinolophus rouxi, was trained to discriminate differences in target distance. During the discrimination trials, the bats emitted complex FM/CF/FM pulses containing first harmonic and dominant second harmonic components.Loud free running artificial pulses, simulating the CF/FM part of the natural echolocation components, interfered with the ability of the bat to discriminate target distance. Changes in the frequency or frequency pattern of the artificial pulses resulted in systematic changes in the degree of interference. Interference occurred when artificial CF/FM pulses were presented at frequencies near those of the bat's own first or second harmonic components.These findings suggest that Rhinolophus rouxi uses both the first and second harmonic components of its complex multiharmonic echolocation sound for distance discrimination. For interference to occur, the sound pattern of each harmonic component must contain a CF signal followed by an FM sweep beginning near the frequency of the CF.Abbreviations CF constant frequency - FM frequency modulated     

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.     

Echolocation sound features processed to provide distance information in the CF/FM bat,Noctilio albiventris: evidence for a gated time window utilizing both CF and FM components

Summary Bats of the speciesNoctilio albiventris, trained to discriminate differences in target distance, emitted pairs of pulses at a rate of 7–10/s, the first a constant frequency (CF) pulse of about 8 ms duration and 75 kHz frequency, followed after about 28 ms by a CF/FM pulse having a 6 ms, 75 kHz CF component that terminates in a 2 ms FM sweep to about 57 kHz.Loud free-running artificial pulses, simulating the bat's natural CF/FM echolocation sound, interfered with distance discrimination at repetition rates exceeding 5/s. Systematic modifications in the temporal and frequency structure of the artificial pulses resulted in orderly changes in the degree of interference. Artificial pulses simulating the natural CF or FM components alone had no effect, nor did 10/s white noise pulses, although constant white noise of the same intensity masked the behavior.Interference occurred when the CF of the artificial pulses was between 52 and 77 kHz, ending with a downward FM sweep of 25 kHz from the CF. For interference to occur there was a much more critical requirement that the FM sweep begin at approximately the frequency of the CF component. The FM sweep needed to be 11 kHz or greater bandwidth. Interference occurred when the duration of the CF component of the CF/FM artificial pulse was between 2 and 30 ms, with maximal effect between 10 and 20 ms. However, a brief (2.0 ms) CF signal 2–27 ms before an isolated FM signal was as effective as a continuous CF component of the same duration.When coupled with the bat's own emissions, artificial CF/FM pulses interfered if they occurred after the bat's CF/FM pulse and before the next natural emission. A 2 ms FM sweep alone was effective in interfering with distance discrimination when it came 8–27 ms after the onset of the bat's own CF/FM pulse. Neither CF/FM nor FM artificial pulses interfered when they began during the bat's own emission. A 10 ms CF pulse alone had no effect at any time.These findings indicate thatN. albiventris uses both the CF and FM components of its short-CF/FM echolocation sound for distance discrimination. The CF onset activates a gating mechanism that, during a narrowly defined subsequent time window, enables the nervous system to process FM pulse-echo pairs for distance information, within a fairly broad frequency range, as long as the frequencies of the CF and the beginning of the FM sweep are nearly identical.Abbreviations CF constant frequency - FM frequency modulation     

普氏蹄蝠(Hipposideros pratti)回声定位声波、形态及捕食策略

研究了普氏蹄蝠（Hipposideros pratti)不同状态（飞行,悬挂）下的回声定位声波特征,形态特征和生态特征（捕食策略,捕食地和食物类型）。结果表明,普氏蹄蝠的回声定位声波为CF－FM型,在不同状态下,主频率有一定的差异,飞行状态的主频率略低于悬挂状态,表明普氏蹄蝠是利用多谱勒补偿效应来适应飞行速度引起的主频率变化,以进行准确的定位和有效的捕食;同时飞行状态下声脉冲时间,声脉冲间隔时间及FM带宽略低于悬挂状态,而声脉冲重复率和能率环略主于悬挂状态,表明普氏蹄蝠在不同状态下利用不同特征的声波进行捕食,由回声定位声波推断和野外观察可知,普氏蹄蝠可能在树冠周围以盘旋方式（在昆虫高峰期）或以捕蝇器式（在昆虫高峰期这后）捕食中等偏大的振翅昆虫（如甲虫）。     

The allometry of echolocation call frequencies of insectivorous bats: why do some species deviate from the pattern?

The peak echolocation frequency of insectivorous bats generally declines as body size increases. However, there are notable exceptions to this rule, with some species, such as Rhinolophus clivosus, having a higher than expected peak frequency for their body size. Such deviations from allometry may be associated with partitioning of foraging habitat (the foraging habitat hypothesis) or insect prey (the prey detection hypothesis). Alternatively, the deviations may be associated with the partitioning of sonar frequency bands to allow effective communication in a social context (the acoustic communication hypothesis). We tested the predictions of these hypotheses through comparisons at the family, clade and species level, using species of rhinolophids in general and R. clivosus, a species with a wide distribution, as a specific test case. We compared the wing parameters, echolocation frequency and ecology of R. clivosus to those of the sympatric R. capensis. Rhinolophus clivosus has a much higher echolocation frequency than predicted from its wing loading or body mass. Furthermore, contrary to the predictions of the foraging habitat hypothesis, we found no difference in foraging habitat between R. clivosus and R. capensis. The size range of insect prey taken by the two species also overlapped almost completely, contrary to the prey detection hypothesis. On the other hand, the variation of echolocation frequencies around the allometric relationship for rhinolophids was smaller than that for Myotis spp., supporting the prediction of the acoustic communication hypothesis. We thus propose that the relatively high peak frequency of R. clivosus is the result of partitioning of sonar frequency bands to minimize the ambiguity of echolocation calls during social interactions.     

Response properties of FM-FM combination-sensitive neurons in the auditory cortex of the mustached bat

Summary For echolocation, the mustached bat,Pteronotus parnellii rubiginosus, emits orientation sounds (pulses) and listens to echoes. Each pulse is made up of 8 components, of which 4 are constant frequencies (CF_1–4) and 4 are frequency-modulated (FM_1–4). Target-range information, conveyed by the time delay of the echo FM from the pulse FM, is processed in this species by specialized neurons in a part of the auditory cortex known as the FM-FM area. These cortical neurons are responsive to pulse-echo pairs at specific echo delays (Fig. 1). The essential components in the sound pair include the pulse FM₁ followed by an echo FM_n (n=2, 3 or 4). Downward sweeping FM₁-FM_n sounds that are similar to those the animal naturally hears during echolocation are the most effective in evoking facilitative responses. Most FM-FM neurons, however, still exhibit facilitative responses to stimulus pairs consisting of upward sweeping FM sounds and/or pure tones at frequencies found in FM sweeps (Figs. 2 and 3). The magnitude of facilitation is altered by changes in echo rather than pulse amplitude (Figs. 5 and 6). Neurons characterized by shorter best delays (or echoes from closer targets) do not require larger best echo amplitudes for facilitation.Abbreviations CF constant frequency - FM frequency modulation - H _n CF — FM harmonics of the mustached bat biosonar signal - CF _n CF components of the harmonics - FM _n FM components of the harmonics - PCF _n pulse CF_n - ECF _n echo CF_n - PFM _n pulse FM_n - EFM _n echo FM_n - PH _n pulse H_n - EH _n echo H_n - BA best amplitude for facilitation - BD best delay for facilitation - PST peri-stimulus-time - PSTC peri-stimulus-time-cumulative - dB SPL dB re 20 Pa     

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.     

Identification of sympatric bat species by the echolocation calls

One hundred and thirty-eight echolocation calls of 63 free-flying individuals of five bat species (Rhinolophus ferrumequinum, Myotis formosus, Myotis ikonnikovi, Myotis daubentoni and Murina leucogaster) were recorded (by ultrasonic bat detector (D980)) in Zhi’an village of Jilin Province, China. According to the frequency-time spectra, these calls were categorized into two types: FM/CF (constant frequency) / FM (R. ferrumequinum) and FM (frequency modulated) (M. formosus, M. ikonnikovi, M. daubentoni and M. leucogaster). Sonograms of the calls of R. ferrumequinum could easily be distinguished from those of the other four species. For the calls of the remaining four species, six echolocation call parameters, including starting frequency, ending frequency, peak frequency duration, longest inter-pulse interval and shortest inter-pulse interval, were examined by stepwise discriminant analysis. The results show that 84.1% of calls were correctly classified, which indicates that these parameters of echolocation calls play an important role in identifying bat species. These parameters can be used to test the accuracy of general predictions based on bats’ morphology in the same forest and can provide essential information for assessing patterns of bat habitat use. __________ Translated from Journal of Northeast Normal University (Natural Science Edition), 2006, 38 (3): 109–114 [译自: 东北师范大学学报(自然科学版)]     

Acoustic irradiation produced by flying moths (Lepidoptera,Noctuidae)

Characteristics of acoustic waves accompanying the flight of noctuid moths (Noctuidae) were measured. The low-frequency part of the spectrum is formed of a series of up to 17 harmonics of the wingbeat frequency (30–50 Hz) with a general tendency toward the decrease in the spectral density and the increase in the sound frequency. The root-mean-square level of the sound pressure from flapping wings was found to be 70–78 dB SPL. Besides low-frequency components, the flight of moths was accompanied by short ultrasonic pulses, which appeared with every wingbeat. Most of the spectral energy was concentrated within a range of 7–150 kHz with the main peaks at 60–110 kHz. The short-term pulses were divided into two or more subpulses with different spectra. The high-frequency pulses were produced at two phases of the wingbeat cycle: during the pronation of the wings at the highest point and at the beginning of their upward movement from the lowest point. In most of the specimens tested, the peak amplitude of sounds varied from 55 to 65 dB SPL at a distance of 6 cm from the insect body. However, in nine noctuid species, no high-frequency acoustic components were recorded. In these experiments, the acoustic flow from the flying moth within a frequency range of 2 to 20 kHz did not exceed the self-noise level of the microphone amplifier (RMS 18 dB SPL). Probable mechanisms of the high frequency acoustic emission during flight, the effect of these sounds on the auditory sensitivity of moths, and the possibility of their self-revealing to insectivorous bats are discussed. In addition, spectral characteristics of the moth echolocation clicks were more precisely determined within the higher frequency range (>100 kHz).     

雌蚊翅振音及其在蚊虫防治中的应用

蚊虫飞翔时 ,翅上下拍打会形成连续的翅振音。雌蚊翅振音频率会随蚊种、蚊体长和日龄及环境温度而变化 ,一般在 3 0 0～ 5 0 0Hz之间。同种个体间翅振音频率变化较小 ,常在平均频率± 5 0Hz范围内。雄蚊只对基本频率的雌蚊翅振音起反应 ,雌蚊翅振音的偶然变化不会降低其对雄蚊的引诱力。雄蚊对雌蚊翅振音具有敏感反应的主要原因是其听觉器官对雌蚊翅振音形成了良好的适应性。因此 ,许多蚊虫研究者希望利用雌蚊翅振音来防治蚊虫。因该方法对环境安全 ,在今后蚊虫防治中其重要性将会日益显现。     

Storage of Doppler-shift information in the echolocation system of the “CF-FM”-bat,Rhinolophus ferrumequinum

Summary The greater horseshoe bat (Rhinolophus ferrumequinum) emits echolocation sounds consisting of a long constant-frequency (CF) component preceeded and followed by a short frequency-modulated (FM) component. When an echo returns with an upward Doppler-shift, the bat compensates for the frequency-shift by lowering the emitted frequency in the subsequent orientation sounds and stabilizes the echo image. The bat can accurately store frequency-shift information during silent periods of at least several minutes. The stored frequency-shift information is not affected by tone bursts delivered during silent periods without an overlap with an emitted orientation sound. The system for storage of Doppler-shift information has properties similar to a sample and hold circuit with sampling at vocalization time and with a rather flat slewing rate for the stored frequency information.Supported by Stiftung Volkswagenwerk, grant No. 111858, Deutsche Forschungsgemeinschaft, grant No. Ne 146/7, National Science Foundation (USA), grant No. GB-40018 and the Alexander von Humboldt-Stiftung.     

Echolocation calls of Rhinolophus ferrumequinum in relation to habitat type and environmental factors

The present experiment was carried out in Luotong Mountain Natural Reserve in Jilin Province of China in 2007. We recorded and analyzed the echolocation calls of Rhinolophus ferrumequinum in different habitats by using Avisoft Bioacoustics USG 116 and Avisoft-SASLAB PRO (Avisoft Bioacoustics, Berlin, Germany). Our results showed that R. ferrumequinum foraged in diverse habitats in the study area, and their echolocation calls were significantly variable in different habitats (One-Way ANOVA, P < 0.05). Vegetative, climatic and topographical factors were selected by using the principal component analysis and the correlations between the parameters of echolocation calls and these environmental factors were analyzed. The results indicated that although R. ferrumequinum always emitted FM/CF/FM echolocation calls in different habitats, the parameters of echolocation calls varied with variable environmental factors. Significant negative correlation existed between FM1 bandwidth and arbor height (r = ?0.948, P < 0.05), FM₂ bandwidth and arbor height (r = ?0.825; P < 0.05), FM1 starting frequency and canopy area (r = ?0.967, P < 0.05), FM₂ ending frequency and canopy area (r = ?0.958, P < 0.05), FM1 starting frequency and air relative humidity (r = ?0.776, P < 0.05), FM₂ ending frequency and air relative humidity (r = ?0.875, P < 0.05), peak frequency and air relative humidity (r = ?0.794, P < 0.05), pulse duration and average shrub height (r = ?0.911, P < 0.05), and inter-pulse interval and average shrub height (r = ?0.990, P < 0.05). Significant positive correlation existed between peak frequency and number of plants (r = 0.756, P < 0.05), and pulse duration and height below the canopy (r = 0.870, P < 0.05). Our results suggested that many kinds of ecological factors (such as vegetation factor, climatic factor and topographical factor) affected the structure of echolocation calls and made them diverse in different habitats, i.e., echolocation calls of bats had phenotypic flexibility and eco-adaptability. These characteristics determined the degree of available habitats and natural resources for R. ferrumequinum.