短嘴金丝燕的回声定位机制与其归巢行为的探究

通过对短嘴金丝燕(Aerodramus brevirostris)的传统遮挡放飞实验和飞行行为的观察,研究短嘴金丝燕回声定位的机制,同时利用SONY MD录音并用Cool Edit2.1软件进行声波分析,证明了该物种是能利用回声定位的鸟类。该物种定位的声波主频较低,所以其定位能力没有蝙蝠(Chiroptera)那样精确,个体主要还是依靠视觉定位,只有在全黑的情况下才利用声音定位的。     

回声定位蝙蝠及其声通讯

综述了回声定位蝙蝠种类及其发声方式,回声定位信号的主要类型及回声定位信号声学特征,多普勒频移对长CF／FM蝙蝠的主要作用,简介了蝙蝠求偶和母婴识别等内声通讯行为,提出了一些尚待解决的重要问题。     

Echolocation versus echo suppression in humans

Several studies have shown that blind humans can gather spatial information through echolocation. However, when localizing sound sources, the precedence effect suppresses spatial information of echoes, and thereby conflicts with effective echolocation. This study investigates the interaction of echolocation and echo suppression in terms of discrimination suppression in virtual acoustic space. In the ‘Listening’ experiment, sighted subjects discriminated between positions of a single sound source, the leading or the lagging of two sources, respectively. In the ‘Echolocation’ experiment, the sources were replaced by reflectors. Here, the same subjects evaluated echoes generated in real time from self-produced vocalizations and thereby discriminated between positions of a single reflector, the leading or the lagging of two reflectors, respectively. Two key results were observed. First, sighted subjects can learn to discriminate positions of reflective surfaces echo-acoustically with accuracy comparable to sound source discrimination. Second, in the Listening experiment, the presence of the leading source affected discrimination of lagging sources much more than vice versa. In the Echolocation experiment, however, the presence of both the lead and the lag strongly affected discrimination. These data show that the classically described asymmetry in the perception of leading and lagging sounds is strongly diminished in an echolocation task. Additional control experiments showed that the effect is owing to both the direct sound of the vocalization that precedes the echoes and owing to the fact that the subjects actively vocalize in the echolocation task.     

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.     

Echolocation constraints of Daubenton's Bat foraging over water

1. Daubenton's Bats ( Myotis daubentonii ) foraging over a stream concentrated their activity over calm surfaces, avoiding an adjacent area with small ripples (< 3 cm high). Aerial insects were most abundant over the ripples, so insect distribution could not explain why the bats avoided this area.
2. The bats flew low over water and always ( N = 22) directed the head forwards, presumably emitting the echolocation beam parallel to the surface, thus minimizing clutter. At an angle of incidence of 30° there was significantly more clutter from the rippled water.
3. The ripples produced ultrasonic noises in the form of transient pulses at an average rate of 6·2 per second. In the present case, such pulses were common enough potentially to interfere with target detection by the bats. Transient noises and echo clutter from moving ripples may be the principal reason why bats generally avoid foraging low over turbulent water.
4. The target strength of a potential insect prey at the water surface and the source levels of the bats' searching signals were measured to use in estimating the echo level at the bat when it detects the prey. The echo level at detection (+ 38 dB sound pressure level) was about the same as the clutter level extrapolated to the detection distance. This suggests that Daubenton's Bat operates at very low signal-to-noise ratios when foraging for insects near the water surface.     

Echolocation by the barbastelle bat, Barbastella barbastellus

When searching for insects along edges, Barbastella barbastellus alternated between two signal types. Type-2 signals had durations around 6 ms and were composed of an initial shallowly downward frequency modulated component, starting at about 45 kHz and followed by a shorter more steeply modulated component that ended at about 32 kHz. Type-1 signals were rather stereotyped with durations around 2.5 ms and a very short rise time. They covered an approximately 8 kHz-wide frequency band positioned just below the 12-15 kHz-wide frequency band of type-2 signals, with no or small frequency overlap. In the recordings, type-1 signals almost had always a higher amplitude than type-2 signals, at least partly caused by head movements. Assuming that signal structure reflects function, we hypothesize that type-2 signals have the same adaptive value as the signals with a broadband and narrowband component of other vespertilionids, but with a reverse arrangement of the signal elements. Like the broadband component of the type-2 signals, type-1 signals are well suited to localize background targets. Thus, the localization component may be distributed among two signals separated in time, which has the advantage that both signals can be varied independently in the direction of emission and in amplitude.     

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.     

大趾鼠耳蝠回声定位声波特征与分析

在12 m×4 m×4 m的围网内录制大趾鼠耳蝠(Myotis macrodactylus)飞行与悬挂状态下的回声定位声波,使用双尾t-检验对不同状态下的声波参数进行差异显著性分析.结果表明,大趾鼠耳蝠回声定位声波为短的、宽带的且能量主要集中在第1谐波的调频型声波,伴有1-2个谐波.第1谐波起始频率、带宽和声脉冲间隔在飞行与悬挂状态下具有显著差异(P<0.05).回声定位声波飞行状态下的第1谐波终止频率、带宽、声脉冲持续时间和声脉冲间隔均存在性别差异,而主频率没有显著的性别差异.回声定位声波的这些特征及差异体现了对其捕食生境、捕食策略及通讯行为的适应.     

华南菊头蝠的回声定位声波特征与分析

使用Petterson D500X超声波接收仪对华南菊头蝠Rhinolophus huananus飞行和静止状态的回声定位声波进行录制,利用独立样本t检验对2种状态的回声定位声波参数进行分析。结果显示,华南菊头蝠的回声定位声波类型为FM-CF-FM型,有1~2个谐波,不同状态下的峰频、声脉冲时间、声脉冲间隔时间差异均有统计学意义(P0.05)。与相似种大耳菊头蝠R.macrotis比较,二者在飞行状态下的回声定位声波峰频存在差异,可作为区分2个种之间的声学依据。     

高颅鼠耳蝠回声定位声波特征与分析

在自建的野外实验事内,录制雌雄成体高颅鼠耳蝠Myotis siligorensis不同行为状态(飞行、爬行、手持)回声定位声波,利用单因素方差分析(One-Way ANOVA)对不同状态及不同性别的回声定位声波参数进行均值多重比较和显著性差异分析.结果 表明,高颅鼠耳蝠回声定位声波为长的宽频带调频(FM)声波,有1~3个谐波,但能量主要集中在第一谐波.不同行为状态下,高颅鼠耳蝠声脉冲持续时间、声脉冲间隔和能率环均存在显著性差异(P<0.05),第一谐波起始频率和终止频率、主频率、带宽差异不显著;第一谐波终止频率、带宽、声脉冲持续时间和能率环的性别差异显著(P<0.05),具有性别二态性.高颅鼠耳蝠回声定位声波特征体现了其在捕食策略和捕食生境方面的生态适应.     

无尾蹄蝠的回声定位声波特征及分析

采用超声波监听仪U30录制无尾蹄蝠自由飞行状态的回声定位声波,经Batsound3.0分析,其声波为高频(145.4±10.9kHz)、宽带(62.6±9.2kHz)、具两个谐波的短(1.67±0.4ms)FM型,不同于蹄蝠科其他蝙蝠的CF型,表明该科内物种声波类型存在多态性。头骨的形态测定分析支持其通过鼻腔发射声波,与蹄蝠科其他蝙蝠一致,表明该科内声波发射方式的单一性。适应环境的选择压力及翼型和声波的适应性可能是其选择FM型叫声的重要原因。     

马铁菊头蝠不同状态下回声定位声波分析

在自建网室(9 m×4 m×4 m)内驯养马铁菊头蝠(Rhinolophus ferrumequinum),利用超声波探测仪录制蝙蝠不同状态下回声定位声波,声波录制与红外摄像保持同步。结果表明,马铁菊头蝠回声定位声波为调频(FM)/恒频(CF)/调频(FM)型;在蝙蝠接近猎物过程中,声脉冲持续时间和间隔时间显著变短,下调FM(即tFM)组分变得愈为显著,捕捉猎物瞬间,产生捕食蜂鸣;飞行与悬挂状态相比,声脉冲重复率、主频率、声脉冲时间、声脉冲间隔和能率环的差异均达到显著水平。     

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.     

彩蝠的形态和回声定位信号特征

2008年11月于海南省临高县美登镇网捕到1只雄性蝙蝠标本,经形态鉴定为蝙蝠科(Vespertilionidae)彩蝠属(Kerivoula)彩蝠(K.picta).本文记述了采自海南岛的该彩蝠的形态与回声定位信号特征.     

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.     

Echolocation Performance of the Vampire Bat (Desmodus rotundus)

The neotropical vampire bats (Desmodus rotundus) echolocate using ultrasonic pulses like those of the Latin American phyllostomatid bats. In this paper the orally produced echolocation sounds of Desmodus are analysed and the performance of the echolocation system is studied in two-choice training experiments on two vampire bats. Ability to detect objects is relatively limited; both animals were capable of discerning the presence of a 1 cm wide metal strip at a distance of 50 cm, but they failed with 0.5 cm wide strips. The ultrasonic pulses produced at a distance of 50 cm appear to sample an area with a diameter of 2.5 to 3.0 cm (i. e., the solid angle tested with each pulse is 3° to 3° 40′ in extent).     

大棕蝠江南亚种回声定位声波特征与分析

在自建录音棚内录制大棕蝠汀南亚种Eptesicus serotinus andersoni不同状态下的同声定位声波,使用单因素方差分析(One-Way ANOVA)方法埘小同状态下的声波参数进行显著性差异分析和均值的多重比较,结果表明,大棕蝠江南亚种回声定位声波为短的、宽带的且能量大部分集中在第1谐波的调频型声波,伴有3～4个谐波.并且飞行、悬挂和手持状态的各声波参数均存在显著差异(P<0.05).悬挂状态同声定位声波的声脉冲持续时间大于飞行和手持状态,飞行状态下回声定位声波声脉冲间隔最小,但同声定位声波的主频率为所有状态中最高的.回声定位声波的这些特征及差异体现了与其捕食策略和捕食生境的适应性.