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）。     

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CFFM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高于悬挂状态,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(在昆虫高峰期之后)捕食中等偏大的振翅昆虫(如甲虫).     

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。     

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.     

普通长翼蝠福建亚种不同行为状态下回声定位声波研究

普通长翼蝠福建亚种为中国的地方性亚种。采用超声波监听仪和Batsound3 10软件对其回声定位声波进行录制和分析,发现回声定位声波为中等长度的FM型,伴有1～2个谐波,声波主频率为49 35±4 24kHz,一次完整声波的声脉冲时间为3 46±1 63ms,两次声波间的声脉冲间隔为96 09±33 84ms。分析表明,普通长翼蝠福建亚种在飞行和手持状态下的回声定位声波声脉冲时间均小于其悬挂状态,飞行状态下声脉冲间隔时间是各种状态中较小的,而飞行状态下回声定位声波的主频率则为所有状态中最高的,说明蝙蝠在飞行中要面临复杂的环境,辨别较多的障碍物,因此利用高频率声波进行回声定位,才能实现灵活复杂的飞行。     

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.     

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。     

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.     

两种扁颅蝠回声定位叫声的比较

对扁颅蝠 (Tylonycterispachypusa)和褐扁颅蝠 (T robustula)在飞行状态下的回声定位叫声进行了比较研究。结果表明 ,2种扁颅蝠的回声定位叫声的声谱图均呈调频 (FM)型 ,且波形相似 ;但叫声的最低频率、最高频率和主频率差异极显著 (P <0 0 1)。扁颅蝠的频率范围较高 ,为 6 2 4～ 91 6kHz ,主频率为 (76 5± 2 1)kHz ;褐扁颅蝠的频率范围较低 ,为 4 2 7～ 72 4kHz ,主频率为 (49 2± 1 8)kHz ;而 2种蝙蝠的声脉冲时程、声脉冲间隔和声脉冲重复率差异不显著 (P >0 0 5 )。回声定位叫声差异与其体型、所处的生境有关     

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。     

IDENTIFICATION OF BRITISH BAT SPECIES BY MULTIVARIATE ANALYSIS OF ECHOLOCATION CALL PARAMETERS

ABSTRACT

Previous studies have found variability and individual distinctiveness in the echolocation calls of bats. We consider two implications of individually distinct echolocation calls: 1) whether bats may be able to use such variation to recognise familiar conspecifics, and 2) whether investigators could use such variation to identify known individuals or to census populations. We compared the discriminability of the echolocation calls of big brown bats (Eptesicus fuscus) recorded in three situations: (a) while held in the hand, (b) while perched on a platform, and (c) while flying in an anechoic chamber. Using variables describing each sonar call, we employed discriminant function analysis (DFA) to assign calls to recording situation or to bat. Discrimination of calls by recording situation was largely unsuccessful, although flying calls could be distinguished from platform calls. Assignment of calls to individual bat across recording situations yielded 72% success, and, within a given recording situation, yielded 87% success. Stepwise DFA reduced the number of variables needed to discriminate between individuals with only a slight decrease in correct classification. These results suggest that bats (or researchers) may be able to use the information contained in the echolocation calls for individual recognition. Individual distinctiveness raises the possibility of censusing bats by sound. We used cluster analysis in an attempt to determine whether, given a sample of calls from an unknown number of bats, a reasonable estimate of the number of bats could be obtained. Results were unsatisfactory, suggesting that cluster analysis probably will not permit acoustic censusing of bats in the field.     

Substrate-gleaning versus aerial-hawking: plasticity in the foraging and echolocation behaviour of the long-eared bat,Myotis evotis

The foraging and echolocation behaviour of Myotis evotis was investigated during substrate-gleaning and aerial-hawking attacks. Bats gleaned moths from both the ground and a bark-covered trellis, however, they were equally adept at capturing flying moths. The calls emitted by M. evotis during substrate-gleaning sequences were short, broadband, and frequency-modulated (FM). Three behavioural phases were identified: search, hover, and attack. Gleaning search calls were significantly longer in duration, lower in highest frequency, and larger in bandwidth than hover/attack calls. Calls were detected in only 68% of gleaning sequences, and when they were emitted, bats ceased calling 200 ms before attacking. Terminal feeding buzzes, the rapid increase in pulse repetition rate associated with an attempted prey capture, were never recorded during gleaning attacks. The echolocation calls uttered by M. evotis during aerial-hawking foraging sequences were also short duration, high frequency, FM calls. Two distinct acoustic phases were identified: approach and terminal. Approach calls were significantly different from terminal calls in all variables measured. Calls were detected in 100% of aerial-hawking attacks and terminal feeding buzzes were invariably produced. Gleaning hover/attack calls were spectrally similar to aerial approach calls, but were shorter in duration and emitted at a significantly lower (but constant) repetition rate than aerial signals. Although the foraging environment (flight cage contents) remained unchanged between tasks (substrate-gleaning vs. aerial-hawking), bats emitted significantly lower amplitude calls while gleaning. We conclude that M. evotis adjusts its echolocation behaviour to meet the perceptual demands (acoustical constraints) imposed by each foraging situations.Abbreviations BW bandwidth - CF constant frequency - dB SPL decibels sound pressure level - FM frequency modulated - HF highest frequency - LF lowest frequency - PF peak frequency Presented at the meeting Acoustic Images in Bat Sonar, a conference on FM echolocation honoring Donald R. Griffin's contributions to experimental biology (June 14–16, Brown University, Providence RI).     

Calling louder and longer: how bats use biosonar under severe acoustic interference from other bats

Active-sensing systems such as echolocation provide animals with distinct advantages in dark environments. For social animals, however, like many bat species, active sensing can present problems as well: when many individuals emit bio-sonar calls simultaneously, detecting and recognizing the faint echoes generated by one''s own calls amid the general cacophony of the group becomes challenging. This problem is often termed ‘jamming’ and bats have been hypothesized to solve it by shifting the spectral content of their calls to decrease the overlap with the jamming signals. We tested bats’ response in situations of extreme interference, mimicking a high density of bats. We played-back bat echolocation calls from multiple speakers, to jam flying Pipistrellus kuhlii bats, simulating a naturally occurring situation of many bats flying in proximity. We examined behavioural and echolocation parameters during search phase and target approach. Under severe interference, bats emitted calls of higher intensity and longer duration, and called more often. Slight spectral shifts were observed but they did not decrease the spectral overlap with jamming signals. We also found that pre-existing inter-individual spectral differences could allow self-call recognition. Results suggest that the bats’ response aimed to increase the signal-to-noise ratio and not to avoid spectral overlap.     

大蹄蝠回声定位信号特征与下丘神经元频率调谐

采用超声监测仪录制超声信号和细胞外电生理记录下丘神经元的频率调谐曲线(frequency tuningcurqes,FTCs)的方法,探讨了大蹄蝠(Hipposideros armiger)回声定位信号与下丘(inferior colliculus,IC)神经元频率调谐之间的相关性.结果发现,大蹄蝠回声定位叫声为恒频-调频(consrant frequency-frequenevmodulated,CF-FM)信号,一般含有2-3个谐波,第二谐波为其主频,cF成分频率(Mean±SD,n=18)依次为:(33.3 4±0.2)、(66.5±0.3)、(99.4 4±0.5)kHz;电生理实验共获得72个神经元的频率调谐曲线,Q10-dB值的范围是0.5-95.4(9.2±14.6,rg=72),最佳频率(best frequency,BF)在回声定位主频附近的神经元具有尖锐的频率调谐特性.结果表明,大蹄蝠回声定位信号与下丘神经元频率调谐存在相关性,表现为最佳频率在回声定位信号主频附近的神经元频率调谐曲线的Q10-dB值较大,具有很强的频率分析能力.     

Can pipistrelles, Pipistrellus pipistrellus (Schreber, 1774) and Pipistrellus pygmaeus (Leach, 1825), foraging in a group, change parameters of their signals?

Echolocation behaviour and the structure of calls of Pipistrellus pygmaeus and Pipistrellus pipistrellus were studied by using a time expansion bat detector. Echolocation signals were recorded in the field in south-eastern Moravia and northern Bohemia (Czech Republic) and in an ad hoc experimental laboratory. For each of the species, multivariate analysis of variance (MANOVA) indicated significant differences in calls produced inside the experimental room and in the open. Paired t -tests and MANOVA were also used to reveal influences of interindividual contacts in each of the cryptic species on the spectral patterns of call variables. Differences were found in the spectral variables of echolocation calls of an individual flying in the room alone and in a group of conspecifics. The possibility that bats use their flexibility to avoid mutual disturbances of echolocation calls was tested. We found that bats flying in a group modify the parameters of their echolocation signals according to the presence of other individuals of the same species. These differences can indicate jamming avoidance and recognition of own echoes. However, they did not change the parameters if individuals of another species were present. Social calls are more numerous when bats fly in a mixed-species group than in a monospecific group.     

普氏蹄蝠下丘谐波主频神经元的时程选择性

为探讨下丘(Inferior colliculus,IC)回声定位信号主频范围内的神经元的时程选择性,在自由声场刺激条件下,我们在4 只普氏蹄蝠的IC 采用不同时程的声刺激,研究了神经元的时程选择性。通过在体细胞外记录,共获得56 个声敏感下丘神经元,其记录深度、最佳频率和最小阈值的范围分别为1547 - 3967 (2878. 9 ±629.1)μm,20 -68 (49.0 ± 11. 1)kHz 和36.5 -95. 5 (59. 8 ±13. 0)dB SPL。根据所记录到的下丘神经元对不同时程的声刺激的反应,即对不同时程的选择性(Duration selectivity),将其分为6 种类型:短通型(Short-pass,SP,n = 11/56)、带通型(Band-pass,BP,n = 1/56)、长通型(Long-pass,LP,n = 5 /56)、反带通型(Band-reject,BR,n = 3 /56)、多峰型(Multi-peak,MP,n =6 /56)和全通型(All-pass,AP,n =30 /56)或非时程选择型(Nonduration-selective,NDS)。通过比较普氏蹄蝠下丘谐波主频内和主频外神经元的时程选择性,我们发现处于回声定位信号主频范围内神经元(n =32)比主频外神经元(n = 24)具有更短的最佳时程和更高的时程选择性。结果提示,在普氏蹄蝠回声定位过程中谐波主频内神经元较谐波主频外神经元发挥了更为重要的作用。     

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

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