皮氏菊头蝠夏季的捕食行为对策

利用蝙蝠超声波探测器和CoolEditor 2 0 0 0声波分析软件研究了皮氏菊头蝠 (Rhinolophuspearsoni)的超声波信号 ,同时在野外研究了其捕食行为。研究结果显示皮氏菊头蝠是FM/CF/FM型的食虫蝙蝠。其回声定位信号的CF声波两端均附有短暂的FM信号 ,每次声波脉冲包含 2段信号 ,第 1段信号的CF频率为 (6 1 0 8±0 0 19)kHz ,持续时间为 (4 6 85± 3 72 )ms ;第 2段信号的CF频率为 (6 0 97± 0 0 3)kHz ,持续时间为 (35 12± 2 6 7)ms。在对皮氏菊头蝠的捕食行为研究中 ,通过运用生物多样性指数分析和Spearman相关性分析 ,结果表明皮氏菊头蝠在常绿阔叶落叶混交林中主要以式捕食鳞翅目 (Lepidoptera)、鞘翅目 (Coleoptera)等中型个体的昆虫 ,对食物种类及其体型具有选择性。此外 ,其形态与回声定位功能之间还表现出相关性。     

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。     

三种蝙蝠飞行状态下回声定位信号的比较

对马铁菊头烟、萨氏伏翼与白腹管鼻蝠在飞行状态下的回声定位信号进行了比较研究。结果表明：马铁菊头蝠的回声定位为长恒频一调频声（CF－FM）叫声（叫声持续时间均长于10ms）,具有两个谐波;萨氏伏翼与白腹管鼻蝠的回声定位均为宽频调频声（FM）叫声,叫声持续时间分别为3．0±1．3ms和3．6±3．0ms,具有两个或三个谐波。这些回声定位叫声的特性可能与它们所处的生境相适应。     

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。     

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。     

拖网式食鱼蝠——大足鼠耳蝠的形态、回声定位声波及捕食策略

大足鼠耳蝠(Myotisricketti)是中国特有蝙蝠,其回声定位声波和捕食策略国内外均无报道,对大足鼠耳蝠该方面的研究报导是国内首次。大足鼠耳蝠体型较大,具有强大的后足,足上有强而有力的弯曲的爪,尾膜和距很长。大足鼠耳蝠回声定位声波为FM(调频)型,一般具有1～2个谐波,主频率较低(37.78±1.04kHz),调频带较宽(第一谐波频带宽为42.02±6.98kHz,第二谐波频带宽为25.79±7.89kHz),声脉冲时间较长(2.91±0.54ms),声脉冲间隔时间变化较大(32.30±15.10ms),能率环较高(11.27±5.84%);野外观察发现,大足鼠耳蝠主要在低水面上空飞行,利用大足从水面捕食猎物(拖网式捕食),猎物主要由鱼类组成。即分析和讨论了大足鼠耳蝠形态特征、回声定位特征和捕食策略的相互适应性。     

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）。     

八种菊头蝠回声定位声波频率与体型的相关性

菊头蝠回声定位声波中含有强的恒频(con-stant frequency,CF)组分,通常在开始和结尾伴有短的FM组分(Schnitzler,1968).飞行状态能影响回音信号(张树义等,1999).在飞行中,蝙蝠发出的频率变低以补偿由飞行速度引起的多谱勒变化,返回的回声接近于蝙蝠停止时的声波频率(Schnitzler,1968).回声定位声波的频率随蝙蝠年龄和季节的变化会产生一些变动,但如果频率被身体结构制约,CF组分频率在蝙蝠静止时会保持相对恒定(Vater,1987;Heller et al.,1989;Joneset al.1994).Francis et al.(1998)对19种菊头蝠、Heller et al.(1989)对26种菊头蝠进行了体型测量和回声定位声波信号的测定,得出结论为:菊头蝠回声定位声波中CF组分的频率与体型大小成负相关.但Jones(1992)和Jones et al.(1993)认为体型大小对菊头蝠回声定位声波没有影响.     

吉林省发现绯鼠耳蝠

在吉林省集安市采到绯鼠耳蝠 (Myotisformosus)样本 5只 ,为吉林省蝙蝠科新记录。对其体型与头骨进行了测量。对回声定位声波进行了录制和分析 ,发现其回声定位声波为FM型 ,一次完整声波包括1～ 2个谐波 ,主频率 (5 4 5 4± 6 3 9)kHz,通过回声定位声波特征推测 ,绯鼠耳蝠在较简单的环境中捕食中等大小的昆虫。     

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Batsound 3.0分析,其声波为高频 (145.4±10.9 kHz)、宽带(62.6±9.2 kHz)、具两个谐波的短(1.67±0.4 ms)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。     

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…     

Ultrasonic bouts of a blind climbing rodent (Typhlomys chapensis): acoustic analysis

ABSTRACT

The peculiar acoustic structure of ultrasonic bouts of blind climbing rodents Typhlomys might provide insight on their potential function. We examined 1481 bouts consisting of 1-6 pulses; 49.7% of them were single-pulse bouts. Bout duration and inter-bout interval depended on the number of pulses per bout, whereas period from start of a previous bout to start of the next bout was constant (80.0±2.9 ms). Ultrasonic pulses (540 pulses measured in a subset of 234 bouts) were short (0.68±0.15 ms) sweeps with fundamental frequency slopes from 127.3±6.3 kHz to 64.1±4.6 kHz and peak frequency at 93.3±7.4 kHz, emitted within bouts with inter-pulse periods of 13.03±3.01 ms. Single pulses and start pulses of multi-pulse bouts were lower in frequency than other pulses of the bouts. In contrast, pulse duration was independent on pulse position within bout. Pulses of Typhlomys were reminiscent of echolocation calls of Murina and Myotis bats, but higher in frequency, much shorter, fainter, displayed a convex contour of frequency modulation and only the fundamental frequency band without harmonics and were organized in bouts, that is not characteristic for bat echolocation. Most probably, Typhlomys uses their ultrasonic pulses for call-based orientation during locomotion, including climbing and jumping among bush branches.     

Echolocation calls of free-flying common vampire bats Desmodus rotundus (Chiroptera: Phyllostomidae) in Chile

We recorded and characterized the echolocation calls emitted by the common vampire bat Desmodus rotundus during foraging in natural habitats in Chile. Signal design typically shows multiple harmonics consisting of a brief quasi-constant frequency (QCF) component at the beginning of the pulse followed by a downward frequency modulated component. Calls are characterized by long durations (5.5 ms) and emitted as single pulses or in groups of 2–3 pulses at a repetition rate of 29 Hz. The higher frequency ranges (85–35 kHz) and the unusual QCF component that characterized multiharmonic signals of free-flying D. rotundus in Chile is a remarkable feature for acoustic identification with other Chilean bats.     

Ontogenesis of the echolocation system in the rufous horseshoe bat,Rhinolophus rouxi (Audition and vocalization in early postnatal development)

1. The development of vocalization and hearing was studied in Sri Lankan horseshoe bats (Rhinolophus rouxi) during the first postnatal month. The young bats were caught in a nursing colony of rhinolophids in which birth took place within a two week period. 2. The new-born bats emitted isolation calls through the mouth. At the beginning these calls consisted of pure tones with frequencies below 10 kHz (Fig. 1). During the first postnatal week the call frequency increased to about 15 kHz, and the fundamental was augmented by two to four harmonics. No evoked potentials to pure tone stimuli could be elicited in the inferior colliculus of this age group, i.e., auditory processing at the midbrain level was not demonstrable. 3. Evoked potentials were first recorded in the second week, broadly tuned to 15-45 kHz, with a maximum sensitivity between 15-25 kHz. In the course of the second week, however, higher frequencies up to 60 kHz became progressively incorporated into the audiogram (Fig. 3). The fundamental frequency of the multiharmonic isolation calls, emitted strictly through the mouth, increased to about 20 kHz. 4. In the bats' third postnatal week an increased hearing sensitivity (auditory filter) emerged, sharply tuned at frequencies between 57 and 60 kHz (Fig. 4e). The same individuals were also the first to emit long constant frequency echolocation calls through the nostrils (Fig. 4c). The energy of the calls was arranged in harmonic frequency bands with the second harmonic exactly tuned to the auditory filter. These young bats continued to emit isolation calls through the mouth, which were, however, not harmonically related to the echolocation calls (Fig. 4b, d). 5. During the fourth week, both the auditory filter and the matched echolocation pulses (the second harmonic) shifted towards higher frequencies (Fig. 5). During the fifth week the fundamental frequency of the calls was progressively attenuated, and both the second harmonic of the pulses and the auditory filter reached the frequency range typical for adult bats of 73-78 kHz (Fig. 6). 6. The development of audition and vocalization is discussed with regard to possible interactions of both subsystems, and their incorporation into the active orientation system of echolocation.     

Vocal communication in adult greater horseshoe bats, Rhinolophus ferrumequinum

Whereas echolocation in horseshoe bats is well studied, virtually nothing is known about characteristics and function of their communication calls. Therefore, the communication calls produced by a group of captive adult greater horseshoe bats were recorded during various social interactions in a free-flight facility. Analysis revealed that this species exhibited an amazingly rich repertoire of vocalizations varying in numerous spectro-temporal aspects. Calls were classified into 17 syllable types (ten simple syllables and seven composites). Syllables were combined into six types of simple phrases and four combination phrases. The majority of syllables had durations of more than 100 ms with multiple harmonics and fundamental frequencies usually above 20 kHz, although some of them were also audible to humans. Preliminary behavioral observations indicated that many calls were emitted during direct interaction with and in response to social calls from conspecifics without requiring physical contact. Some echolocation-like vocalizations also appeared to clearly serve a communication role. These results not only shed light upon a so far widely neglected aspect of horseshoe bat vocalizations, but also provide the basis for future studies on the neural control of the production of communicative vocalizations in contrast to the production of echolocation pulse sequences.     

菲菊头蝠回声定位声波频率的性二态有利于性别识别

许多动物的叫声频率呈现性二态现象。蝙蝠夜间活动,主要利用声音信号导航空间、追踪猎物、传递交流信息。本研究选择成体菲菊头蝠作为研究对象,检验回声定位声波频率性二态是否有利于性别识别。研究发现,菲菊头蝠回声定位声波频率参数具有显著性别差异。播放白噪音、雄性回声定位声波及雌性回声定位声波期间,实验个体的反应叫声数量依次递减。播放白噪音、雌性回声定位声波及雄性回声定位声波后,实验个体的反应叫声数量依次递增。白噪音诱导反应叫声强度高于回声定位声波诱导反应叫声强度。研究结果表明,菲菊头蝠回声定位声波的频率参数编码发声者性别信息,有利于种群内部的性别识别。本研究暗示,回声定位声波可能在蝙蝠配偶选择中扮演一定作用。     

三种共栖蝙蝠的回声定位信号特征及其夏季食性的比较

2005年6至9月,对桂林市郊区两个山洞中高颅鼠耳蝠(Myotissiligorensis)、菲菊头蝠(Rhinolo-phuspusillus)和黑髯墓蝠(Taphozousmelanopogon)的回声定位叫声特征和食性进行分析,并结合其形态特征与野外观察,推断其捕食生境和捕食策略。研究结果发现:黑髯墓蝠体型最大,声音特征属短调频型多谐波,一般为4个谐波,能量主要集中在第二谐波上,主频率为(32·84±1·17)kHz,选择鞘翅目和双翅目昆虫为主要食物;高颅鼠耳蝠(长调频型)和菲菊头蝠(长恒频-调频型),体型都较小,主频率分别是(84·44±8·13)kHz和(110·78±1·65)kHz,以双翅目昆虫为主要食物;而菲菊头蝠则以鞘翅目和双翅目昆虫为主要食物。上述结果证明,高颅鼠耳蝠、菲菊头蝠和黑髯墓蝠在声音和食物组成等方面出现了明显分化。     

Different Auditory Feedback Control for Echolocation and Communication in Horseshoe Bats

Auditory feedback from the animal''s own voice is essential during bat echolocation: to optimize signal detection, bats continuously adjust various call parameters in response to changing echo signals. Auditory feedback seems also necessary for controlling many bat communication calls, although it remains unclear how auditory feedback control differs in echolocation and communication. We tackled this question by analyzing echolocation and communication in greater horseshoe bats, whose echolocation pulses are dominated by a constant frequency component that matches the frequency range they hear best. To maintain echoes within this “auditory fovea”, horseshoe bats constantly adjust their echolocation call frequency depending on the frequency of the returning echo signal. This Doppler-shift compensation (DSC) behavior represents one of the most precise forms of sensory-motor feedback known. We examined the variability of echolocation pulses emitted at rest (resting frequencies, RFs) and one type of communication signal which resembles an echolocation pulse but is much shorter (short constant frequency communication calls, SCFs) and produced only during social interactions. We found that while RFs varied from day to day, corroborating earlier studies in other constant frequency bats, SCF-frequencies remained unchanged. In addition, RFs overlapped for some bats whereas SCF-frequencies were always distinctly different. This indicates that auditory feedback during echolocation changed with varying RFs but remained constant or may have been absent during emission of SCF calls for communication. This fundamentally different feedback mechanism for echolocation and communication may have enabled these bats to use SCF calls for individual recognition whereas they adjusted RF calls to accommodate the daily shifts of their auditory fovea.     

Genetic divergence and echolocation call frequency in cryptic species of Hipposideros larvatus s.l. (Chiroptera: Hipposideridae) from the Indo-Malayan region

The intermediate leaf-nosed bat ( Hipposideros larvatus ) is a medium-sized bat distributed throughout the Indo-Malay region. In north-east India, bats identified as H. larvatus captured at a single cave emitted echolocation calls with a bimodal distribution of peak frequencies, around either 85 kHz or 98 kHz. Individuals echolocating at 85 kHz had larger ears and longer forearms than those echolocating at 98 kHz, although no differences were detected in either wing morphology or diet, suggesting limited resource partitioning. A comparison of mitochondrial control region haplotypes of the two phonic types with individuals sampled from across the Indo-Malay range supports the hypothesis that, in India, two cryptic species are present. The Indian 98-kHz phonic bats formed a monophyletic clade with bats from all other regional populations sampled, to the exclusion of the Indian 85-kHz bats. In India, the two forms showed 12–13% sequence divergence and we propose that the name Hipposideros khasiana for bats of the 85-kHz phonic type. Bats of the 98-kHz phonic type formed a monophyletic group with bats from Myanmar, and corresponded to Hipposideros grandis , which is suggested to be a species distinct from Hipposideros larvatus . Differences in echolocation call frequency among populations did not reflect phylogenetic relationships, indicating that call frequency is a poor indicator of evolutionary history. Instead, divergence in call frequency probably occurs in allopatry, possibly augmented by character displacement on secondary contact to facilitate intraspecific communication.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 119–130.