环境噪声对蝙蝠回声定位叫声及飞行活动的影响

环境噪声影响动物的活动及其叫声特性,已成为动物面对的一种重要选择压力。为应对噪声的干扰,多数动物类群会远离噪声区域和改变其叫声的频谱时间结构,如延长叫声持续时间、提高叫声频率等,但有些动物的活动和叫声频谱时间结构并不受环境噪声的影响。本研究在自然条件下,研究不同环境噪声强度对蝙蝠活动和回声定位声波的影响。选取噪声强度有差异的12个样点,分别录制各样点大卫鼠耳蝠、西南鼠耳蝠、亚洲长翼蝠及未知蝙蝠的回声定位声波,分析其持续时间、起始频率、峰频、终止频率和带宽,统计蝙蝠通过次数。回归分析结果显示：环境噪声强度与大卫鼠耳蝠、西南鼠耳蝠、亚洲长翼蝠及未知蝙蝠的活动无显著相关性P > 0.05）,与回声定位声波的脉冲持续时间、起始频率、峰频、终止频率及带宽均不显著相关（P > 0.05）。暗示低频低强度（< 20 kHz, < 67.5 dB）的环境噪声可能对高频回声定位蝙蝠的叫声及活动没有显著影响。     

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。     

降雨噪声对菲菊头蝠出飞行为的影响

降雨噪声属于常见的自然噪声,由雨滴撞击物体表面产生。目前,有关降雨噪声对动物的潜在影响被普遍忽视。回声定位蝙蝠主要利用声信号在黑暗环境导航空间、探测猎物及社群交流,是开展降雨噪声影响研究的理想类群。本研究选择菲菊头蝠 (Rhinolophus pusillus)作为研究对象,检验降雨噪声是否影响蝙蝠出飞行为。我们在集群栖息地外,播放强降雨噪声、空白对照和种内回声定位声波,开展野外回放实验。利用单因素方差分析及其事后检验,评价菲菊头蝠对不同回放刺激的反应差异。研究发现,相比空白对照,强降雨噪声导致菲菊头蝠的通勤数量百分比平均降低2.82倍,回声定位脉冲数量平均减少4.86倍,集群出飞时间延长3.75 min。相比空白对照,同种回声定位声波对菲菊头蝠出飞行为的影响并不显著。研究结果证实强降雨噪声抑制菲菊头蝠的出飞行为。本研究表明,降雨引起 的噪声干扰可能是导致蝙蝠躲避降雨的重要因素,为野生蝙蝠物种保育与管理提供启示。     

马铁菊头蝠对同种咀嚼声的反应研究

咀嚼是哺乳动物食物吸收的重要组成部分,对动物的生存和繁殖极其重要。动物咀嚼时会发出低频低强度的咀嚼声,研究表明人类的咀嚼声可以增强自身或他人的食欲和愉悦度。蝙蝠作为哺乳动物中的第二大类群,其咀嚼声的特征和功能仍不清楚。本研究以吉林省集安市治安村的马铁菊头蝠Rhinolophus ferrumequinum为研究对象,通过回放蝙蝠咀嚼声和空白对照实验,同步录制蝙蝠的进食行为和回声定位声波,试图阐明蝙蝠咀嚼声对其进食行为和回声定位声波的影响。结果表明:进食次数在2种回放条件下的差异无统计学意义,但捕食尝试在2种回放条件下的差异有统计学意义。因此,咀嚼声能够显著地提高蝙蝠的进食欲望。此外,在蝙蝠咀嚼声的刺激下,其回声定位声波的频率增加,持续时间延长,脉冲速率减慢。这可能是因为蝙蝠在咀嚼声的影响下改变发声动机,从而改变了其回声定位声波的频谱时间结构。本研究第一次报道了蝙蝠咀嚼声对其进食行为的影响,为进一步阐明蝙蝠咀嚼声的功能和进化具有重要意义。     

蝙蝠回声定位声波的可塑性及其生态适应

蝙蝠通过调节回声定位声波特征来满足自身的感官需求,表现出回声定位声波的可塑性及其对生态环境与需求的适应。声波频率、强度、脉冲持续时间和间隔时间等特征与蝙蝠所处的生态位密切相关,声波可塑性在蝙蝠进化过程中起着至关重要的作用。本文结合马铁菊头蝠(Rhinolophus ferrumequinum)和大趾鼠耳蝠(Myotis macrodactylus)回声定位声波可塑性的研究,从回声定位声波的方向性、目标距离、环境复杂度和应对干扰4个方面总结了蝙蝠如何通过改变回声定位声波特征来满足自身在导航和捕捉猎物过程中的感官需求与生态适应,并阐述了回声定位声波可塑性的研究现状,为开展蝙蝠声学和行为学研究提供参考。     

吉林省新纪录东方蝙蝠Vespertilio sinensis (Peters, 1880)的回声定位声波特征与分析

通过对吉林省长春市采集到的11只蝙蝠标本的外形、头骨、牙齿和阴茎骨进行测量与对照,鉴定为东方蝙蝠(Vespertilio sinensis),是吉林省翼手目新纪录.用实时录音的超声波仪录制其正常飞行状态下的回声定位声波.结果表明,东方蝙蝠发出短的、宽带的、多谐波的陡坡调频型回声定位声波,能量主要集中在第1谐波.起始频率为83.66±2.08 kHz,峰频为34.54±0.88 kHz,终止频率为24.78±0.41 kHz,带宽为58.84±2.10 kHz,声脉冲持续时间和声脉冲间隔分别为2.63±0.27 ms和61.67±7.5 ms.     

蝙蝠声音信号功能研究进展

声音信号介导动物的资源竞争、配偶选择及反捕食等系列生活史事件,对维持动物种群稳定与群落平衡至关重要。蝙蝠占据夜空生态位,视觉退化,听觉发达,一直被视为声学研究的模式生物。本文围绕蝙蝠回声定位声波与交流声波的功能,综述当前研究现状,展望未来发展方向。纵观已有研究,蝙蝠回声定位声波具有多重功能,包括空间导航与猎物探测、水体与植被识别、协调觅食活动及传递交流信息。蝙蝠利用交流声波从事社群活动,包括社群联系、资源防御、繁殖活动及求救呼叫。绝大多数研究关注蝙蝠回声定位声波的功能,有关蝙蝠交流声波的功能近年才引起重视。未来工作有待进一步探究蝙蝠声音信号产生的影响机制。     

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     

菲菊头蝠不同地理种群回声定位声波差异及其影响因子

回声定位声波地理差异及其形成原因是蝙蝠生态学研究领域一个基本而关键的问题,对于探索物种生存机制、物种形成及其保护具有重要科学意义。本研究从较大地理尺度上(9个地理种群)研究了菲菊头蝠(Rhinolophus pusillus)回声定位声波结构的地理差异,并进一步探讨了影响回声定位声波地理种群差异的因素。结果表明,菲菊头蝠雌性的体型较雄性略大,其主频较高。不同地理种群之间回声定位声波差异明显,包括脉冲持续时间、脉冲间隔、主频以及带宽在不同的地理种群之间均表现出一定程度的差异。进一步分析发现,不同地理种群之间的雌性菲菊头蝠前臂长和体重均与主频呈较弱的负相关,降雨量与雌性的主频呈较强的正相关;而不同地理种群之间的雄性前臂长、体重和降雨量与回声定位声波参数均无相关性;此外,地理距离、温度、湿度均与雌雄回声定位声波参数无相关性。本研究结果表明,菲菊头蝠不同地理种群间的回声定位声波出现明显差异,其中,体型和降雨量为主要影响因子,说明蝙蝠回声定位叫声的进化主要受到了当地生境的影响,表现出动物对不同生境的适应性进化。     

蝙蝠回声定位与捕食对策的研究

蝙蝠的回声定位可以在相当程度上反映出其捕食对策以及栖息环境的特点,回声定位在强度、持续时间及频率等方面的变化模式显示出这类声学信号的多样性,而这种多样性与蝙蝠的捕食对策相关。这方面的研究在国际上历经几十年不衰,然而,在我国,蝙蝠回声定位的研究基本上是空白?..     

Acoustic identification of eight species of bat (mammalia: chiroptera) inhabiting forests of southern hokkaido, Japan: potential for conservation monitoring

Assessing the impact of forest management on bat communities requires a reliable method for measuring patterns of habitat use by individual species. A measure of activity can be obtained by monitoring echolocation calls, but identification of species is not always straightforward. We assess the feasibility of using analysis of time-expanded echolocation calls to identify free-flying bats in the Tomakomai Experimental Forest of Hokkaido University, Hokkaido, northern Japan. Echolocation calls of eight bat species were recorded in one or more of three conditions: from hand-released individuals, from bats flying in a confined space and from bats emerging from their roost. Sonograms of 171 calls from 8 bat species were analyzed. These calls could be categorized into 3 types according to their structure: FM/CF/FM type (Rhinolophus ferrumequinum), FM types (Murina leucogaster, Murina ussuriensis, Myotis macrodactylus and Myotis ikonnikovi) and FM/QCF types (Eptesicus nilssonii, Vespertilio superans and Nyctalus aviator). Sonograms of the calls of R. ferrumequinum could easily be distinguished from those of all other species by eye. For the remaining calls, seven parameters (measures of frequency, duration and inter-call interval) were examined using discriminant function analysis, and 92% of calls were correctly classified to species. For each species, at least 80% of calls were correctly classified. We conclude that analysis of echolocation calls is a viable method for distinguishing between species of bats in the Tomakomai Experimental Forest, and that this approach could be applied to examine species differences in patterns of habitat-use within the forest.     

Testing the performances of automated identification of bat echolocation calls: A request for prudence

Echolocating bats are surveyed and studied acoustically with bat detectors routinely and worldwide, yet identification of species from calls often remains ambiguous or impossible due to intraspecific call variation and/or interspecific overlap in call design. To overcome such difficulties and to reduce workload, automated classifiers of echolocation calls have become popular, but their performance has not been tested sufficiently in the field. We examined the absolute performance of two commercially available programs (SonoChiro and Kaleidoscope) and one freeware package (BatClassify). We recorded noise from rain and calls of seven common bat species with Pettersson real-time full spectrum detectors in Sweden. The programs could always (100%) distinguish rain from bat calls, usually (68–100%) identify bats to group (Nyctalus/Vespertilio/Eptesicus, Pipistrellus, Myotis, Plecotus, Barbastella) and usually (83–99%) recognize typical calls of some species whose echolocation pulses are structurally distinct (Pipistrellus pygmaeus, Barbastella barbastellus). Species with less characteristic echolocation calls were not identified reliably, including Vespertilio murinus (16–26%), Myotis spp. (4–93%) and Plecotus auritus (0–89%). All programs showed major although different shortcomings and the often poor performance raising serious concerns about the use of automated classifiers for identification to species level in research and surveys. We highlight the importance of validating output from automated classifiers, and restricting their use to specific situations where identification can be made with high confidence. For comparison we also present the result of a manual identification test on a random subset of the files used to test the programs. It showed a higher classification success but performances were still low for more problematic taxa.     

Traffic noise differentially impacts call types in a Japanese treefrog (Buergeria japonica)

Acoustic noise from automobile traffic impedes communication between signaling animals. To overcome the acoustic interference imposed by anthropogenic noise, species across taxa adjust their signaling behavior to increase signal saliency. As most of the spectral energy of anthropogenic noise is concentrated at low acoustic frequencies, species with lower frequency signals are expected to be more affected. Thus, species with low-frequency signals are under stronger pressure to adjust their signaling behaviors to avoid auditory masking than species with higher frequency signals. Similarly, for a species with multiple types of signals that differ in spectral characteristics, different signal types are expected to be differentially masked. We investigate how the different call types of a Japanese stream breeding treefrog (Buergeria japonica) are affected by automobile traffic noise. Male B. japonica produce two call types that differ in their spectral elements, a Type I call with lower dominant frequency and a Type II call with higher dominant frequency. In response to acoustic playbacks of traffic noise, B. japonica reduced the duration of their Type I calls, but not Type II calls. In addition, B. japonica increased the call effort of their Type I calls and decreased the call effort of their Type II calls. This result contrasts with prior studies in other taxa, which suggest that signalers may switch to higher frequency signal types in response to traffic noise. Furthermore, the increase in Type I call effort was only a short-term response to noise, while reduced Type II call effort persisted after the playbacks had ended. Overall, such differential effects on signal types suggest that some social functions will be disrupted more than others. By considering the effects of anthropogenic noise across multiple signal types, these results provide a more in-depth understanding of the behavioral impacts of anthropogenic noise within a species.     

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。     

Morphology, echolocation and foraging behaviour in two sympatric sibling species of bat (Tylonycteris pachypus and Tylonycteris robustula) (Chiroptera: Vespertilionidae)

We studied the wing morphology, echolocation calls, foraging behaviour and flight speed of Tylonycteris pachypus and Tylonycteris robustula in Longzhou County, South China during the summer (June–August) of 2005. The wingspan, wing loading and aspect ratio of the two species were relatively low, and those of T. pachypus were lower compared with T. robustula . The echolocation calls of T. pachypus and T. robustula consist of a broadband frequency modulated (FM) sweep followed by a short narrowband FM sweep. The dominant frequency of calls of T. pachypus was 65.1 kHz, whereas that of T. robustula was 57.7 kHz. The call frequencies (including highest frequency of the call, lowest frequency of the call and frequency of the call that contained most energy) of T. pachypus were higher than those of T. robustula , and the pulse duration of the former was longer than that of the latter. The inter-pulse interval and bandwidth of the calls were not significantly different between the two species. Tylonycteris pachypus foraged in more complex environments than T. robustula , although the two species were both netted in edge habitats (around trees or houses), along pathways and in the tops of trees. Tylonycteris pachypus flew slower (straight level flight speed, 4.3 m s⁻¹) than T. robustula (straight level flight speed, 4.8 m s⁻¹). We discuss the relationship between wing morphology, echolocation calls, foraging behaviour and flight speed, and demonstrate resource partitioning between these two species in terms of morphological and behavioural factors.     

沼水蛙繁殖期鸣声特征及鸣叫节律

鸣叫对无尾两栖类动物的生存与繁殖起重要作用。蛙类的鸣叫行为受到环境因素影响表现出一定的节律性。2016年8和9月,采用录音机和指向性话筒,在野外录制了57只沼水蛙（Hylarana guentheri）的鸣声并对其鸣声特征进行分析;通过悬挂录音笔和自动温湿度记录仪研究了沼水蛙鸣叫节律（17 d）及其与环境温度、相对湿度的关系。结果显示,沼水蛙的鸣声由1 ~ 4个音节组成,不同类型鸣声间的音节主频、音节时长存在显著差异（P < 0.05）。该物种全天具有鸣叫行为,13:00 ~ 14:00时为鸣叫高峰期。白天单音节鸣声、双音节鸣声、三音节鸣声、总鸣声和总音节的数量较夜晚显著增加（P < 0.01）。鸣声数量和音节数量均与环境温度呈正相关（P < 0.01）。结果表明,沼水蛙通过改变音节数量、音节主频和音节时长改变鸣叫策略。沼水蛙的鸣叫行为具有昼夜节律性且受环境温度的影响。     

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.     

马铁菊头蝠(Rhinolophus ferrumequinum)回声定位声波与生境类型和环境因子的关系

2007年在吉林省罗通山自然保护区,利用超声波探测仪（Avisoft-SASLAB PRO）录制并分析不同生境中马铁菊头蝠的回声定位声波。结果显示马铁菊头蝠在不同类型生境中活动;各生境中回声定位声波参数存在显著差异（one-way ANOVA,P〈0.05）。从环境因子中通过主成分分析筛选出与其回声定位声波相关的植被、气候和地形因子,探讨回声定位声波与这些因子的相关性。结果显示FM1和FM2带宽与乔木高（r=-0.948,-0.825;P〈0.05）、FM1起始频率和FM2终止频率与林冠面积（r=-0.967,-0.958;P〈0.05）、FM1起始频率、FM2终止频率和峰频与湿度（r=-0.776、-0.875和-0.794,P〈0.05）、脉冲持续时间和脉冲间隔与平均灌木高均呈显著负相关（r=-0.911,-0.990;P〈0.05）,峰频与植被株数（r=0.756,P〈0.05）、脉冲持续时间与冠下高呈显著正相关（r=0.870,P〈0.05）。表明各种环境因子（植被因子、气候因子和地形因子）都在一定程度上影响回声定位声波,回声定位声波具有表型可塑性和生境适应性,这些特性决定了马铁菊头蝠生境利用的程度和可利用的资源。