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。     

五种蝙蝠形态与回声定位叫声的性别差异

为研究角菊头蝠(Rhinolophus cornutus)、三叶蹄蝠(Aselliscus uheeleri)、大蹄蝠(Hipposideros armiger)、大鼠耳蝠(Myotis myotis)和大足鼠耳蝠(M．riketti)的形态和回声定位叫声的性别差异性,明确同种蝙蝠雌雄个体对食物、栖息地等资源利用的细微差异,我们利用超声波探测仪、Batsound分析软件及SPSS11．0统计软件对5种95只蝙蝠进行了录音、声波分析和统计分析。5种蝙蝠形态性别差异性不显著,角菊头蝠、三叶蹄蝠、大蹄蝠和大足鼠耳蝠叫声频率性别差异性显著,大鼠耳蝠叫声频率性别差异性不显著。角菊头蝠雌性叫声的基频、分音、主频率高于雄性,声脉冲时间、间隔时间大于雄性,调频(FM)带宽小于雄性;三叶蹄蝠、大蹄蝠叫声的基频、主频率雄性高于雌性,调频带宽雌性小于雄性;大足鼠耳蝠叫声的主频率雄性高于雌性,FM带宽雌性大于雄性[动物学报49(6)：742～747,2003]。     

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。     

两种同域分布蹄蝠在开阔度不同的环境中回声定位声波的可塑性

在广西桂林研究了同域分布的大蹄蝠(Hipposideros armiger)和中蹄蝠(H.larvatus)在不同开阔度环境中回声定位声波信号的变化。用超声波仪录制自由悬挂和分别释放于人工"大棚"和"小棚"内飞行的蝙蝠的回声定位声波,使用超声分析软件分析声脉冲时程、主频率及声脉冲间隔,通过重复测量方差分析比较不同状态下的声波参数。结果表明:中蹄蝠声波的主频在悬挂状态下最高,小棚内飞行时次之,大棚内飞行最低;两种蹄蝠声波的脉冲时程和脉冲间隔在悬挂状态下最长,大棚内飞行次之,小棚内飞行最低。总之,这两种蹄蝠的回声定位声波能够随所处状态的变化而变化,可根据生境的复杂度调节声讯号,具有明显的声波可塑性。     

江苏省蝙蝠新纪录——皮氏菊头蝠

2008年5月在江苏省采到8只蝙蝠,经鉴定为皮氏菊头蝠(Rhinolophus pearsoni),为该省翼手目新纪录,这意味着我国大陆秦岭淮河以南地区均有该物种分布.江苏样本比文献记载的其他地区样本体重稍轻,体型略小,但回声定位声波主频率略高,符合菊头蝠主频率与体型大小成负相关的普遍规律.     

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。     

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。     

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。     

Determinants of echolocation call frequency variation in the Formosan lesser horseshoe bat (Rhinolophus monoceros)

The origin and maintenance of intraspecific variation in vocal signals is important for population divergence and speciation. Where vocalizations are transmitted by vertical cultural inheritance, similarity will reflect co-ancestry, and thus vocal divergence should reflect genetic structure. Horseshoe bats are characterized by echolocation calls dominated by a constant frequency component that is partly determined by maternal imprinting. Although previous studies showed that constant frequency calls are also influenced by some non-genetic factors, it is not known how frequency relates to genetic structure. To test this, we related constant frequency variation to genetic and non-genetic variables in the Formosan lesser horseshoe bat (Rhinolophus monoceros). Recordings of bats from across Taiwan revealed that females called at higher frequencies than males; however, we found no effect of environmental or morphological factors on call frequency. By comparison, variation showed clear population structure, with frequencies lower in the centre and east, and higher in the north and south. Within these regions, frequency divergence was directional and correlated with geographical distance, suggesting that call frequencies are subject to cultural drift. However, microsatellite clustering analysis showed that broad differences in constant frequency among populations corresponded to discontinuities in allele frequencies resulting from vicariant events. Our results provide evidence that the processes shaping genetic subdivision have concomitant consequences for divergence in echolocation call frequency.     

皮氏菊头蝠回声定位声波与年龄的关系

皮氏菊头蝠 (Rhinolophuspearsoni)雌性成体 5只和幼体 2只采自贵州省贞丰县珉谷镇。采用超声波探测仪 (D980 ,ULTRASOUNDDETECTOR)接收皮氏菊头蝠的回声定位声波 ,转换到原频率的 1 / 1 0后导入计算机 ,然后用专业声谱分析软件 (Batsound 3 1 0 )进行分析。成蝠在飞行和悬挂状态下的声波结构相似 ,只是声波各项参数值略有不同 :它们发射FM CF FM型声波 ,具有 2～ 3个谐波 ,主频率在飞行时为 5 6 80± 0 6 2kHz ,悬挂时为 5 8 0 5± 0 2 4kHz ;声脉冲时间和间隔在飞行时分别为 3 4 6 2± 5 2 9ms和 86 5 0± 1 9 72ms ,悬挂时分别为 4 1 0 8± 5 87ms和 1 1 7 2 9± 6 6 4 4ms ;能率环飞行时为 ( 4 4 0 6± 1 2 5 8) % ,悬挂时为 ( 4 6 0 0±2 4 2 5 ) %。幼蝠声波为CF FM型 ,谐波数为 5～ 8个 ,主频率明显低于成体 ,FM带宽窄于成体 ,声脉冲时间和间隔短于成体 ,能率环低于成体。皮氏菊头蝠回声定位声波与年龄有关 ,这可能因成体的声波主要是探测食物和周围环境的详细信息 ,而幼体主要是与母蝠进行交流。     

Morphological correlates of echolocation frequency in the endemic Cape horseshoe bat, <Emphasis Type="Italic">Rhinolophus capensis</Emphasis> (Chiroptera: Rhinolophidae)

We investigated intraspecific variation in echolocation calls of the Cape horseshoe bat, Rhinolophus capensis, by comparing echolocation and associated morphological parameters among individuals from three populations of this species. The populations were situated in the center and at the western and eastern limits of the distribution of R. capensis. The latter two populations were situated in ecotones between vegetation biomes. Ecotone populations deviated slightly from the allometric relationship between body size and peak frequency for the genus, and there was no relationship between these variables within R. capensis. Nasal chamber length was the best predictor of peak frequency but not correlated with body size. The evolution of echolocation thus appears to have been uncoupled from body size in R. capensis. Furthermore, females used higher frequencies than males, which imply a potential social role for peak frequency. The differences in peak frequency may have originated from random founder effects and then compounded by genetic drift and/or natural selection. The latter may have acted directly on peak frequency altering skull parameters involved in echolocation independently of body size, resulting in the evolution of local acoustic signatures.     

贵州发现毛翼管鼻蝠和华南菊头蝠及其回声定位声波特征

2017年5月分别在贵州省兴义市和安龙县用雾网采集到1只大体型雌性管鼻蝠和2只小体型雄性菊头蝠,经鉴定为毛翼管鼻蝠(Harpiocephalus harpia)和华南菊头蝠(Rhinolophus huananus),属贵州省翼手目新分布记录。所采集的毛翼管鼻蝠雌性个体可能为怀孕个体,故测量相关数据并鉴定后原地放归野外。华南菊头蝠标本保存于东北师范大学环境学院。毛翼管鼻蝠(1♀):鼻部呈短管状,全身被毛厚密而柔软,后足、翼膜和尾膜均被覆黄褐色绒毛;体重14.01 g,前臂长48.23 mm;回声定位声波为调频(FM)型,静止状态下声波峰频为(56.04±4.52)k Hz。华南菊头蝠(2♂):耳大,对耳屏相对较小;鼻叶之蹄状叶宽大,完全覆盖吻部;鞍状叶小,其高略超过其宽,近乎矩形;体毛烟褐色;2只样本体重分别为4.52 g、4.12 g,前臂长40.70 mm、40.00 mm;头骨狭长,颅全长16.35 mm、16.46 mm,颅宽8.11 mm、8.14 mm;回声定位声波属调频-恒频-调频(FM-CF-FM)型,静止状态下峰频分别为(68.48±0.08)k Hz、(67.18±0.06)k Hz。2种蝙蝠物种在贵州省为首次发现,扩大了其在国内的分布范围,丰富了生态资料,为进一步的研究和保护提供了基础资料。     

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.     

大足鼠耳蝠的翼型和回声定位声波特征

大足鼠耳蝠(Myotis ricketti Thomas,1894)属翼手目(Chiroptera),蝙蝠科(Vespertilionidae),鼠耳蝠属(Myotis).     

Female Mate Choice Can Drive the Evolution of High Frequency Echolocation in Bats: A Case Study with Rhinolophus mehelyi

Animals employ an array of signals (i.e. visual, acoustic, olfactory) for communication. Natural selection favours signals, receptors, and signalling behaviour that optimise the received signal relative to background noise. When the signal is used for more than one function, antagonisms amongst the different signalling functions may constrain the optimisation of the signal for any one function. Sexual selection through mate choice can strongly modify the effects of natural selection on signalling systems ultimately causing maladaptive signals to evolve. Echolocating bats represent a fascinating group in which to study the evolution of signalling systems as unlike bird songs or frog calls, echolocation has a dual role in foraging and communication. The function of bat echolocation is to generate echoes that the calling bat uses for orientation and food detection with call characteristics being directly related to the exploitation of particular ecological niches. Therefore, it is commonly assumed that echolocation has been shaped by ecology via natural selection. Here we demonstrate for the first time using a novel combined behavioural, ecological and genetic approach that in a bat species, Rhinolophus mehelyi: (1) echolocation peak frequency is an honest signal of body size; (2) females preferentially select males with high frequency calls during the mating season; (3) high frequency males sire more off-spring, providing evidence that echolocation calls may play a role in female mate choice. Our data refute the sole role of ecology in the evolution of echolocation and highlight the antagonistic interplay between natural and sexual selection in shaping acoustic signals.