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。     

马铁菊头蝠捕食活动与猎物资源的关系

捕食者与猎物的关系研究对了解物种捕食行为及种群空间格局具有重要意义。采用Avisoft Bioacoustics超声波仪录制马铁菊头蝠自然状态下的声波以确定其捕食活动强度,用灯诱法、扫网法和飞行阻隔法相结合采集昆虫,搜集蝙蝠粪便并分析其食物组成。结果表明,马铁菊头蝠在8月份活动最频繁,昆虫丰富度在8月份最丰富,马铁菊头蝠捕食活动与鳞翅目丰富度呈显著正相关。马铁菊头蝠主要以鳞翅目和鞘翅目为食,但食性存在明显的月份变化。卡方检验结果表明,马铁菊头蝠捕食的猎物与环境中可利用的昆虫猎物存在显著差异。在食物资源丰富时,马铁菊头蝠选择性地捕食营养丰富的鞘翅目昆虫。     

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。     

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。     

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。     

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.     

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。     

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。     

形态和声波相似的中华菊头蝠与中菊头蝠的共存机制

研究了同域分布的中华菊头蝠(Rhinolophus sinicus)与中菊头蝠(Rhinolophus affinis)的食性、形态、回声定位声波及捕食时间.中华菊头蝠与中菊头蝠均属于中等体型的菊头蝠,前臂长分别为(51.25±0.22) mm和(52.40±0.37) mm;悬挂状态下的回声定位声波均为典型的调频-恒频-调频(FM-CM-FM)型叫声,峰频分别为(82.07±0.17) kHz和(84.41±0.48) kHz.粪便分析显示中华菊头蝠与中菊头蝠分别捕食9目和7目昆虫,均以鳞翅目(Lepidiptera)和鞘翅目(Coleoptera)昆虫为主要食物(体积百分比总和> 90%),捕食鳞翅目昆虫的体积百分比差异显著,对猎物大小(以鞘翅目昆虫体长衡量)的选择无显著差异.中华菊头蝠与中菊头蝠的营养生态位宽度分别为2.38和2.28,重叠度达0.91,营养生态位未发生明显分化,但充足的食物资源促进了二者的共存.另外,2种菊头蝠的感官生态位和时间生态位未发生明显分化.由2种菊头蝠的翼载和峰频的差异推测二者发生了空间生态位和捕食微生境的分化,这也可能促进了二者的共存.     

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。     

‘No cost of echolocation for flying bats’ revisited

Echolocation is energetically costly for resting bats, but previous experiments suggested echolocation to come at no costs for flying bats. Yet, previous studies did not investigate the relationship between echolocation, flight speed, aerial manoeuvres and metabolism. We re-evaluated the 'no-cost' hypothesis, by quantifying the echolocation pulse rate, the number of aerial manoeuvres (landings and U-turns), and the costs of transport in the 5-g insectivorous bat Rhogeessa io (Vespertilionidae). On average, bats (n = 15) travelled at 1.76 ± 0.36 m s?1 and performed 11.2 ± 6.1 U-turns and 2.8 ± 2.9 ground landings when flying in an octagonal flight cage. Bats made more U-turns with decreasing wing loading (body weight divided by wing area). At flight, bats emitted 19.7 ± 2.7 echolocation pulses s?1 (range 15.3-25.8 pulses s?1), and metabolic rate averaged 2.84 ± 0.95 ml CO? min?1, which was more than 16 times higher than at rest. Bats did not echolocate while not engaged in flight. Costs of transport were not related to the rate of echolocation pulse emission or the number of U-turns, but increased with increasing number of landings; probably as a consequence of slower travel speed when staying briefly on ground. Metabolic power of flight was lower than predicted for R. io under the assumption that energetic costs of echolocation call production is additive to the aerodynamic costs of flight. Results of our experiment are consistent with the notion that echolocation does not add large energetic costs to the aerodynamic power requirements of flight in bats.     

Interspecific responses to distress calls in bats (Chiroptera: Vespertilionidae): a function for convergence in call design?

Distress calls were recorded from three sympatric species of pipistrelle bat (Pipistrellus nathusii, P. pipistrellus and P. pygmaeus) in England and Northern Ireland. At foraging sites, we conducted playback experiments, consisting of experimental distress call sequences from each species and control sequences of random noise and sound recorded with no bats present. We measured response by simultaneously recording ultrasound during playbacks and counting the echolocation pulses above a predetermined threshold which were then identified to species. All three species responded to each other's calls. The number of recorded echolocation pulses of all species increased eight-fold, on average, during the playback of distress call sequences compared with the playback of ultrasonic noise, and four-fold compared with the playback of silence. In a separate playback experiment, the number of echolocation pulses of P. pygmaeus increased 14-fold during the playback of distress calls of four endemic species of bat from Madagascar (Emballonura atrata, Myotis goudoti, Miniopterus majori and M. manavi) compared with the playback of silence. This increased response might have been caused by the high calling rates of the Malagasy species. Distress calls of P. nathusii, P. pipistrellus and P. pygmaeus were structurally convergent, consisting of a series of downward-sweeping, frequency-modulated elements of short duration and high intensity with a relatively strong harmonic content. Selection may favour convergence in the structure of distress calls among bat species, if attracting heterospecifics increases the chance of repelling predators by mobbing.     

Female big brown bats, Eptesicus fuscus, recognize sex from a caller's echolocation signals

The echolocation calls of bats function in prey capture and navigation but are not commonly regarded as communicative signals. However, because bats' echolocation calls show patterns of variability, they may transmit information about a bat, such as its age, individual identity or sex. For echolocation calls to function in this manner, variation in calls must be reliably linked to the characteristics of the bat, as has been shown in a number of studies. However, few studies have asked whether bats respond to this variation. We tested whether female big brown bats can identify the sex of an unfamiliar bat from playbacks of its echolocation calls. Playback consisted of a 30-s preplayback period, a 60-s playback period of either male or female echolocation calls, and a 30-s postplayback period. In the playback and postplayback periods the vocalization rates of female bats changed significantly relative to the preplayback period depending on the sex of the playback stimulus, indicating that they could determine sex from the echolocation calls. These findings support the possibility that echolocation calls play a role in communication in big brown bats.     

Tonotopical organization and pure tone response characteristics of single units in the auditory cortex of the Greater Horseshoe Bat

Summary Tonotopical organization and frequency representation in the auditory cortex of Greater Horseshoe Bats was studied using multi-unit recordings.The auditory responsive cortical area can be divided into a primary and a secondary region on the basis of response characteristics forming a core/belt structure.In the primary area units with best frequencies in the range of echolocation signals are strongly overrepresented (Figs. 6–8). There are two separate large areas concerned with the processing of the two components of the echolocation signals. In one area frequencies between the individual resting frequency and about 2 kHz above are represented, which normally occur in the constant frequency (CF) part of the echoes (CF-area), in a second one best frequencies between resting frequency and about 8 kHz below are found (FM-area).In the CF-area tonotopical organization differs from the usual mammalian scheme of dorso-ventral isofrequency slabs. Here isofrequency contours are arranged in a semicircular pattern.The representation of the cochlear partition (cochleotopic organization) was calculated. In the inferior colliculus and auditory cortex there is a disproportionate representation of the basilar membrane. This finding is in contradiction to the current opinion that frequency representation in the auditory system of Horseshoe Bats is only determined by the mechanical tuning properties of the basilar membrane.Response characteristics for single units were studied using pure tone stimuli. Most units showed transient responses. In 25% of units response characteristics depended on the combination of frequency and sound pressure level used.Frequency selectivity of units with best frequencies in the range of echolocation sounds is very high. Q-10dB values of up to 400 were found in a small frequency band just above resting frequency.Abbreviations BF best frequency - CF constant frequency - FM frequency modulated - MT minimal threshold     

东方蝙蝠在交通噪声环境中的回声定位行为

环境噪声会降低动物声信号可探测性,是动物声信号进化的压力。为了避免人工噪声干扰,多数脊椎动物调整其叫声的频谱-时间结构,如采用延长叫声持续时间、提高频率和增加强度等策略。本研究选择回声定位声波频率范围与交通噪声频率范围相互重叠的东方蝙蝠(Vespertilio sinensis)为研究对象,在自然条件下开展实验,研究交通噪声对蝙蝠回声定位声波的影响。本实验选择交通噪声强度存在差异的两个样点,分别录制东方蝙蝠的回声定位声波,分析其结构参数。在交通噪声强度较高的样点,东方蝙蝠回声定位声波的持续时间、斜率均未显著改变,但起始频率、主频、终止频率及带宽均显著提高。结果表明,东方蝙蝠的回声定位行为在交通噪声干扰下具有明显的可塑性。     

Prestin Shows Divergent Evolution Between Constant Frequency Echolocating Bats

The gene Prestin encodes a motor protein that is thought to confer the high-frequency sensitivity and selectivity that characterizes the mammalian auditory system. Recent research shows that the Prestin gene has undergone a burst of positive selection on the ancestral branch of the Old World horseshoe and leaf-nosed bats (Rhinolophidae and Hipposideridae, respectively), and also on the branch leading to echolocating cetaceans. Moreover, these two groups share a large number of convergent amino acid sequence replacements. Horseshoe and leaf-nosed bats exhibit narrowband echolocation, in which the emitted calls are based on the second harmonic of a predominantly constant frequency (CF) component, the frequency of which is also over-represented in the cochlea. This highly specialized form of echolocation has also evolved independently in the neotropical Parnell's mustached bat (Pteronotus parnellii). To test whether the convergent evolution of CF echolocation between lineages has arisen from common changes in the Prestin gene, we sequenced the Prestin coding region (~2,212?bp, >99% coverage) in P. parnellii and several related species that use broadband echolocation calls. Our reconstructed Prestin gene tree and amino acid tree showed that P. parnellii did not group together with Old World horseshoe and leaf-nosed bats, but rather clustered within its true sister species. Comparisons of sequences confirmed that P. parnellii shared most amino acid changes with its congeners, and we found no evidence of positive selection in the branch leading to the genus of Pteronotus. Our result suggests that the adaptive changes seen in Prestin in horseshoe and leaf-nosed bats are not necessary for CF echolocation in P. parnellii.     

Rediscovery of short-tailed bats (

Short-tailed bats (Mystacina sp.) were rediscovered in Nothofagus dominant rainforest in the Eglinton Valley in February 1997, representing the first records of these bats in Fiordland since 1871. Breeding females, adult males and juveniles were captured. This paper presents preliminary observations of taxonomy, echolocation calls, population size, habitat use, activity patterns, home range size, movements, roosting, and singing behaviour. Compared to lesser short- tailed bats (M. tuberculata) on Codfish and Little Barrier Islands, the Fiordland bats were heavier, had larger wings and smaller ears, and were sexually dimorphic. The Mystacina echolocation calls were of low intensity (quiet), making them difficult to detect. Call durations in free-flying bats were only 1.0-2.9 ms long. In a comparative trial the majority of calls that were detected at 25 kHz using the Batbox III bat detector were not recorded at 40 kHz, indicating that there was little overlap with the calls of long-tailed bats (Chalinolobus tuberculatus). In February, roosting groups numbered from 107 to 279 individuals and the bats ranged over 130 km(2) of the valley. Bats began emerging c. 20 minutes after sunset and were active at the roost sites throughout the night. Radio-tagged bats were active for an average of 372 minutes at a time. All roosts were in large diameter (67-146 cm dbh) red beech (N. fusca) trees.     

Eavesdropping by Bats: The Influence of Echolocation Call Design and Foraging Strategy

We used playback presentations to free-flying bats of 3 species to assess the influence of echolocation call design and foraging strategy on the role of echolocation calls in communication. Near feeding sites over water, Myotis lucifugus and M. yumanensis responded positively only to echolocation calls of conspecifics. Near roosts, these bats did not respond before young of the year became volant, and after this responded to presentations of echolocation calls of similar and dissimilar design. At feeding sites Lasiurus borealis responded only to echolocation calls of conspecifics and particularly to “feeding buzzes”. While Myotis, particularly subadults, appear to use the echolocation calls of conspecifics to locate feeding sites, L. borealis appears to use the calls of a foraging neighbour attacking prey to identify opportunities for ‘stealing’ food.