六种共栖蝙蝠的回声定位信号及翼型特征的比较

本研究于2006 年5 ～ 8 月在桂林市七星公园七星岩洞进行,比较分析了共栖2 科(蹄蝠科和蝙蝠科)6 种共75 只蝙蝠的回声定位信号和翼型特征。普氏蹄蝠的回声定位叫声为短而多谐波的CF/ FM 型,主频率为61.2±0.8 kHz, 具有高翼载、低翼展比和中等翼尖指数; 大蹄蝠的回声定位叫声为单CF/ FM 型,主频率为68. 6 ±0.7 kHz,具有高翼载、低翼展比和中等翼尖指数;中蹄蝠的回声定位叫声为单CF / FM 型,主频率为85.2 ±0.5 kHz,具有中等翼载、低翼展比和中等翼尖指数;高颅鼠耳蝠的回声定位叫声为长带宽的FM 型,主频率为50.7 ±3.8 kHz,具有低翼载、中等翼展比和低翼尖指数;大足鼠耳蝠回声定位叫声为FM 型,主频率为39.9 ±3.2 kHz,具有中等翼载、低翼展比和高翼尖指数;绒山蝠回声定位叫声为短而多谐波的FM 型,主频率为49.0± 0. 4 kHz,具有高翼载、中等翼展比和低翼尖指数。经单因素方差分析表明,6 种蝙蝠之间绝大部分的形态和声音参数差异显著(One-way ANOVA,P < 0. 05)。以上结果说明,6 种同地共栖蝙蝠种属特异的回声定位叫声
和形态结构体现出了相互之间的生态位分离,从而降低了种间竞争压力,使得6 种蝙蝠能够同地共存。     

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.     

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).     

Identification of sympatric bat species by the echolocation calls

One hundred and thirty-eight echolocation calls of 63 free-flying individuals of five bat species (Rhinolophus ferrumequinum, Myotis formosus, Myotis ikonnikovi, Myotis daubentoni and Murina leucogaster) were recorded (by ultrasonic bat detector (D980)) in Zhi’an village of Jilin Province, China. According to the frequency-time spectra, these calls were categorized into two types: FM/CF (constant frequency) / FM (R. ferrumequinum) and FM (frequency modulated) (M. formosus, M. ikonnikovi, M. daubentoni and M. leucogaster). Sonograms of the calls of R. ferrumequinum could easily be distinguished from those of the other four species. For the calls of the remaining four species, six echolocation call parameters, including starting frequency, ending frequency, peak frequency duration, longest inter-pulse interval and shortest inter-pulse interval, were examined by stepwise discriminant analysis. The results show that 84.1% of calls were correctly classified, which indicates that these parameters of echolocation calls play an important role in identifying bat species. These parameters can be used to test the accuracy of general predictions based on bats’ morphology in the same forest and can provide essential information for assessing patterns of bat habitat use. __________ Translated from Journal of Northeast Normal University (Natural Science Edition), 2006, 38 (3): 109–114 [译自: 东北师范大学学报(自然科学版)]     

Identification of sympatric bat species by the echolocation calls

One hundred and thirty-eight echolocation calls of 63 free-flying individuals of five bat species (Rhinolophus ferrumequinum,Myotis formosus,Myotis ikonnikovi,Myotis daubentoni and Murina leucogaster)were recorded (by ultrasonic bat detector (D980)) in Zhi'an village of Jilin Province,China.According to the frequency-time spectra,these calls were categorized into two types:FM/CF (constant frequency) / FM (R.ferrumequinum) and FM (frequency modulated)(M.formosus,M.ikonnikovi,M.daubentoni and M.leucogaster).Sonograms of the calls of R.ferrumequinum could easily be distinguished from those of the other four species.For the calls of the remaining four species,six echolocation call parameters,including starting frequency,ending frequency,peak frequency duration,longest inter-pulse interval and shortest inter-pulse interval,were examined by stepwise discriminant analysis.The results show that 84.1% of calls were correctly classified,which indicates that these parameters of echolocation calls play an important role in identifying bat species.These parameters can be used to test the accuracy of general predictions based on bats' morphology in the same forest and can provide essential information for assessing patterns of bat habitat use.     

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。     

Variations in the echolocation calls of the European free-tailed bat

The echolocation calls of Tadarida teniotis were studied in an outdoor flight enclosure (captive individuals) and in the wild using single microphones or an array of four microphones. Calls were characterized by measures of 10 call variables. Comparison of individual calls recorded on four microphones arrayed in a tetrahedron with 1 m between each microphone revealed that all calls were not equally detectable by all microphones but that there were no significant differences in call features obtained from calls recorded on all four microphones. A comparison of 47 calls recorded by all four microphones showed no significant differences in the features of the four recordings of each call. Analysis of calls of five individuals flying individually in an outdoor flight cage revealed significant individual differences in call features. In the field, T. teniotis used long, narrowband search-phase calls, usually without harmonics. Analysis of 1876 search-phase echolocation calls of T. teniotis recorded in the field in Israel and Greece in 2002, 2005 and 2006 showed significant year-to-year and site-to-site differences in some call features. When flying in the presence of conspecifics, T. teniotis changed their echolocation calls. We found a range of different buzzes in the wild, and based on their structure we attempted to classify them as feeding and social buzzes. The features of individual calls comprising buzzes differed significantly among buzzes, and yet there were no consistent differences between what we classified as feeding and social buzzes.     

Postnatal development in the big-footed bat,Myotis macrodactylus: wing morphology,echolocation calls,and flight

We studied the postnatal development of wing morphology and echolocation calls during flight in a free-ranging population of the big-footed bat, Myotis macrodactylus, using the mark-recapture methodology. Young bats were reluctant to move until 7 days of age and started fluttering at a mean age of 10 days. The wingspan and wing area of pups followed a linear pattern of growth until 22 days of age, by which time the young bats exhibited flapping flight, with mean growth rates of 0.62 mm/day and 3.15 mm²/day, for wingspan and area, respectively, after which growth rates decreased. Pups achieved sustained flight at 40 days of age. Of the three nonlinear growth models (logistic, Gompertz, and von Bertalanffy), the logistic equation provided the best fit to the empirical curves for wingspan and wing area. Neonates emitted long echolocation calls with multiple harmonics. The duration of calls decreased significantly between flutter (19 days) and flight (22 days) stages. The peak and start frequency of calls increased significantly over the 3-week period of development, but the terminal frequency did not change significantly over the development period.     

A comparison of supervised learning techniques in the classification of bat echolocation calls

Today's acoustic monitoring devices are capable of recording and storing tremendous amounts of data. Until recently, the classification of animal vocalizations from field recordings has been relegated to qualitative approaches. For large-scale acoustic monitoring studies, qualitative approaches are very time-consuming and suffer from the bias of subjectivity. Recent developments in supervised learning techniques can provide rapid, accurate, species-level classification of bioacoustics data. We compared the classification performances of four supervised learning techniques (random forests, support vector machines, artificial neural networks, and discriminant function analysis) for five different classification tasks using bat echolocation calls recorded by a popular frequency-division bat detector. We found that all classifiers performed similarly in terms of overall accuracy with the exception of discriminant function analysis, which had the lowest average performance metrics. Random forests had the advantage of high sensitivities, specificities, and predictive powers across the majority of classification tasks, and also provided metrics for determining the relative importance of call features in distinguishing between groups. Overall classification accuracy for each task was slightly lower than reported accuracies using calls recorded by time-expansion detectors. Myotis spp. were particularly difficult to separate; classifiers performed best when members of this genus were combined in genus-level classification and analyzed separately at the level of species. Additionally, we identified and ranked the relative contributions of all predictor features to classifier accuracy and found measurements of frequency, total call duration, and characteristic slope to be the most important contributors to classification success. We provide recommendations to maximize accuracy and efficiency when analyzing acoustic data, and suggest an application of automated bioacoustics monitoring to contribute to wildlife monitoring efforts.     

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]。     

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。     

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.     

Evolutionary aspects of bat echolocation

This review is yet another attempt to explain how echolocation in bats or bat-like mammals came into existence. Attention is focused on neuronal specializations in the ascending auditory pathway of echolocating bats. Three different mechanisms are considered that may create a specific auditory sensitivity to echos: (1). time-windows of enhanced echo-processing opened by a corollary discharge of neuronal vocalization commands; (2). differentiation and expansion of ensembles of combination-sensitive neurons in the midbrain; and (3). corticofugal top-down modulations. The second part of the review interprets three different types of echolocation as adaptations to ecological niches, and presents the sophisticated cochlear specializations in constant-frequency/frequency-modulated bats as a case study of finely tuned differentiation. It is briefly discussed how a resonant mechanism in the inner ear of constant-frequency/frequency-modulated bats may have evolved in common mammalian cochlea.     

Geographical and individual variation in echolocation calls of the intermediate leaf‐nosed bat,Hipposideros larvatus

The cause and significance of variation in echolocation call frequency within hipposiderid bats is not well understood despite an increasing number of allopatric and sympatric examples being documented. We examined variation patterns in the resting frequency (RF) of echolocation calls emitted by the intermediate leaf‐nosed bat, Hipposideros larvatus, on a broad geographical scale. Data mining technology and Kruskal–Wallis test both showed substantial variation with a longitudinal pattern in RF in H. larvatus among colonies, and this variation was associated with geographical distance and not body size. In addition, we found that a high degree of variability between individuals was hidden under the geographical variation. The results support an effect of random cultural drift, and challenge the prey detection hypothesis. Moreover, an acoustic difference among local island colonies may be indicative of a vocal dialect. We found that each colony of H. larvatus seems to maintain a ‘private bandwidth’, which could be used for colony identity and individual communication thus helping individuals and colonies to get a number of fitness benefits.     

To females of a noctuid moth,male courtship songs are nothing more than bat echolocation calls

It has been proposed that intraspecific ultrasonic communication observed in some moths evolved, through sexual selection, subsequent to the development of ears sensitive to echolocation calls of insectivorous bats. Given this scenario, the receiver bias model of signal evolution argues that acoustic communication in moths should have evolved through the exploitation of receivers'' sensory bias towards bat ultrasound. We tested this model using a noctuid moth Spodoptera litura, males of which were recently found to produce courtship ultrasound. We first investigated the mechanism of sound production in the male moth, and subsequently the role of the sound with reference to the female''s ability to discriminate male courtship songs from bat calls. We found that males have sex-specific tymbals for ultrasound emission, and that the broadcast of either male songs or simulated bat calls equally increased the acceptance of muted males by the female. It was concluded that females of this moth do not distinguish between male songs and bat calls, supporting the idea that acoustic communication in this moth evolved through a sensory exploitation process.     

Listening for bats: the hearing range of the bushcricket Phaneroptera falcata for bat echolocation calls measured in the field

The hearing range of the tettigoniid Phaneropterafalcata for the echolocation calls of freely flying mouseeared bats (Myotis myotis) was determined in the field. The hearing of the insect was monitored using hook electrode recordings from an auditory interneuron, which is as sensitive as the hearing organ for frequencies above 16 kHz. The flight path of the bat relative to the insect's position was tracked by recording the echolocation calls with two microphone arrays, and calculating the bat's position from the arrival time differences of the calls at each microphone. The hearing distances ranged from 13 to 30 m. The large variability appeared both between different insects and between different bat approaches to an individual insect. The escape time of the bushcricket, calculated from the detection distance of the insect and the instantaneous flight speed of the bat, ranged from 1.5 to more than 4s. The hearing ranges of bushcrickets suggest that the insect hears the approaching bat long before the bat can detect an echo from the flying insect.     

The communicative potential of bat echolocation pulses

Ecological constraints often shape the echolocation pulses emitted by bat species. Consequently some (but not all) bats emit species-specific echolocation pulses. Because echolocation pulses are often intense and emitted at high rates, they are potential targets for eavesdropping by other bats. Echolocation pulses can also vary within species according to sex, body size, age, social group and geographic location. Whether these features can be recognised by other bats can only be determined reliably by playback experiments, which have shown that echolocation pulses do provide sufficient information for the identification of sex and individual in one species. Playbacks also show that bats can locate conspecifics and heterospecifics at foraging and roost sites by eavesdropping on echolocation pulses. Guilds of echolocating bat species often partition their use of pulse frequencies. Ecology, allometric scaling and phylogeny play roles here, but are not sufficient to explain this partitioning. Evidence is accumulating to support the hypothesis that frequency partitioning evolved to facilitate intraspecific communication. Acoustic character displacement occurs in at least one instance. Future research can relate genetic population structure to regional variation in echolocation pulse features and elucidate those acoustic features that most contribute to discrimination of individuals.