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.     

日本伏翼的翼型、回声定位信号及出飞时间特征

对日本伏翼的翼型、回声定位信号及晚间出飞时间进行研究。结果表明,日本伏翼的翼型具有高翼载、低翼展比和中等偏高的翼型特征。日本伏翼发出具有1 - 2 个谐波结构的调频型(FM)回声定位信号叫声,其叫声时程、主频率的平均值分别为3.26 ms 和56. 27 kHz,所有叫声特征参数,个体间变异系数CVb 比个体内变 异系数CVw 大。日本伏翼的晚间出飞时间具有明显月变化,与当地日落时间、气温呈现显著相关。通过与文献比较,发现日本伏翼的回声定位信号特征与录音状态、飞行生境有关;此外,晚间出飞时间存在一定的地理差异。本研究结果将为蝙蝠回声定位信号特征的种属特异性及其生境选择的进一步研究提供有用的信息。     

Breaking the trade-off: rainforest bats maximize bandwidth and repetition rate of echolocation calls as they approach prey

Both mammals and birds experience a performance trade-off between producing vocalizations with high bandwidths and at high repetition rate. Echolocating bats drastically increase repetition rate from 2–20 calls s⁻¹ up to about 170 calls s⁻¹ prior to intercepting airborne prey in order to accurately track prey movement. In turn, bandwidth drops to about 10–30 kHz for the calls of this ‘final buzz’. We have now discovered that Southeast Asian rainforest bats (in the vespertilionid subfamilies Kerivoulinae and Murininae) are able to maintain high call bandwidths at very high repetition rates throughout approach to prey. Five species of Kerivoula and Phoniscus produced call bandwidths of between 78 and 170 kHz at repetition rates of 140–200 calls s⁻¹ and two of Murina at 80 calls s⁻¹. The ‘typical’ and distinct drop in call frequency was present in none of the seven species. This stands in striking contrast to our present view of echolocation during approach to prey in insectivorous bats, which was established largely based on European and American members of the same bat family, the Vespertilionidae. Buzz calls of Kerivoula pellucida had mean bandwidths of 170 kHz and attained maximum starting frequencies of 250 kHz which makes them the most broadband and most highly pitched tonal animal vocalization known to date. We suggest that the extreme vocal performance of the Kerivoulinae and Murininae evolved as an adaptation to echolocating and tracking arthropods in the dense rainforest understorey.     

Echolocation call description of 15 species of Middle-Eastern desert dwelling insectivorous bats

Modern advances in acoustic technology have made possible new and broad ranges of research in bioacoustics, particularly with regard to echolocating bats. In the present study, we present an acoustic guide to the calls of 15 species of bats in the Arava rift valley, Israel, with a focus on their bioacoustics, habitat use and explaining differences between similar species. We also describe a potential case of frequency separation where four bat species using six call types appear to separate the frequencies of their calls to minimize overlap. The studied community of bat species is also found in other Middle Eastern deserts including the deserts of Jordan, Syria and Saudi Arabia and we hope that data gathered will benefit other bat researchers in the region.     

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.     

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。     

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]。     

Female greater wax moths reduce sexual display behavior in relation to the potential risk of predation by echolocating bats

Female greater wax moths Galleria mellonella display by wing fanning in response to bursts of ultrasonic calls produced bymales. The temporal and spectral characteristics of these callsshow some similarities with the echolocation calls of batsthat emit frequency-modulated (FM) signals. Female G. mellonellatherefore need to distinguish between the attractive signalsof male conspecifics, which may lead to mating opportunities,and similar sounds made by predatory bats. We therefore predictedthat (1) females would display in response to playbacks of male calls; (2) females would not display in response to playbacksof the calls of echolocating bats (we used the calls of Daubenton'sbat Myotis daubentonii as representative of a typical FM echolocatingbat); and (3) when presented with male calls and bat callsduring the same time block, females would display more whenperceived predation risk was lower. We manipulated predationrisk in two ways. First, we varied the intensity of bat callsto represent a nearby (high risk) or distant (low risk) bat.Second, we played back calls of bats searching for prey (lowrisk) and attacking prey (high risk). All predictions weresupported, suggesting that female G. mellonella are able todistinguish conspecific male mating calls from bat calls, andthat they modify display rate in relation to predation risk.The mechanism (s) by which the moths separate the calls ofbat and moth must involve temporal cues. Bat and moth signalsdiffer considerably in duration, and differences in durationcould be encoded by the moth's nervous system and used in discrimination.     

Phenotypic traits evolution and morphological traits associated with echolocation calls in cryptic horseshoe bats (Rhinolophidae)

Bats provide an excellent casestudy for studying evolution due to their remarkable flight and echolocation capabilities. In this study, we sought to understand the phenotypic evolution of key traits in Rhinolophidae (horseshoe bats) using phylogenetic comparative methods. We aimed to test the phylogenetic signals of traits, and evaluated the best-fit evolutionary models given the data for each trait considering different traits may evolve under different models (i.e., Brownian Motion [BM], Ornstein-Uhlenbeck [OU], and Early Burst [EB]) and reconstruct ancestral character states. We examined how phenotypic characters are associated with echolocation calls and minimum detectable prey size. We measured 34 traits of 10 Asian rhinolophids species (187 individuals). We found that the majority of traits showed a high phylogenetic signal based on Blomberg′s K and Pagel′s λ, but each trait may evolve under different evolutionary models. Sella traits were shown to evolve under stabilizing selection based on OU models, indicating sella traits have the tendency to move forward along the branches toward some medial value in equilibrium. Our findings highlight the importance of sella characters in association with echolocation call emissions in Rhinolophidae, as calls are important for spatial cognition and also influence dietary preferences. Minimum detectable prey size in Rhinolophidae was associated with call frequency, bandwidth, call duration, wingspan, and wing surface area. Ultimately, understanding trait evolution requires sensitivity due to the differential selective pressures which may apply to different characteristics.     

The evolution of flight and echolocation in bats: another leap in the dark

The earliest known complete bats, from the Eocene (49–53 Mya), were already capable of flapping flight and echolocation. In the absence of direct fossil evidence there have been many speculative scenarios advanced to explain the evolution of these behaviours and their distributions in extant bats. Theories assuming chiropteran monophyly have generally presumed the ancestral pre‐bat was nocturnal, arboreal and insectivorous. Following this assumption hypotheses can be divided into the echolocation first, flight first and tandem development hypotheses, all of which assume that flight evolved only once in the lineage. In contrast, the chiropteran diphyly hypothesis suggests that flight evolved twice. Evidence supporting and refuting the different hypotheses are reviewed. It is concluded that there are significant problems attached to all the current models. A novel hypothesis is advanced, which starts from the assumption that bats are monophyletic and the ancestral pre‐bat was arboreal, but diurnal and frugivorous. After the evolution of flight it is suggested that these animals were driven into the nocturnal niche by the evolution of raptorial birds, and different groups evolved either specialised nocturnal vision (megachiropterans) or echolocation (microchiropterans). A block on sensory modality transfer has retained this distribution of perceptual capabilities ever since, despite some Megachiroptera evolving rudimentary echolocation, and the dietary convergence of some Microchiroptera with the Megachiroptera. The new hypothesis overcomes many of the problems identified in previous treatments.     

Analysis of the distribution of insectivorous bats in Israel

Data on localities from which insectivorous bats were collected or sighted in Israel were compiled into a Geographical Information System (GIS) in order to analyse patterns of species distribution. By intersecting precipitation and temperature data with spatially-referenced data on species observations stored in the GIS we determined the 'climatic envelope' of each species and constructed predictive maps which show the potential distribution of each species. Using cluster analysis, the bats were classified into three main biogeographic groups according to their distribution, namely desert, Mediterranean, and widespread in Israel. The potential distribution maps of all the species indicate that there are areas which have suitable bat habitats but from which bats have never been collected or observed. In the Mediterranean region of Israel, this is attributed to a large reduction in population size due to fumigation of caves, cave visitation and secondary poisoning.     

Energetic cost of hovering flight in nectar-feeding bats (Phyllostomidae: Glossophaginae) and its scaling in moths, birds and bats

Three groups of specialist nectar-feeders covering a continuous size range from insects, birds and bats have evolved the ability for hovering flight. Among birds and bats these groups generally comprise small species, suggesting a relationship between hovering ability and size. In this study we established the scaling relationship of hovering power with body mass for nectar-feeding glossophagine bats (Phyllostomidae). Employing both standard and fast-response respirometry, we determined rates of gas exchange in Hylonycteris underwoodi (7 g) and Choeronycteris mexicana (13–18 g) during hover-feeding flights at an artificial flower that served as a respirometric mask to estimate metabolic power input. The O₂ uptake rate (V˙ _o2) in ml g⁻¹ h⁻¹ (and derived power input) was 27.3 (1.12 W or 160 W kg⁻¹) in 7-g Hylonycteris and 27.3 (2.63 W or 160 W kg⁻¹) in 16.5-g Choeronycteris and thus consistent with measurements in 11.9-g Glossophagasoricina (158 W kg⁻¹, Winter 1998). V˙ _o2 at the onset of hovering was also used to estimate power during forward flight, because after a transition from level forward to hovering flight gas exchange rates initially still reflect forward flight rates. V˙ _o2 during short hovering events (<1.5 s) was 19.0 ml g⁻¹ h⁻¹ (1.8 W) in 16-g Choeronycteris, which was not significantly different from a previous, indirect estimate of the cost of level forward flight (2.1 W, Winter and von Helversen 1998). Our estimates suggest that power input during hovering flight P _h (W) increased with body mass M (kg) within 13–18-g Choeronycteris (n = 4) as P _h  = 3544 (±2057 SE) M ^{1.76 (±0.21 SE)} and between different glossophagine bat species (n = 3) as P _h  = 128 (±2.4 SE) M ^{0.95 (±0.034 SE)}. The slopes of three scaling functions for flight power (hovering, level forward flight at intermediate speed and submaximal flight power) indicate that: 1. The relationship between flight power to flight speed may change with body mass in the 6–30-g bats from a J- towards a U-shaped curve. 2. A metabolic constraint (hovering flight power equal maximal flight power) may influence the upper size limit of 30–35 g for this group of flower specialists. Mass-specific power input (W kg⁻¹) during hovering flight appeared constant with regard to body size (for the mass ranges considered), but differed significantly (P < 0.001) between groups. Group means were 393 W kg⁻¹ (sphingid moths), 261 W kg⁻¹ (hummingbirds) and 159 W kg⁻¹ (glossophagine bats). Thus, glossophagine bats expend the least metabolic power per unit of body mass supported during hovering flight. At a metabolic power input of 1.1 W a glossophagine bat can generate the lift forces necessary for balancing 7 g against gravitation, whereas a hummingbird can support 4 g and a sphingid moth only 3 g of body mass with the same amount of metabolic energy. These differences in power input were not fully explained by differences in induced power output estimated from Rankine-Froude momentum-jet theory. Accepted: 10 November 1998     

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。     

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。