大菊头蝠和中菊头蝠分娩声波的特征

动物的声学信号在信息传递和维持社群稳定等方面扮演了非常重要的角色,特别是对于占据夜空生态位并集群生活的蝙蝠,它们日间栖息于几乎没有光照的洞穴中。这些夜行性蝙蝠主要借助声信号介导重要生活史事件,包括生殖、觅食、通讯和空间导航等。分娩是动物生殖活动中的关键一环,决定了后代甚至母体的生死。一直以来野生动物的分娩行为,特别是分娩叫声鲜有研究。本研究以大菊头蝠(Rhinolophus luctus)和中菊头蝠(R. affinis)为研究对象,捕捉并录制其在分娩过程中的声信号,进而开展声谱分析和统计分类。研究发现,大菊头蝠和中菊头蝠在分娩过程中分别发出了6种和4种类型的交流声波。基于子集验证的判别式分析表明,大多数的音节类型都被正确划分。纳入多种声学参数的多维尺度分析显示,不同音节类型在三维图像中差异显著。此外,欧几里得距离分析发现,种间的叫声差异远大于种内的叫声差异。本研究结果可为深入揭示夜行哺乳动物的叫声功能和发声机制提供参考。     

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。     

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.     

四川产中菊头蝠喜马拉雅亚种和马铁菊头蝠日本亚种外部形态及头骨的比较

对四川产中菊头蝠喜马拉雅亚种(Rhinolophus affinis himalayanus)和马铁菊头蝠日本亚种(R.ferrumequinum nippon)的23项外部形态指标和22项头骨形态指标进行了测量,并进行数理统计分析。结果表明,(1)中菊头蝠喜马拉雅亚种和马铁菊头蝠日本亚种的23项外部形态测量指标中有2项差异显著(P<0.05),16项差异极显著(P<0.01),其中:中菊头蝠喜马拉雅亚种(n=14)体长小于59 mm(47.30~58.40),尾长小于26 mm(21.69~25.68),耳长小于22 mm(18.36~21.88),前臂长小于56 mm(50.9~54.20),第Ⅲ掌骨第一指节长小于16 mm(13.2~15.53),第Ⅲ掌骨第二指节小于32 mm(27.38~31.74);而马铁菊头蝠日本亚种(n=6)体长大于62 mm(62.43~66.76),尾长大于30 mm(30.56~36.18),耳长大于24 mm(24.47~27.20),前臂长大于56 mm(58.78~63.46),第Ⅲ掌骨第一指节大于19 mm(19.38~21.39),第Ⅲ掌骨第二指节大于34 mm(34.42~38.11),这些差异可作为区分四川产这两个种的依据。(2)中菊头蝠喜马拉雅亚种和马铁菊头蝠日本亚种的22项头骨形态测量指标中,除颅高、听泡间距差异不显著(P>0.05)外,其他20项指标都存在极显著差异(P<0.01)。     

中菊头蝠中国三亚种的形态特征比较

比较了四川、广东和海南3地的中菊头蝠(Rhinolophus affinis)种群的外部和头骨形态差异。在测量23个外部可量性状以及22个头骨可量性状的基础上,运用统计分析软件SPSS12.0对其中12个外部可量性状以及22个头骨可量性状进行数理统计。主成分分析的结果表明四川、广东和海南3地的中菊头蝠在外部形态和头骨形态上均有明显差异,用判别分析对主成分分析结果的正确性进行验证,证明了中国的中菊头蝠划为喜马拉雅、华南、海南3个亚种是合理的。     

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。     

贵州五种菊头蝠的核型分析 *

采用常规骨髓细胞空气干燥法,研究了贵州5种菊头蝠的核型。贵州菊头蝠和中菊头蝠2n=62,两者染色体臂数(NF)均为60;托氏菊头蝠、小菊头蝠和栗黄菊头蝠的染色体数是2n=36,其中托氏菊头蝠和小菊头蝠染色体臂数(NF)是58,栗黄菊头蝠是60。5种菊头蝠的性别决定机制均是xY。     

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。     

四种菊头蝠染色体组型分析

本文分析了皮氏菊头蝠(R. pearsoni chinensis),鲁氏菊头蝠(R.rouxi sinicus),角菊头蝠(R.cornutus pumilus)及中菊头蝠(R.affinis)的常规核型,现报道如下。     

中国大陆菊头蝠科一新纪录——单角菊头蝠(Rhinolophus monoceros Andersen,1905)

During a survey of bat species diversity from July 2008 to February 2009 in Songtao county of Guizhou Province, one specimen of Rhinolophus was collected from Guizhou. It was identified as a Formosan lesser horseshoe bat, based on the following features: small body size, forearm length of 36. 3mm, the greatest length of skull was 15.16 mm, the cells of lancet were shorter and appeared triangular and showed an acute angle, and forward bending, the lancet was shorter and approximately triangular and had a narrow top, the anterior median swellings of the skull were higher and more vertical, the rear of the sagittal crest was thin, the upper molars were shorter and the length of C~1 - M~3 was 5. 36 mm, the width of M~3 M~3 was 5.24 mm. Rhinolophus monoceros, which is new record in Guizhou and the first found on the Chinese mainland, supports Simmons' prediction that Rhinolophus monoceros may occur in the south of China.     

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.     

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。     

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.     

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.     

The effects of cultural drift on geographic variation in echolocation calls of the Chinese rufous horseshoe bat (Rhinolophus sinicus)

Drift, selection, or their combined effects commonly drive geographic variation in traits. Clarifying the relative roles of each process is a long‐standing research goal in evolutionary biology. Acoustic signals of bats are a phenotypic characteristic that plays an important role in social organization and species recognition. We extensively sampled the Chinese rufous horseshoe bat (Rhinolophus sinicus) throughout China and Vietnam and reconstructed a species phylogeny to better understand the patterns and causes of the geographic variation of acoustic signals. Our results showed that the resting frequency (RF) of calls varied with latitude, sex, and distance among different colony locations. RF differences were not correlated with genetic distance (based on only one nuclear locus and the mitochondrial locus), climatic factors (mean annual temperature and mean annual precipitable water), or body size, although differences in calls increased with distance among various populations. This suggests that cultural drift may play more important roles than genetic drift and acoustic adaptation in shaping acoustic differences within regions in R. sinicus.     

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。     

Origins of Horseshoe Bats (Rhinolophus, Rhinolophidae) in Southern Africa: Evidence from Allozyme Variability

Morphological analyses indicate that horsehose bats in the genus Rhinolophus constitute a monophyletic group which most likely originated in southeastern Asia but which presently inhabits Oriental, Australian, Palaearctic, and Ethiopian zoogeographical provinces. Ten species occur in southern Africa, but it is uncertain which species represent dispersals from Eurasia through North Africa and which have resulted from speciation in Africa. Analyses of 34 allozyme encoding loci in these 10 species and in 2 southern African species of leafnose bats in the sister genus Hipposideros reveal the presence of at least three lineages of Rhinolophus in southern Africa. One lineage includes R. clivosus, R. darlingi, R. fumigatus, and R. hildebrandtii, all of which, except R. clivosus, are endemic to sub-Saharan Africa. Rhinolophus blasii is genetically allied with, but distinct from this group, and appears to be a recent migrant from another lineage centered on the Mediterranean. A third lineage, including at least R. capensis, R. denti, R. simulator, and R. swinnyi, is endemic to sub-Saharan Africa. The phylogenetic position of R. landeri is uncertain, most likely because of the small sample size used to estimate allelic frequencies for this species. The biochemical genetic definitions of these lineages largely agree with previous morphological analyses of Rhinolophus species. Divergences between species within two lineages (R. clivosus, R. darlingi, R. fumigatus, and R. hildebrandtii; and R. capensis, R. denti, R. simulator, and R. swinnyi) appear to reflect two bursts of speciation in the Plio-Pleistocene period within Africa.