七种蝙蝠回声定位行为生态研究
摘要
蝙蝠回声定位的研究一直是国际上蝙蝠研究的重点和热点。由于声波的特征和蝙蝠的生态行为有着密切的联系，如声波的主频率影响蝙蝠可探测到的猎物的大小、猎物能被探测到的距离及猎物探测和躲避蝙蝠的能力，声波的结构则影响蝙蝠在干扰环境中探测猎物的能力，因此研究影响蝙蝠声波结构和特征的因素、探讨蝙蝠声波探测和定位的机制，可以更加深入地了解蝙蝠的生态行为。回声定位声波的发出和分析处理不仅受声波发射器官及声波接收器官结构的限制，也受蝙蝠的体型、年龄、蝙蝠的状态、蝙蝠捕食阶段等因素的影响。
本论文采用回声定位声波探测仪(D980，ULTRASOUND DETECTOR，Pettersson Elektronik AB，瑞典)和数字式录音机(SONY，TCD-D8)录制回声定位声波，用专业声谱分析软件Batsound 3．10(Pettersson Elektronik AB，瑞典)对录制的声波进行分析。用游标卡尺、直尺和天平等测量蝙蝠的形态特征。采用专业统计软件SPSS11．5进行数据处理和统计分析。
对云南菊头蝠、大耳菊头蝠、普氏蹄蝠、双色蹄蝠的回声定位声波进行分析，描述其声波特征，是国内首次对这4种蝙蝠回声定位声波的报导，云南菊头蝠是中国特有蝙蝠，对其进行声波描述具有中国特色，为国际蝙蝠回声定位声波库的建立提供新的数据和资料。对云南菊头蝠的母幼蝠进行形态和声波上的比较研究，探讨随着云南菊头蝠生长发育其形态和回声定位声波的变化，描述母幼的交流行为，确定其母幼交流的方式主要为声音。比较了云南菊头蝠和普氏蹄蝠飞行状态和悬挂状态下的回声定位声波，描述这2种蝙蝠的多普勒补偿效应，悬挂状态声波频率显著高于飞行状态，声脉冲时间和间隔时间显著长于飞行状态，FM带宽窄于飞行状态。比较和分析3种蝙蝠(角菊
头幅、三叶蹄幅和大蹄幅）形态和回声定位声波的性别二态性，结果显
示编幅形态没有显著的性别二态性，回声定位声波特征存在明显的性
别二态性，可能由于群居骗幅的性别分工或为通讯方便造成的，编幅
形态和回声定位声波性别二态性的研究在国际上开展不多，本文对3
种崛幅的性别二态性的比较填补了国内的空白，厂辟了中国编幅研究
的新领域。
The echolocation of bats is a major field in the research of bats. The characteristics of calls are closely correlated to the ecology of bats, for example, the dominant frequency influences the body size of bats' prey, the detecting range of bats; the structure of calls decides bats' ability to detect prey in the complicated environment. Therefore, the research on factors, which influence the structure and characteristics of ultrasound and on the mechanism by which bats detect and orient the prey will promote the understanding of the ecological behavior of bats. The emission and receiving of calls are not only influenced by the sound-emitting apparatus and sound-receiving apparatus, but also by the body size, age, state and foraging stages of bats.
The echolocation calls detector (D980, ULTRASOUND DETECTOR, Pettersson Elektronik AB, Sweden) and digital recorder (SONY, TCD-D8) were used to record the calls and the software Batsound 3.10 (Pettersson Elektronik AB, Sweden) was used to analyze the calls. The morphological features were measured with the ruler, vernier caliper and balance. The software SPSS 11.5 was adopted to compute and analyze the data.
The characteristics of echolocating calls of R.yunanensis, R.macrotis, H.pratti and H.bicolor were analyzed and described; it is the first time in China to do this kind of research. R.yunanensis is native to China, so the research on its echolocation is characteristic of China and provides new data to set up the echolocating calls database in the world. The bat mother and its child were compared concerning their morphology and echolocating calls, and the change in morphology and echolocating calls, which happened as the young bat grew were discussed. The communicating behavior between the mother and the young was described and thus the calls played an important role in the communication between
the mother and the young. The echolocating calls in the state of fly were compared with those in the state of hang in two kinds of bats, R.yunanensis and H.pratti,and the Doppler Compensation Effect was described. The analyzing result indicated that the frequency in hang was higher that that in fly, the duration and interpulse duration in hang were longer than those in fly, and the FM bandwidth in hang was narrower than that in fly. The difference in morphology and echolocating calls of female and male individuals was analyzed in three kinds of bats, R.cornutus, A.wheeleeri and H.armiger. The result showed that there was no significant difference in morphology, but there was significant difference in echolocating calls between the two sexes in all the three kinds of bats. The significant difference in calls may be caused by the work division or communication in the colony of bats, as bats are animals which are adapted to aggregation life.
本论文采用回声定位声波探测仪(D980，ULTRASOUND DETECTOR，Pettersson Elektronik AB，瑞典)和数字式录音机(SONY，TCD-D8)录制回声定位声波，用专业声谱分析软件Batsound 3．10(Pettersson Elektronik AB，瑞典)对录制的声波进行分析。用游标卡尺、直尺和天平等测量蝙蝠的形态特征。采用专业统计软件SPSS11．5进行数据处理和统计分析。
对云南菊头蝠、大耳菊头蝠、普氏蹄蝠、双色蹄蝠的回声定位声波进行分析，描述其声波特征，是国内首次对这4种蝙蝠回声定位声波的报导，云南菊头蝠是中国特有蝙蝠，对其进行声波描述具有中国特色，为国际蝙蝠回声定位声波库的建立提供新的数据和资料。对云南菊头蝠的母幼蝠进行形态和声波上的比较研究，探讨随着云南菊头蝠生长发育其形态和回声定位声波的变化，描述母幼的交流行为，确定其母幼交流的方式主要为声音。比较了云南菊头蝠和普氏蹄蝠飞行状态和悬挂状态下的回声定位声波，描述这2种蝙蝠的多普勒补偿效应，悬挂状态声波频率显著高于飞行状态，声脉冲时间和间隔时间显著长于飞行状态，FM带宽窄于飞行状态。比较和分析3种蝙蝠(角菊
头幅、三叶蹄幅和大蹄幅）形态和回声定位声波的性别二态性，结果显
示编幅形态没有显著的性别二态性，回声定位声波特征存在明显的性
别二态性，可能由于群居骗幅的性别分工或为通讯方便造成的，编幅
形态和回声定位声波性别二态性的研究在国际上开展不多，本文对3
种崛幅的性别二态性的比较填补了国内的空白，厂辟了中国编幅研究
的新领域。
The echolocation of bats is a major field in the research of bats. The characteristics of calls are closely correlated to the ecology of bats, for example, the dominant frequency influences the body size of bats' prey, the detecting range of bats; the structure of calls decides bats' ability to detect prey in the complicated environment. Therefore, the research on factors, which influence the structure and characteristics of ultrasound and on the mechanism by which bats detect and orient the prey will promote the understanding of the ecological behavior of bats. The emission and receiving of calls are not only influenced by the sound-emitting apparatus and sound-receiving apparatus, but also by the body size, age, state and foraging stages of bats.
The echolocation calls detector (D980, ULTRASOUND DETECTOR, Pettersson Elektronik AB, Sweden) and digital recorder (SONY, TCD-D8) were used to record the calls and the software Batsound 3.10 (Pettersson Elektronik AB, Sweden) was used to analyze the calls. The morphological features were measured with the ruler, vernier caliper and balance. The software SPSS 11.5 was adopted to compute and analyze the data.
The characteristics of echolocating calls of R.yunanensis, R.macrotis, H.pratti and H.bicolor were analyzed and described; it is the first time in China to do this kind of research. R.yunanensis is native to China, so the research on its echolocation is characteristic of China and provides new data to set up the echolocating calls database in the world. The bat mother and its child were compared concerning their morphology and echolocating calls, and the change in morphology and echolocating calls, which happened as the young bat grew were discussed. The communicating behavior between the mother and the young was described and thus the calls played an important role in the communication between
the mother and the young. The echolocating calls in the state of fly were compared with those in the state of hang in two kinds of bats, R.yunanensis and H.pratti,and the Doppler Compensation Effect was described. The analyzing result indicated that the frequency in hang was higher that that in fly, the duration and interpulse duration in hang were longer than those in fly, and the FM bandwidth in hang was narrower than that in fly. The difference in morphology and echolocating calls of female and male individuals was analyzed in three kinds of bats, R.cornutus, A.wheeleeri and H.armiger. The result showed that there was no significant difference in morphology, but there was significant difference in echolocating calls between the two sexes in all the three kinds of bats. The significant difference in calls may be caused by the work division or communication in the colony of bats, as bats are animals which are adapted to aggregation life.
引文
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[2] Arita H and M B Fenton. Flight and echolocation in the ecology and evolution of bats. Tree, 1997. 12:53-58.
[3] Bell G P and M B Fenton. The use of Doppler-shifted echoes as a flutter detection and clutter rejection system: the echolocation and feeding behavior of Hipposideros ruber (Chiroptera: Hipposideridae). Behav Ecol Sociobiol, 1984. 15:109-114.
[4] Bogdanowicz W, R D Csada and M B Fenton. Structure of nose leaf, echolocation, and foraging behavior in the Phyllostomidae (Chiroptera). Journal of Mammalogy, 1997. 78(3):942-953.
[5] Bringham R M, J E Cebek and M B C Hichey. Intraspecific variation and resource partitioning in insectivorous bats. Journal of Mammalogy, 1989. 70:426-28.
[6] Emde G von der and H-U Schnitzler. Fulttering target detection in Hipposiderid bats. J Comp Physiol A, 1986. 159:765-772.
[7] Fenton M B and I L Rautenbach. A comparison of the roosting and foraging behaviour of three species of African insectivorous bats (Rhinolophidae, Vespertilionidae, and Molossidae). Can J Zool, 1986. 64:2860-2867.
[8] Fenton M B. Hipposideros caffer (Chiroptera: Hipposideridae) in Zimbabwe: morphology and echolocation calls. J. Zool, Lond. 1986. A 210:347-353.
[9] Francis C M and J Habersetzer. Interspecific and
Intraspecific Variation in Echolocation Call Frequency and morphology of horseshoe bats, Rhinolophus and Hipposederos. Bat: Biology and Conservation. Smithsonian Institution Press. Washington and London. 1998. 169-179.
[10] Grinnell A D. Hearing in Bats: An Overview. Hearing by Bats. Springer-Verlag, 1995. 1-36.
[11] Guillén A, J Juste B and C Ibánez. Variation in the frequency of the echolocation calls of Hipposideros ruber in the Gulf of Guinea: an exploration of the adaptive meaning of the constant frequency value in rhinolophoid CF bats. J Evol Biol, 2000. 13:70-80.
[12] Guppy A and R B Coles. Acoustical aspects of hearing and echolocation in bats. Animal Sonar: Processes and Performance. Plenum Press, New York. 1988. 289-294.
[13] Habersetzer J. Adaptive echolocation sounds in the bat Rhinopoma hardiwchei. J Comp Physiol, 1981. 144:559-566.
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