洞栖蝙蝠回声定位声波及生态位分化研究
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摘要
蝙蝠的回声定位系统和飞行是区别于其他哺乳动物的两大特征。蝙蝠利用回声定位声波进行空中飞行定位和探测猎物，声波特征不同，对周围环境及猎物的定位和识别能力也不同。飞行特点与形态特征（翼形）的关系密切。翼形不同，蝙蝠飞行的速度、灵活性不同，其最适捕食生境及猎物大小就不同。在同一生境中栖息的蝙蝠种群，为了避免激烈的种间竞争，可能在回声定位声波、形态、食性和捕食等方面具有一定的生态位分化(niche separation)现象。因此，本文分析了集安市大砬子洞水鼠耳蝠（Myotis daubentonii），绯鼠耳蝠（M. formosus），白腹管鼻蝠（Murina leucogaster），伊氏鼠耳蝠（M. ikonnikovi），马铁菊头蝠（Rhinolophus ferrumequinum）不同状态（飞行、静止、悬挂、爬行）、不同性别之间的声波差异，对其食性、形态特征、捕食生境、捕食策略、活动规律等进行对比研究。最后提出这5种蝙蝠的生态位分化显著，并对蝙蝠种群稳定共存的机制进行了讨论。
马铁菊头蝠（Rhinolophus ferrumequinum）是大中型蝙蝠，使用CF声波，主要以大中型的鳞翅目、鞘翅目昆虫为食，在开阔空间和稍密集的林内及灌木丛表面进行空中捕食(aerial feeding)或拾遗捕食(gleaning)。其他4种蝙蝠均利用FM声波。其中，白腹管鼻蝠（Murina leucogaster）和绯鼠耳蝠（M. formosus）均为中小型蝙蝠，前者声波频带极宽，以中小型的鳞翅目、双翅目昆虫为食，在密集的林内进行空中捕食；后者声波带宽中等，以中小型的鞘翅目、鳞翅目昆虫为食，在中等复杂的林中进行空中或拾遗捕食。水鼠耳蝠（Myotis daubentonii）和伊氏鼠耳蝠（M. ikonnikovi）为小型蝙蝠，前者声波带宽中等，主要以水生小型的双翅目、毛翅目为食，在水面5—20cm和中等复杂的林中进行拖网(trawling)和空中捕食，后者声波带宽最窄，主要以小型的鳞翅目、双翅目昆虫为食，在开阔空间内或稀疏林中进行空中捕食，视觉可能在捕食中具有重要的辅助作用。此外，5种蝙蝠在山洞内的栖息位置也不同，种群之间距离较大。总之，这些种间差异保证了5种
蝙蝠能够在同一山洞中稳定共存，降低种间竞争。
同时，水鼠耳蝠、白腹管鼻蝠、伊氏鼠耳蝠、马铁菊头蝠不同状态下回声定位声波的种内差异显著。水鼠耳蝠和伊氏鼠耳蝠不同性别之间声波和形态的差异显著，存在典型的性别二态性现象。马铁菊头蝠的性别差异不显著。声波和形态的种内差异可能是避免种内竞争和声波干扰的手段，使得营集群生活的蝙蝠能够达到稳定的种内共存。飞行状态下5种蝙蝠种间声波的差异则预示着蝙蝠可能偏好不同的捕食策略、捕食生境、猎物大小和类型等。
Echolocation system and flight are two characters which distinguish bats from other mammalians. Bats use echolocation calls to orient and detect targets. Bat’s ability to locate enviroments and discriminate targets changes along with echolocation calls. Flight characters associate with morphologic features. With different wing shape, the velocity and maneuvorbility change evidently, and so do suitable foraging habitat and targets type. Bats those inhabit in the same habitat will present ecopartition in echolocation, morphology, diets and so on in order to avoid interspecies competition. Therefore, in the thesis, first descripe morphologic characters and analyse echolocation calls of five bat species, Myotis daubentonii, M. formosus, Murina leucogaster, M. ikonnikovi, Rhinolophus ferrumequinum in different of sex and states which includes flight, stationary, hanging and crawling; sencondly, study the difference of echolocation calls during flight, diets, morhpologic characters, foraging habitat, foraging strategy and action rules of the five bat species by contrast. At last, conclude that that niche separation of the bats are evident, what make bats coexist stably.
Rhinolophus ferrumequinum is biggest in these 5 species(mess is about 20g), use CF echolocation calls, which is sensitive to wing-vibrated insects. It feeds mainly on big-medium insects of order Lepidoptera and Coleoptera, foraging in wide space and sparse-foliage forest by aerial feeding and gleaning. Other four species all use FM echolocation calls. Murina leucogaster and M. formosus are nearly the same in body size, the former’s wave width is wondrously big. It feeds mainly on medium-small insects of order Lepidoptera and Diptera, foraging in complicated forest by aerial feeding. The latter's wave width is medium. It feeds mainly on medium-small insects of order Coleoptera and Lepidoptera, foraging in medium-complicated forest by aerial feeding or gleaning. M. daubentonii
and M. ikonnikovi are the smallest ones in size. The former’ wave width is medium. It feeds mainly on aquatic small insects of order Diptera and Trichoptera, foraging in 5-20cm above water surface and midium- complicated forest by trawling and aerial feeding. The latter’s wave width is shortest, which have disadvantage to discriminate targets from enviroment. It feeds mainly on even smaller insects of order Lepidoptera and Diptera, foraging in wide space or sparse-foliage forest by aerial feeding. Vision is an important assistant means during foraging. These interspecies differences decrease interspecies competition, and ensure they can coexist in one cave steadily.
At the same time, the result show that there is significant difference of echolocation calls in different states in all species, and only in M. daubentonii and M. ikonnikovi that there is significant difference in body size and echolocation calls betweed two sexes, but not in R. Ferrumequinum. The difference in calls and morphology is possibly the means of escaping interspecies sound jamming and ardent competition. While the difference of sound during flight indicate that bats may prefer to different foraging styles, foraging habitat, targets size and so on.
马铁菊头蝠（Rhinolophus ferrumequinum）是大中型蝙蝠，使用CF声波，主要以大中型的鳞翅目、鞘翅目昆虫为食，在开阔空间和稍密集的林内及灌木丛表面进行空中捕食(aerial feeding)或拾遗捕食(gleaning)。其他4种蝙蝠均利用FM声波。其中，白腹管鼻蝠（Murina leucogaster）和绯鼠耳蝠（M. formosus）均为中小型蝙蝠，前者声波频带极宽，以中小型的鳞翅目、双翅目昆虫为食，在密集的林内进行空中捕食；后者声波带宽中等，以中小型的鞘翅目、鳞翅目昆虫为食，在中等复杂的林中进行空中或拾遗捕食。水鼠耳蝠（Myotis daubentonii）和伊氏鼠耳蝠（M. ikonnikovi）为小型蝙蝠，前者声波带宽中等，主要以水生小型的双翅目、毛翅目为食，在水面5—20cm和中等复杂的林中进行拖网(trawling)和空中捕食，后者声波带宽最窄，主要以小型的鳞翅目、双翅目昆虫为食，在开阔空间内或稀疏林中进行空中捕食，视觉可能在捕食中具有重要的辅助作用。此外，5种蝙蝠在山洞内的栖息位置也不同，种群之间距离较大。总之，这些种间差异保证了5种
蝙蝠能够在同一山洞中稳定共存，降低种间竞争。
同时，水鼠耳蝠、白腹管鼻蝠、伊氏鼠耳蝠、马铁菊头蝠不同状态下回声定位声波的种内差异显著。水鼠耳蝠和伊氏鼠耳蝠不同性别之间声波和形态的差异显著，存在典型的性别二态性现象。马铁菊头蝠的性别差异不显著。声波和形态的种内差异可能是避免种内竞争和声波干扰的手段，使得营集群生活的蝙蝠能够达到稳定的种内共存。飞行状态下5种蝙蝠种间声波的差异则预示着蝙蝠可能偏好不同的捕食策略、捕食生境、猎物大小和类型等。
Echolocation system and flight are two characters which distinguish bats from other mammalians. Bats use echolocation calls to orient and detect targets. Bat’s ability to locate enviroments and discriminate targets changes along with echolocation calls. Flight characters associate with morphologic features. With different wing shape, the velocity and maneuvorbility change evidently, and so do suitable foraging habitat and targets type. Bats those inhabit in the same habitat will present ecopartition in echolocation, morphology, diets and so on in order to avoid interspecies competition. Therefore, in the thesis, first descripe morphologic characters and analyse echolocation calls of five bat species, Myotis daubentonii, M. formosus, Murina leucogaster, M. ikonnikovi, Rhinolophus ferrumequinum in different of sex and states which includes flight, stationary, hanging and crawling; sencondly, study the difference of echolocation calls during flight, diets, morhpologic characters, foraging habitat, foraging strategy and action rules of the five bat species by contrast. At last, conclude that that niche separation of the bats are evident, what make bats coexist stably.
Rhinolophus ferrumequinum is biggest in these 5 species(mess is about 20g), use CF echolocation calls, which is sensitive to wing-vibrated insects. It feeds mainly on big-medium insects of order Lepidoptera and Coleoptera, foraging in wide space and sparse-foliage forest by aerial feeding and gleaning. Other four species all use FM echolocation calls. Murina leucogaster and M. formosus are nearly the same in body size, the former’s wave width is wondrously big. It feeds mainly on medium-small insects of order Lepidoptera and Diptera, foraging in complicated forest by aerial feeding. The latter's wave width is medium. It feeds mainly on medium-small insects of order Coleoptera and Lepidoptera, foraging in medium-complicated forest by aerial feeding or gleaning. M. daubentonii
and M. ikonnikovi are the smallest ones in size. The former’ wave width is medium. It feeds mainly on aquatic small insects of order Diptera and Trichoptera, foraging in 5-20cm above water surface and midium- complicated forest by trawling and aerial feeding. The latter’s wave width is shortest, which have disadvantage to discriminate targets from enviroment. It feeds mainly on even smaller insects of order Lepidoptera and Diptera, foraging in wide space or sparse-foliage forest by aerial feeding. Vision is an important assistant means during foraging. These interspecies differences decrease interspecies competition, and ensure they can coexist in one cave steadily.
At the same time, the result show that there is significant difference of echolocation calls in different states in all species, and only in M. daubentonii and M. ikonnikovi that there is significant difference in body size and echolocation calls betweed two sexes, but not in R. Ferrumequinum. The difference in calls and morphology is possibly the means of escaping interspecies sound jamming and ardent competition. While the difference of sound during flight indicate that bats may prefer to different foraging styles, foraging habitat, targets size and so on.
引文
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[2] Arita H T, et al. Flight and echolocation in the ecology and evolution of bats. Tree. 1997,12 (2). 53~59
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