6种共栖菊头蝠共存机制及菲律宾菊头蝠组蝙蝠声波主频偏离机制研究
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摘要
翼手目(Chiroptera)动物是哺乳动物中仅次于啮齿目的第二大目,具有极高的物种多样性,对维持生态系统平衡具有重要作用。然而,近年来由于生境丧失、滥用农药等人为干扰因素,翼手目动物的生物多样性急剧丧失。因此,亟需开展对翼手目动物的生态学研究。物种的稳定共存是维持生物多样性的基础之一,而翼手目动物常多个物种混居,且群落成员多为生态需求相似的同一共位群(Guild)物种,为研究物种共存机制提供了理想的模型。通过对翼手目动物共存机制的研究,可为制定物种保护措施提供科学依据及理论指导。此外,回声定位蝙蝠的声波结构是其生态适应力的重要体现,有的蝙蝠声波结构会发生异于常规的变化,这种非常规现象可能暗示了蝙蝠在进化过程中对自然选择压力的应答,研究这种现象的产生机制为全面揭示蝙蝠的进化辐射及适应性进化提供了新视角。
本研究以云南省6种共栖菊头蝠科蝙蝠:马铁菊头蝠(Rhinolophus ferrumequinum)、中华菊头蝠(R. sinisus)、中菊头蝠(R. affinis)、大耳菊头蝠(R. macrotis)、短翼菊头蝠(R. lepidus)及菲菊头蝠(R. pusillus)为对象,基于生态位分化理论研究物种的共存机制。同时,首次通过猎物选择行为学实验研究了大耳菊头蝠声波主频偏离与体型负相关规律的生态驱动力;首次基于生理结构角度揭示了发声器官及声信号接收器官形态的非常规变化是菲律宾菊头蝠组蝙蝠声波产生非常规分化的生理基础。主要内容如下:
1.通过几何形态测量法比较了6种同域共存菊头蝠头骨的形态差异。结果表明,6种共栖菊头蝠头骨的大小和形状均存在显著差异,头骨形状的变化协同于尺寸的变化。上颌骨形状的种间差异体现了蝙蝠对回声定位功能、猎物咀嚼等多重功能的适应,下颌骨的形状变化主要体现了蝙蝠对食物咀嚼功能的适应。
2.通过Variance Test检验了6种共栖菊头蝠的5个重要特征参数:回声定位声波主频、前臂长、翼载及头骨质心的种间差异及其排布模式,结果表明在6种共栖蝙蝠中,声波主频的种间差异呈均匀排布,而其余参数的种间差异为随机排布。该结果不能回答竞争作用是否是群落结构的决定作用,但声信号的群落特征替代(Community-wide characters displacement))暗示了共栖蝙蝠感官能力的差异对维持物种共存的重要作用。
3.通过猎物选择行为学实验验证了菲律宾菊头蝠组蝙蝠声波主频偏离常规的生态驱动力。猎物选择实验表明,菲律宾菊头蝠组蝙蝠——大耳菊头蝠的猎物选择模式与和其同域共存,体型相似但声波主频存在明显分化的短翼菊头蝠的猎物选择模式之间存在差异,表明了群落内食物资源利用分化的竞争作用是促使大耳菊头蝠声波主频偏离常规的生态驱动力之一。
4.鼻腔及外耳是菊头蝠科蝙蝠声波发射及回声信号接收系统中的重要组成部分。通过对比10种菊头蝠科蝙蝠的鼻腔形态及外耳长度研究了3种菲律宾菊头蝠组蝙蝠声波主频偏离常规的生理结构基础。结果表明,10种蝙蝠的鼻腔尺寸及外耳长度分别与其体型呈正相关,而声波主频偏离与体型负相关规律的3种菲律宾菊头蝠组蝙蝠的鼻腔大小及外耳长度均相应的偏离了其与体型的正相关规律。该结果暗示形态偏离常规的声信号处理系统是3种菲律宾菊头蝠组蝙蝠声波主频偏离与体型负相关规律的生理基础之一。对前臂长、鼻腔大小及外耳长度的偏相关分析结果表明发声器官与回声信号接收器官之间的相关性强于其与体型之间的相关性,暗示了声信号接收器官与发声器官的形态之间具有良好的适应性。
5.研究了10种蝙蝠前臂长、鼻腔大小及耳长与蝙蝠的回声定位声波主频之间的关系。简单相关分析及一元线性回归分析的结果表明,蝙蝠的回声定位声波主频分别负相关于这3个形态参数;3种菲律宾菊头蝠组蝙蝠的声波主频偏离了与体型的负相关关系,但均符合基于鼻腔尺寸或外耳长度的预测。Mantel检验结果表明,10种蝙蝠的声波主频种间分歧程度与其鼻腔大小及耳长的种间差异程度呈正相关,而与体型差异相关性较弱。偏相关分析、多元线性回归分析及通径分析结果表明,声信号处理系统及体型均作用于蝙蝠的回声定位声波主频,但声信号处理系统形态与声波主频相关性更高且影响更加直接。在声信号处理系统中,发声器官与回声定位声波主频的相关性高于回声接收器官与声波主频的相关性,暗示了发声器官的形态结构是影响回声定位蝙蝠声波主频的首要生理因素。
Bats constitute the second largest order of mammals worldwide with high species richness. As an important biological indicator for ecosystem, the bats diversity has declined dramatically due to human disturbance. So, the ecology study for bats is seriously needed. The stable coexistence of sympatric species maintains species diversity. Bats are ideal organisms with which to test hypotheses related to coexistence. Research on the coexistent mechanism in bats communities can provide the significant knowledge for conservation biology. Furthermore, the echolocation calls and morphologies of bats have profound effects on the way they live and they often have an allometry relationship. However, some bats species deviated from this correlation. The physics of echolocation is intimately related to the ecology of insectivorous bats. Deviations from allometric relationships may imply the response of bats to nature selection pressure, providing insight into the evolutionary radiation and the adaptive evolution of bats.
From 2004 to 2008, we studied the coexistent mechanism in six sympatric rhinolophids species: Rhinolophus ferrumequinum, R. sinisus, R. affinis, R. macrotis, R. lepidus, R. pusillus in Shuanghe town, Yunnan Province. We also studied the ecological and physical factors which are responsible for deviation of echolocation call frequencies from allometry in rhinolophids from Phillippinensis-group. The details are as follow.
1. The skull form of six sympatric rhinolophids species was compared by geometric morphometrics method. Both size and shape of crania and mandible were significant different among these sympatric species. The variances in shape of crania and mandible were related to the size variances of crania and mandible, respectively. The variances in crania may imply the complex adaptation for multiply functions, such as echolocation and food chew, while the variances in mandible may be the result of adaptation for diet.
2. The Variance Tests were used to detect community-wide characters displacement in five character parameters collected from 6 sympatric species, including peak frequency, forearm length, wing loading and centroid size of crania and skull, which associated with the resource using. The results showed support for evenly displaced acoustic character, no support on other four parameters. These results showed the importance of sensory echology in maintaining animal coexistence and the mixed support for competition.
3. The prey selection experiment was conducted to test the ecological factor leading to deviating peak frequency of echolocation call in Rhinolophus macrotis, one species from Phillippinensis-group. The prey selection strategy of R. macrotis was compared with the R.lepidus. These two sympatric species are morphologically similar but acoustically divergent: R.macrotis has an echolocation frequency significantly lower than that predicted by the allometric relationship, whereas that of R. lepidus agreed with expectations. These results confirmed the existence of finely tuned trophic niche differentiation and suggested that food resource partitioning is one of the factors leading to lower peak frequency of calls in R. macrotis.
4. Nasal cavity and outer ear are important part of sound emission and echo reception for rhinolophids, respectively. We compared the nasal chamber size and ear length of 10 species rhinolophids, including three species from Phillippinensis-group, to investigate the physical factor relating the divergent peak frequency of echolocation call in Phillippinensis-group bats. The results showed that the nasal cavity size and ear length were positively related to the forearm length, respectively. We found that both of the nasal chamber size and ear length of three species of rhinolophids from Phillippinensis-group also deviated from the allomety relationship with body size. This result indicated that the distinct developed sound signal system is one of the physical bases for divergent peak frequency of echolocation call from prediction by forearm in Phillippinensis-group bats. The result of partial correlation among three morphology parameters showed that the relationship between sound emission and echo reception were stronger than that with body size, implying the closely adaption between those different functional parts.
5. We also studied the relationship between echolocation call frenquency and different morphology parameters. The results of bivariate correlation analysis and linear regression analysis showed that the call peak frequency was negtively related to the forearm length, nasal cavity size and ear length, respectively. The call peak frequencies of three rhinolophid species from Phillippinensis-group were accordance with the prediction basing on nasal cavity size or ear length. The results of Mantel test suggested that the divergence in call frequency were positively with divergences in nasal cavity size or era length but not to that in forearm length. The results of partial correlation analysism, multiply regression analysis and path analysis suggested that the peak frequency was stronger with the morphology of sound manage system than that with body size. Those results also implied that the nasal cativty form may be the most important physical factor influencing the call frequency.
本研究以云南省6种共栖菊头蝠科蝙蝠:马铁菊头蝠(Rhinolophus ferrumequinum)、中华菊头蝠(R. sinisus)、中菊头蝠(R. affinis)、大耳菊头蝠(R. macrotis)、短翼菊头蝠(R. lepidus)及菲菊头蝠(R. pusillus)为对象,基于生态位分化理论研究物种的共存机制。同时,首次通过猎物选择行为学实验研究了大耳菊头蝠声波主频偏离与体型负相关规律的生态驱动力;首次基于生理结构角度揭示了发声器官及声信号接收器官形态的非常规变化是菲律宾菊头蝠组蝙蝠声波产生非常规分化的生理基础。主要内容如下:
1.通过几何形态测量法比较了6种同域共存菊头蝠头骨的形态差异。结果表明,6种共栖菊头蝠头骨的大小和形状均存在显著差异,头骨形状的变化协同于尺寸的变化。上颌骨形状的种间差异体现了蝙蝠对回声定位功能、猎物咀嚼等多重功能的适应,下颌骨的形状变化主要体现了蝙蝠对食物咀嚼功能的适应。
2.通过Variance Test检验了6种共栖菊头蝠的5个重要特征参数:回声定位声波主频、前臂长、翼载及头骨质心的种间差异及其排布模式,结果表明在6种共栖蝙蝠中,声波主频的种间差异呈均匀排布,而其余参数的种间差异为随机排布。该结果不能回答竞争作用是否是群落结构的决定作用,但声信号的群落特征替代(Community-wide characters displacement))暗示了共栖蝙蝠感官能力的差异对维持物种共存的重要作用。
3.通过猎物选择行为学实验验证了菲律宾菊头蝠组蝙蝠声波主频偏离常规的生态驱动力。猎物选择实验表明,菲律宾菊头蝠组蝙蝠——大耳菊头蝠的猎物选择模式与和其同域共存,体型相似但声波主频存在明显分化的短翼菊头蝠的猎物选择模式之间存在差异,表明了群落内食物资源利用分化的竞争作用是促使大耳菊头蝠声波主频偏离常规的生态驱动力之一。
4.鼻腔及外耳是菊头蝠科蝙蝠声波发射及回声信号接收系统中的重要组成部分。通过对比10种菊头蝠科蝙蝠的鼻腔形态及外耳长度研究了3种菲律宾菊头蝠组蝙蝠声波主频偏离常规的生理结构基础。结果表明,10种蝙蝠的鼻腔尺寸及外耳长度分别与其体型呈正相关,而声波主频偏离与体型负相关规律的3种菲律宾菊头蝠组蝙蝠的鼻腔大小及外耳长度均相应的偏离了其与体型的正相关规律。该结果暗示形态偏离常规的声信号处理系统是3种菲律宾菊头蝠组蝙蝠声波主频偏离与体型负相关规律的生理基础之一。对前臂长、鼻腔大小及外耳长度的偏相关分析结果表明发声器官与回声信号接收器官之间的相关性强于其与体型之间的相关性,暗示了声信号接收器官与发声器官的形态之间具有良好的适应性。
5.研究了10种蝙蝠前臂长、鼻腔大小及耳长与蝙蝠的回声定位声波主频之间的关系。简单相关分析及一元线性回归分析的结果表明,蝙蝠的回声定位声波主频分别负相关于这3个形态参数;3种菲律宾菊头蝠组蝙蝠的声波主频偏离了与体型的负相关关系,但均符合基于鼻腔尺寸或外耳长度的预测。Mantel检验结果表明,10种蝙蝠的声波主频种间分歧程度与其鼻腔大小及耳长的种间差异程度呈正相关,而与体型差异相关性较弱。偏相关分析、多元线性回归分析及通径分析结果表明,声信号处理系统及体型均作用于蝙蝠的回声定位声波主频,但声信号处理系统形态与声波主频相关性更高且影响更加直接。在声信号处理系统中,发声器官与回声定位声波主频的相关性高于回声接收器官与声波主频的相关性,暗示了发声器官的形态结构是影响回声定位蝙蝠声波主频的首要生理因素。
Bats constitute the second largest order of mammals worldwide with high species richness. As an important biological indicator for ecosystem, the bats diversity has declined dramatically due to human disturbance. So, the ecology study for bats is seriously needed. The stable coexistence of sympatric species maintains species diversity. Bats are ideal organisms with which to test hypotheses related to coexistence. Research on the coexistent mechanism in bats communities can provide the significant knowledge for conservation biology. Furthermore, the echolocation calls and morphologies of bats have profound effects on the way they live and they often have an allometry relationship. However, some bats species deviated from this correlation. The physics of echolocation is intimately related to the ecology of insectivorous bats. Deviations from allometric relationships may imply the response of bats to nature selection pressure, providing insight into the evolutionary radiation and the adaptive evolution of bats.
From 2004 to 2008, we studied the coexistent mechanism in six sympatric rhinolophids species: Rhinolophus ferrumequinum, R. sinisus, R. affinis, R. macrotis, R. lepidus, R. pusillus in Shuanghe town, Yunnan Province. We also studied the ecological and physical factors which are responsible for deviation of echolocation call frequencies from allometry in rhinolophids from Phillippinensis-group. The details are as follow.
1. The skull form of six sympatric rhinolophids species was compared by geometric morphometrics method. Both size and shape of crania and mandible were significant different among these sympatric species. The variances in shape of crania and mandible were related to the size variances of crania and mandible, respectively. The variances in crania may imply the complex adaptation for multiply functions, such as echolocation and food chew, while the variances in mandible may be the result of adaptation for diet.
2. The Variance Tests were used to detect community-wide characters displacement in five character parameters collected from 6 sympatric species, including peak frequency, forearm length, wing loading and centroid size of crania and skull, which associated with the resource using. The results showed support for evenly displaced acoustic character, no support on other four parameters. These results showed the importance of sensory echology in maintaining animal coexistence and the mixed support for competition.
3. The prey selection experiment was conducted to test the ecological factor leading to deviating peak frequency of echolocation call in Rhinolophus macrotis, one species from Phillippinensis-group. The prey selection strategy of R. macrotis was compared with the R.lepidus. These two sympatric species are morphologically similar but acoustically divergent: R.macrotis has an echolocation frequency significantly lower than that predicted by the allometric relationship, whereas that of R. lepidus agreed with expectations. These results confirmed the existence of finely tuned trophic niche differentiation and suggested that food resource partitioning is one of the factors leading to lower peak frequency of calls in R. macrotis.
4. Nasal cavity and outer ear are important part of sound emission and echo reception for rhinolophids, respectively. We compared the nasal chamber size and ear length of 10 species rhinolophids, including three species from Phillippinensis-group, to investigate the physical factor relating the divergent peak frequency of echolocation call in Phillippinensis-group bats. The results showed that the nasal cavity size and ear length were positively related to the forearm length, respectively. We found that both of the nasal chamber size and ear length of three species of rhinolophids from Phillippinensis-group also deviated from the allomety relationship with body size. This result indicated that the distinct developed sound signal system is one of the physical bases for divergent peak frequency of echolocation call from prediction by forearm in Phillippinensis-group bats. The result of partial correlation among three morphology parameters showed that the relationship between sound emission and echo reception were stronger than that with body size, implying the closely adaption between those different functional parts.
5. We also studied the relationship between echolocation call frenquency and different morphology parameters. The results of bivariate correlation analysis and linear regression analysis showed that the call peak frequency was negtively related to the forearm length, nasal cavity size and ear length, respectively. The call peak frequencies of three rhinolophid species from Phillippinensis-group were accordance with the prediction basing on nasal cavity size or ear length. The results of Mantel test suggested that the divergence in call frequency were positively with divergences in nasal cavity size or era length but not to that in forearm length. The results of partial correlation analysism, multiply regression analysis and path analysis suggested that the peak frequency was stronger with the morphology of sound manage system than that with body size. Those results also implied that the nasal cativty form may be the most important physical factor influencing the call frequency.
引文
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