菲菊头蝠不同地理种群回声定位声波差异及其影响因子

回声定位声波地理差异及其形成原因是蝙蝠生态学研究领域一个基本而关键的问题,对于探索物种生存机制、物种形成及其保护具有重要科学意义。本研究从较大地理尺度上(9个地理种群)研究了菲菊头蝠(Rhinolophus pusillus)回声定位声波结构的地理差异,并进一步探讨了影响回声定位声波地理种群差异的因素。结果表明,菲菊头蝠雌性的体型较雄性略大,其主频较高。不同地理种群之间回声定位声波差异明显,包括脉冲持续时间、脉冲间隔、主频以及带宽在不同的地理种群之间均表现出一定程度的差异。进一步分析发现,不同地理种群之间的雌性菲菊头蝠前臂长和体重均与主频呈较弱的负相关,降雨量与雌性的主频呈较强的正相关;而不同地理种群之间的雄性前臂长、体重和降雨量与回声定位声波参数均无相关性;此外,地理距离、温度、湿度均与雌雄回声定位声波参数无相关性。本研究结果表明,菲菊头蝠不同地理种群间的回声定位声波出现明显差异,其中,体型和降雨量为主要影响因子,说明蝙蝠回声定位叫声的进化主要受到了当地生境的影响,表现出动物对不同生境的适应性进化。     

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.     

三种共栖蝙蝠的回声定位信号特征及其夏季食性的比较

2005年6至9月,对桂林市郊区两个山洞中高颅鼠耳蝠(Myotissiligorensis)、菲菊头蝠(Rhinolo-phuspusillus)和黑髯墓蝠(Taphozousmelanopogon)的回声定位叫声特征和食性进行分析,并结合其形态特征与野外观察,推断其捕食生境和捕食策略。研究结果发现:黑髯墓蝠体型最大,声音特征属短调频型多谐波,一般为4个谐波,能量主要集中在第二谐波上,主频率为(32·84±1·17)kHz,选择鞘翅目和双翅目昆虫为主要食物;高颅鼠耳蝠(长调频型)和菲菊头蝠(长恒频-调频型),体型都较小,主频率分别是(84·44±8·13)kHz和(110·78±1·65)kHz,以双翅目昆虫为主要食物;而菲菊头蝠则以鞘翅目和双翅目昆虫为主要食物。上述结果证明,高颅鼠耳蝠、菲菊头蝠和黑髯墓蝠在声音和食物组成等方面出现了明显分化。     

基于核基因RAG1部分序列探讨菊头蝠科和蹄蝠科的系统发育关系

应用核基因重组激活基因1(Recombination activating gene,RAG1)部分序列对贵州9种菊头蝠和5种蹄蝠的系统发育关系进行了研究,运用贝叶斯法(Bayesian inference,BI)、最大似然法(Maximum likelihood,ML)和邻接法(Neighbor-joining,NJ)构建了系统进化树。研究结果表明:1.菊头蝠科和蹄蝠科形成两个平行的分支,支持它们是两个平行科的结论。2.马铁菊头蝠是菊头蝠科中第一个独立出来的亚分支,与其余8种菊头蝠的亲缘关系最远;Rhinolophus sp.与大菊头蝠聚合为一支,推测Rhinolophus sp.可能是大菊头蝠,或者是大菊头蝠的近缘种,它们是继马铁菊头蝠之后分离出来的第二亚分支;中华菊头蝠、栗黄菊头蝠、中菊头蝠聚合为第三亚分支,贵州菊头蝠、大耳菊头蝠、菲菊头蝠聚合为第四亚分支,此两分支形成姊妹群。3.在蹄蝠科的亚分支中,三叶蹄蝠、小蹄蝠、普氏蹄蝠依次分离出来,最后分出的是大蹄蝠和中蹄蝠。     

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.     

三种共栖蝙蝠的回声定位信号特征及其夏季食性的比较

2005年6至9月，对桂林市郊区两个山洞中高颅鼠耳蝠(Myotis siligorensis)、菲菊头蝠(Rhinolophus pusillus)和黑髯墓蝠(Taphozous melanopogon)的回声定位叫声特征和食性进行分析，并结合其形态特征与野外观察，推断其捕食生境和捕食策略。研究结果发现：黑髯墓蝠体型最大，声音特征属短调频型多谐波，一般为4个谐波，能量主要集中在第二谐波上，主频率为(32.84±1.17)kHz，选择鞘翅目和双翅目昆虫为主要食物；高颅鼠耳蝠(长调频型)和菲菊头蝠(长恒频-调频型)，体型都较小，主频率分别是(84.44±8.13)kHz和(110.78±1.65)kHz，以双翅目昆虫为主要食物；而菲菊头蝠则以鞘翅目和双翅目昆虫为主要食物。上述结果证明，高颅鼠耳蝠、菲菊头蝠和黑髯墓蝠在声音和食物组成等方面出现了明显分化。     

安徽的菊头蝠

菊头蝠属于翼手目菊头蝠科,本科和蹄蝠科HIPPOSIDERIDAE的脸部均具复杂的鼻叶(皮肤叶)。有人将它们分列为亚科,隶属于菊头蝠科内。本文仍按Miller(1907)等人意见各单列为一科。本科仅具一属,约69种,国内有14种。我省已知9种,主要分布于皖南地区,马铁菊头蝠及角菊头蝠等少数种类延伸至江淮及皖西。它们的鼻叶主要分为马蹄叶、鞍状叶和顶叶三个部分(见图),鼻叶的特点是种的一个重要分类依据。齿式为:1、1、2、3/2、1、2、3=32枚。图鼻叶示意图(左:角菊头蝠中:绒菊头蝠右:鲁氏菊头蝠)种的检索表(括号内系分布地区)1.鞍状叶与连接突间具凹…     

基于线粒体16S rRNA基因部分序列探讨贵州9种菊头蝠(翼手目:菊头蝠科)的分子系统进化关系

依据部分线粒体16S rRNA基因序列,我们对贵州九种菊头蝠进行了分子系统进化关系研究.结果表明:马铁菊头蝠与其他菊头蝠亲缘关系较远,它可能是最原始的种类.中华菊头蝠和贵州菊头蝠聚在同一分支,表明它们之间亲缘关系非常接近,高鞍菊头蝠和贵州菊头蝠的DNA差异百分比最小(1.13%),因此认为这两个种亲缘关系也非常接近.     

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。     

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。     

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。     

三峡工程重庆库区民翼手类研究

1996年至1997年,通过对三峡工程重庆库区13个县（区）68个洞穴的考察和标本采集,结合历史资料记述,确认重庆库区共有翼手目动物4科26种。其中8种为重庆市新记录,分别是绒菊头蝠、皮菊头蝠、西南鼠耳蝠、水鼠耳蝠、尖耳鼠耳蝠、大足鼠耳蝠、长折翼蝠和皱唇蝠。研究发现冀手类更喜欢潮湿、低海拔、人类干扰少、洞深在50m以上、周围是农田－草灌的洞穴。     

Mechanical adaptations for echolocation in the cochlea of the bat Hipposideros lankadiva

The cochlear mechanics of bats with long constant-frequency components in their echolocation calls are sharply tuned to the dominant second harmonic constant frequency. Hipposiderid bats employ a shorter constant-frequency call component whose frequency is less stable than in long-constant-frequency bats. To investigate to what degree cochlear mechanics in hipposiderid bats are already specialized for the processing of constant frequencies, we recorded distortion-product otoacoustic emissions in Hipposideros lankadiva. Iso-distortion threshold curves for the 2f1-f2 distortion-product otoacoustic emission reveal a threshold maximum close to the second harmonic constant frequency, between 65.0 and 70.0 kHz, and a second insensitivity close to the first harmonic constant frequency. The group delay of the 2f1-f2 distortion is prolonged for both frequency ranges, indicating that a specialized cochlear resonance may act to absorb the constant-frequency call components. Compared to long-constant-frequency bats, the threshold maximum at the second harmonic constant frequency is less pronounced and the optimum cochlear frequency separation is larger. Distortion-product otoacoustic emission suppression tuning curves and neuronal tuning curves recorded from neurons in the cochlear nucleus display an increase of tuning sharpness close to the second harmonic constant-frequency range which is smaller than that reported for long-constant-frequency bats. Our data suggest that the cochlea of hipposiderid bats represents an intermediate state between that of non-specialized bats and long-constant-frequency bats.     

食物大小对棕果蝠和犬蝠取食行为的影响

按照最优化觅食理论,动物在取食时需在能量获取与捕食风险之间权衡。本文通过室内行为实验,研究两种旧大陆果蝠棕果蝠和犬蝠对食物大小的选择规律与取食策略。按体积由小到大将苹果分为Ⅰ型、Ⅱ型、Ⅲ型、Ⅳ型4 种类型的食物块,通过红外相机观察果蝠对不同大小食块的取食情况,并就其对各类型食块的取食率、取食次数和停留时间进行统计分析。结果表明:这两种果蝠对Ⅱ型和Ⅲ型食块的取食率显著高于Ⅰ型和Ⅳ型;对Ⅰ型食块的取食次数显著高于Ⅱ型和Ⅳ型;对Ⅳ型食块的停留时间显著高于Ⅰ型和Ⅱ型。它们在摄取体积较小的食块时,以取走后进食为主要取食方式,但摄取大体积食块时则主要在原地进食。取食过程中,果蝠优先选择大小适于搬运的食块,是捕食风险与能量收益权衡的结果。     

广东省发现长指鼠耳蝠及其回声定位声波特征

2013年7月在广东南岭国家级自然保护区阳山县境内(24°48′39.5′′N,112°51′01.3′′E,海拔155 m)捕捉到4只雄性蝙蝠,体型小,前臂长34.1~34.7 mm,颅全长14.3~14.7 mm;体毛浓密,背毛毛基黑色,毛尖灰色,腹毛黑或棕色,毛尖奶油白,靠近肛门处腹毛浅灰色,些许白色;翼膜附着于跖部末端;后足特别延展,长度超过胫骨长之半;鉴定为长指鼠耳蝠[Myotis longipes(Dobson1873)]。同时,基于Cyt b基因序列(1 140 bp)构建的部分鼠耳蝠物种系统进化关系,进一步确认上述标本为长指鼠耳蝠,为广东省翼手目新纪录。该种蝙蝠在我国贵州、广西、重庆有分布记载,但标本和相关资料很少。本文给出了长指鼠耳蝠的外形和头骨特征测量数据,并与印度的标本进行了对比。长指鼠耳蝠的回声定位声波为调频型(FM),主频率为68.2 k Hz;此外,对其分类地位和分布状况进行了讨论。标本保存于广东省生物资源应用研究所。     

在不同状态和生境复杂度中大蹄蝠回声定位叫声的可塑性

蝙蝠回声定位声波的可塑性对其适应不同状态、生境以及捕食和社会交流具有重要的作用。为进一步研究大蹄蝠的回声定位声波在不同状态和生境下的可塑性,通过室内行为实验,对大蹄蝠在4 种不同状态(室内飞行、静息、布袋内和手持)和4 种不同生境复杂度(室外、室内0 棵树、室内1 棵树、室内5 棵树)条件下飞行的回声定位声波特征进行研究。结果表明:大蹄蝠的回声定位声波为CF - FM 型,通常连续发出2 - 4 个脉冲组成一个脉冲组。对比大蹄蝠在4 种不同状态下的回声定位叫声发现,主频按静息、布袋内、手持、飞行的顺序依次降低,后端FM 频宽则按手持、布袋内、飞行和静息的顺序依次变短;而脉冲间隔和脉冲时程则均按静 息、飞行、布袋内、手持的顺序依次增加。对比大蹄蝠在4 种不同生境复杂度中飞行的回声定位叫声发现,主频按室外、室内0 棵树、室内1 棵树、室内5 棵树依次提高,而脉冲时程及脉冲间隔则依次缩短;室外放飞条件下的后端FM 频宽比室内飞行的短。研究结果说明,大蹄蝠在不同状态、不同生境复杂度条件下的回声定位叫声具有明显的可塑性和生境适应性。     

广东省蝙蝠新纪录——大黑伏翼

<正>广东南岭地区蝙蝠多样性较为丰富,2000年后发表的9种广东蝙蝠新纪录中即有5种采集于南岭(吴毅等,2001,2005;张燕均等,2010;周全等,2011,2012;陈毅等,2013)。2013年7月21日,我们在保护区动物资源调查过程中,于广东南岭国家级自然保护区八宝山保护站(北纬24°55'43.4″,东经113°0'57.0″,海拔1 000 m)附近的路上夜晚挂网捕捉到一批蝙蝠     

The relative influence of competition and prey defences on the trophic structure of animalivorous bat ensembles

Deterministic filters such as competition and prey defences should have a strong influence on the community structure of animals like animalivorous bats which have life histories characterized by low fecundity, low predation risk, long life expectancy and stable populations. We investigated the relative influence of these two deterministic filters on the trophic structure of animalivorous bat assemblages in South Africa. We used null models to test if patterns of dietary overlap were significantly different from patterns expected by chance and multivariate analyses to test the correlations between diet and phenotype (body size, wing morphology and echolocation). We found little evidence that competition structured the trophic niche of coexisting bats. Contrary to predictions from competition, dietary overlap between bats of ensembles and functional groups (open-air, clutter-edge, and clutter foragers) were significantly higher than expected by chance. Instead, we found support for the predictions of the allotonic frequency hypothesis: there were significant relationships between peak echolocation frequency and the proportion of moths in the diets of bats at local and regional scales, and peak echolocation frequency was the best predictor of diet even after we controlled for the influence of body size and phylogeny. These results suggest that echolocation frequency and prey hearing exert more influence on the trophic structure of sympatric animalivorous bats than competition. Nonetheless, differential habitat use and sensory bias may also be major determinants of trophic structure because these are also correlated with frequencies of bat calls.     

The Evolution of Bat Vestibular Systems in the Face of Potential Antagonistic Selection Pressures for Flight and Echolocation

The vestibular system maintains the body’s sense of balance and, therefore, was probably subject to strong selection during evolutionary transitions in locomotion. Among mammals, bats possess unique traits that place unusual demands on their vestibular systems. First, bats are capable of powered flight, which in birds is associated with enlarged semicircular canals. Second, many bats have enlarged cochleae associated with echolocation, and both cochleae and semicircular canals share a space within the petrosal bone. To determine how bat vestibular systems have evolved in the face of these pressures, we used micro-CT scans to compare canal morphology across species with contrasting flight and echolocation capabilities. We found no increase in canal radius in bats associated with the acquisition of powered flight, but canal radius did correlate with body mass in bat species from the suborder Yangochiroptera, and also in non-echolocating Old World fruit bats from the suborder Yinpterochiroptera. No such trend was seen in members of the Yinpterochiroptera that use laryngeal echolocation, although canal radius was associated with wing-tip roundedness in this group. We also found that the vestibular system scaled with cochlea size, although the relationship differed in species that use constant frequency echolocation. Across all bats, the shape of the anterior and lateral canals was associated with large cochlea size and small body size respectively, suggesting differential spatial constraints on each canal depending on its orientation within the skull. Thus in many echolocating bats, it seems that the combination of small body size and enlarged cochlea together act as a principal force on the vestibular system. The two main groups of echolocating bats displayed different canal morphologies, in terms of size and shape in relation to body mass and cochlear size, thus suggesting independent evolutionary pathways and offering tentative support for multiple acquisitions of echolocation.