广西扁颅蝠与褐扁颅蝠的食物选择

2002年3～10日在广西南宁地区的宁明县和龙州县,用超声波监测和网捕法确定扁颅蝠与褐扁颅蝠的捕食区,在捕食区内用粘捕法调查潜在的食物量;用粪便分析法确定食物组成。在宁明县和龙州县扁颅蝠和褐扁颅蝠的潜在食物量都是以双翅目昆虫为主(45．93％以上),其次为鞘翅目(12．59％以上)和膜翅目(7．47％以上);但双翅目、膜翅目在两地差异显著。宁明县扁颅蝠的食物组成以双翅目(40．33％)和膜翅目(38．46％)为主,鞘翅目(16．07％)次之;龙州县扁颅蝠和褐扁颅蝠食物组成中以膜翅目(63．37％,62．34％)为主,其次为双翅目(21．57％,29．62％)、鞘翅目(11．59％,5．96％);对比两地扁颅蝠的食物组成发现,膜翅目和双翅目差异极显著。对比食物组成与潜在食物量发现,两种蝙蝠对膜翅目为正选择,对其他目负选择或无选择,均为选择性捕食者。     

扁颅蝠和褐扁颅蝠体表寄生革螨的宿主选择

该文通过野外调查和室内行为实验,研究了扁颅蝠(Tylonycteris pachypus)和褐扁颅蝠(T.robustula)与其体表寄生革螨(雷氏巨刺螨(Macronyssus radovskyi)及拟雷氏巨刺螨(M.pararadovskyi))之间的关系.在野外自然条件下,雌性扁颅蝠的体表寄生革螨负荷量与宿主健康指数呈正相关(Spearman:rs=0.55,P＜0.01,n=24),而在雄性扁颅蝠以及雌、雄褐扁颅蝠中则无相关性(P＞0.05).室内原宿主感染实验发现,扁颅蝠和褐扁颅蝠体表寄生革螨均明显倾向于选择各自的雄性宿主,扁颅蝠雌、雄性感染率分别为(42±12)％和(58±12)％(t=-3.6,df=31,P＜0.01);褐扁颅蝠雌、雄性感染率分别为(37±11)％和(63±11)％(t=-6.1,df=26,P＜0.001).用扁颅蝠体表寄生革螨(拟雷氏巨刺螨)对扁颅蝠与褐扁颅蝠交叉感染后发现,寄生革螨明显选择其原宿主扁颅蝠,扁颅蝠与褐扁颅蝠感染率分别为(71±13)％和(29±13)％(t=9.1,df=29,P＜0.001).以上结果表明,扁颅蝠和褐扁颅蝠的体表寄生革螨负荷量与宿主身体状态无明显相关性,而对宿主性别表现不同偏好;扁颅蝠的体表寄生革螨对宿主表现明显的专一性.     

扁颅蝠与褐扁颅蝠的集群结构

2001—2002年在广西宁明县和龙州县利用直接观察、捕捉测量(共捕到197群蝙蝠,全捕180群)和标记重捕法(标记了31群的101只扁颅蝠,重捕到36只)比较研究了扁颅蝠(tylonycteris pachypusa)与褐扁颅蝠(T．robustula)的集群结构。结果发现：扁颅蝠与褐扁颅蝠主要栖宿在刺竹(Bambusa stenostachya)的竹筒内,通过竹筒上的裂缝进出。扁颅蝠栖宿的竹筒长平均为27．7cm,外围直径平均为23．6cm;褐扁颅蝠的分别为28．3cm和23．8cm。扁颅蝠栖宿群大小为1—24只,褐扁颅蝠栖宿群大小为1—13只;2种蝙蝠的栖宿群中皆为独居所占比例最大(扁颅蝠为22．30％,褐扁颅蝠为40．63％),2只所占比例次之(分别为14．87％和18．75％),其它大小类型呈不规则变化。扁颅蝠栖宿群的性别组成,以雌雄混居最常见(占54．72％),其次为独居雄性(占20．95％),而褐扁颅蝠栖宿群雌雄混居群与独居雄性所占比例相当(均为40．63％)。2种蝙蝠的雄性趋向于独居,而雌性趋向于群居。扁颅蝠与褐扁颅蝠可以栖宿在同一片竹林内,并且可以在不同时间轮流使用同一个栖宿竹筒,但2种蝙蝠从未共栖于同一个竹筒内。另外,标记重捕扁颅蝠发现：扁颅蝠经常变换栖宿竹筒(栖宿竹筒不固定);同时栖宿群之间经常发生个体交换[动物学报50(3)：326—333．2004]。     

Roost-site selection among flat-haded bats (Tylonycteris spp.)

During six years, 1962–68, the biology of the bats Tylonycteris pachypus and T. robustula has been studied at the University of malaya Field Studies Centre, Ulu Gombak, Selangor, Malaysia. Both species were often found roosting within internodes of the bamboo Gigantochloa scortechinii , entering and leaving through narrow vertical slits caused by the beetle Lasiochila goryi . In this paper the roost sites are described and evidence is presented that each species of bat actively selected roosts of a different range of dimensions. The ecological implications of the separation of the species by this factor are discussed.     

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。     

Roosting associations of flat-headed bats, Tylonycteris species (Chiroptera: Vespertilionidae) in Malaysia

Roosting associations of Tylonycteris pachypus and T. robustula were studied in West Malaysia. Both species roosted within the internodes of one species of bamboo, but were only once found together at roost. Roosting groups, which formed before entry in the morning, were not limited in size by the dimensions of the roost site. In both species the sexes exhibited different roosting behaviour, the males tending to be solitary and the females gregarious. Seasonal variations in associations occurred correlated with the reproductive cycle. Banding demonstrated that the populations were highly mobile, that the bats frequently changed roost sites, and that the roosting associations were of an ephemeral nature.     

无尾蹄蝠的回声定位声波特征及分析

采用超声波监听仪U30录制无尾蹄蝠自由飞行状态的回声定位声波,经Batsound3.0分析,其声波为高频(145.4±10.9kHz)、宽带(62.6±9.2kHz)、具两个谐波的短(1.67±0.4ms)FM型,不同于蹄蝠科其他蝙蝠的CF型,表明该科内物种声波类型存在多态性。头骨的形态测定分析支持其通过鼻腔发射声波,与蹄蝠科其他蝙蝠一致,表明该科内声波发射方式的单一性。适应环境的选择压力及翼型和声波的适应性可能是其选择FM型叫声的重要原因。     

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。     

中国巨刺螨属一新种和雷氏巨刺螨的重新描述(中气门目,巨刺螨科)(英文)

记述了采自广西的扁颅蝠Tylonycteris robustula(Thomas,1915)体表的寄生革螨1新种:拟雷氏巨刺螨Macronyssus pararadovskyi Tian,Jin et Zhang,sp.nov..新种与雷氏巨刺螨Macronyssus radovskyi(Domrow,1963)近似,二者的区别在于:新种背毛有27对,具有Z3;雌螨生殖腹板具有3根副毛,或部分标本为1根副毛在板的末端,另1对副毛位于板外;雄螨胸生殖腹板上具有4对胸毛和20根腹毛.而雷氏巨刺螨背毛有26对,不具有Z3.雌螨生殖腹板不具有副毛,即副毛均不在生殖腹板上;雄螨胸生殖腹板上具有4对胸毛和18根腹毛.另外,依据采自广西褐扁颅蝠Tylonycterispachypus(Temminck,1840)体表的标本,对雷氏巨刺螨Macronyssus radovskyi(Domrow,1963)的雌、雄螨成体和第1若螨进行了重新描述.所有标本均保存于贵州大学昆虫研究所标本馆.     

Flight performance, echolocation and foraging behaviour in noctule bats Nyctalus noctula

The noctule Nyctulus noctula (Schreber, 1774) is a relatively large (c. 25 g) insectivorous bat which catches insects on the wing (by aerial hawking). Emergence at a maternity roost was earliest relative to sunset when females were lactating, and bats may then have risked predation by flying at higher light levels during a period of high energy demand. Flight performance was quantified by using stereophotogrammetry. At feeding sites bats flew at 6.0 ± 2.1 m/s. This was faster than predicted minimum power speed (V _mp), and either between V _mp and maximum range speed (V _mr), or close to their predicted V _mr, depending on which aerodynamic model of flight power requirements was used. The echolocation behaviour of noctules is flexible. Long duration, low frequency calls (c. 20 kHz) with little frequency modulation were emitted while cruising, but at foraging sites the calls became more frequency-modulated. As the noctule is traditionally thought of as using low frequency echolocation, it was expected to receive weak echoes from small targets and therefore to specialize in eating large insect prey. Although the bats ate mainly beetles, large numbers of small dipterans were also eaten. The noctule is probably able to detect such small items because, when foraging, its calls become broadband and sweep from high frequencies. Higher harmonics are also present, and these may assist in the detection of small prey. In noctules, as in many bats, there appears to be a 1:1 link between wingbeat and call production during the search phase of foraging.     

Development and characterization of 13 novel microsatellite loci from the flat-headed bat (Tylonycteris pachypus) with cross-species amplification in closely related taxa

13 novel microsatellite loci was isolated using the enriched library method from genomic DNA of the flat-headed bat (Tylonycteris pachypus). These loci showed high levels of genetic polymorphism testing on 54 individuals sampled from Guangxi province, China. The number of observed alleles per locus ranged from 2 to 15. The expected and observed heterozygosity values ranged from 0.170 to 0.900 and from 0.185 to 0.944, respectively. One loci departed significantly from Hardy-Weinberg equilibrium (HWE) after Bonferroni correction and no Linkage disequilibrium was found between any pairs of loci. In addition, these loci were tested in the sister species, Tylonycteris robustula, seven loci amplified successfully and were also polymorphic.     

普通长翼蝠食性结构及其回声定位与体型特征

在普通长翼蝠(Miniopterus fuliginosus)的捕食区内用灯诱法和网捕法调查潜在食物(昆虫)种类; 用粪便分析法鉴定普通长翼蝠的食物组成,发现其主要捕食体型较大的鳞翅目和鞘翅目昆虫,体积百分比分别为55%和38%.普通长翼蝠具有相对狭长的翼,翼展比为6.94 ± 0.13;翼载为(9.85 ± 0.83)N/m2,相对较大.飞行状态下普通长翼蝠的回声定位叫声为调频下扫型,声脉冲时程为(1.45 ± 0.06)ms,脉冲间隔为(63.08 ± 21.55)ms,主频较低,为(44.50 ± 2.26)kHz.研究表明,普通长翼蝠的形态特征和回声定位特征与其捕食行为有着密切的联系.     

Genetic divergence and echolocation call frequency in cryptic species of Hipposideros larvatus s.l. (Chiroptera: Hipposideridae) from the Indo-Malayan region

The intermediate leaf-nosed bat ( Hipposideros larvatus ) is a medium-sized bat distributed throughout the Indo-Malay region. In north-east India, bats identified as H. larvatus captured at a single cave emitted echolocation calls with a bimodal distribution of peak frequencies, around either 85 kHz or 98 kHz. Individuals echolocating at 85 kHz had larger ears and longer forearms than those echolocating at 98 kHz, although no differences were detected in either wing morphology or diet, suggesting limited resource partitioning. A comparison of mitochondrial control region haplotypes of the two phonic types with individuals sampled from across the Indo-Malay range supports the hypothesis that, in India, two cryptic species are present. The Indian 98-kHz phonic bats formed a monophyletic clade with bats from all other regional populations sampled, to the exclusion of the Indian 85-kHz bats. In India, the two forms showed 12–13% sequence divergence and we propose that the name Hipposideros khasiana for bats of the 85-kHz phonic type. Bats of the 98-kHz phonic type formed a monophyletic group with bats from Myanmar, and corresponded to Hipposideros grandis , which is suggested to be a species distinct from Hipposideros larvatus . Differences in echolocation call frequency among populations did not reflect phylogenetic relationships, indicating that call frequency is a poor indicator of evolutionary history. Instead, divergence in call frequency probably occurs in allopatry, possibly augmented by character displacement on secondary contact to facilitate intraspecific communication.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 119–130.     

扁颅蝠的扩散行为研究

我们于2002 ～ 2007 年在广西龙州县和宁明县采用标记重捕法对扁颅蝠的扩散行为进行研究, 共标记669 只(成体316 只,亚成体353 只;雌雄分别为293 只和376 只),重捕到139 只(重捕率20. 8% )。结果表明,大部分扁颅蝠雌雄后代在性成熟前发生扩散,其亚成体扩散率无性别差异(雄
性82.2% ,雌性66.7% ;P > 0.05)。对成年雌雄两性扩散率(雄性76.5% ,雌性58.5% )的分析亦未见性别差异(P > 0.05),但亚成体和成体合并结果显示雄蝠扩散率(80.7% )高于雌蝠(62.3% ,P < 0.01)。此外,我们还测量了扩散的距离,雄性后代的扩散距离(787. 5 ± 26.980 m,n = 37)比雌
性(517.4 ± 25.308 m,n = 24)远(P < 0. 01);在出现扩散的61 只后代中,仅有一只(0. 7% )雄性亚成体扩散到其它的竹林,其余个体均在出生竹林内的不同竹筒之间进行扩散。     

Auditory sensitivity and frequency selectivity in greater spear-nosed bats suggest specializations for acoustic communication

We investigated the relationship between auditory sensitivity, frequency selectivity, and the vocal repertoire of greater spear-nosed bats (Phyllostomus hastatus). P. hastatus commonly emit three types of vocalizations: group-specific foraging calls that range from 6 to 11 kHz, low amplitude echolocation calls that sweep from 80 to 40 kHz, and infant isolation calls from 15 to 100 kHz. To determine if hearing in P. hastatus is differentially sensitive or selective to frequencies in these calls, we determined absolute thresholds and masked thresholds using an operant conditioning procedure. Both absolute and masked thresholds were lowest at 15 kHz, which corresponds with the peak energy of isolation calls. Auditory and masked thresholds were higher at sound frequencies used for group-specific foraging calls and echolocation calls. Isolation calls meet the requirements of individual signatures and facilitate parent-offspring recognition. Many bat species produce isolation calls with peak energy between 10 and 25 kHz, which corresponds with the frequency region of highest sensitivity in those species for which audiogram data are available. These findings suggest that selection for accurate offspring recognition exerts a strong influence on the sensory system of P. hastatus and likely on other species of group-living bats.     

Flight performance, foraging tactics and echolocation in the trawling insectivorous bat Myotis adversus (Chiroptera: Vespertilionidae)

The bat Myotis adversus hunts for prey by aerial hawking and by taking prey from the water surface with its feet (trawling). The flight performance and echolocation of this species were studied in Queensland, Australia, and comparisons were made with Myotis daubentoni , a bat filling a similar ecological niche in the Palaearctic Region. The bats foraged in very similar ways, using the same foraging tactics and feeding in similar habitats, yet they were not geometrically similar in shape. The slightly larger Myotis adversus had relatively larger wings than M. daubentoni , conferring a slightly lower wing-loading. Nevertheless, M. adversus flew faster than M. daubentoni during the searching phase of foraging. Myotis daubentoni turned in tighter circles than M. adversus . Both species used short frequency-modulated (FM) echolocation calls of a characteristic sigmoidal structure, and nulls typically observed in the calls were an observational artefact. Myotis adversus also adopted an unusual 'long'FM call while foraging. The relations between echolocation frequencies and body size were explored in male M. adversus . Specialized morphological and acoustic adaptations for prey capture by trawling in insectivorous bats are discussed.     

Driving Factors for the Evolution of Species-Specific Echolocation Call Design in New World Free-Tailed Bats (Molossidae)

Phylogeny, ecology, and sensorial constraints are thought to be the most important factors influencing echolocation call design in bats. The Molossidae is a diverse bat family with a majority of species restricted to tropical and subtropical regions. Most molossids are specialized to forage for insects in open space, and thus share similar navigational challenges. We use an unprecedented dataset on the echolocation calls of 8 genera and 18 species of New World molossids to explore how habitat, phylogenetic relatedness, body mass, and prey perception contribute to echolocation call design. Our results confirm that, with the exception of the genus Molossops, echolocation calls of these bats show a typical design for open space foraging. Two lines of evidence point to echolocation call structure of molossids reflecting phylogenetic relatedness. First, such structure is significantly more similar within than among genera. Second, except for allometric scaling, such structure is nearly the same in congeneric species. Despite contrasting body masses, 12 of 18 species call within a relatively narrow frequency range of 20 to 35 kHz, a finding that we explain by using a modeling approach whose results suggest this frequency range to be an adaptation optimizing prey perception in open space. To conclude, we argue that the high variability in echolocation call design of molossids is an advanced evolutionary trait allowing the flexible adjustment of echolocation systems to various sensorial challenges, while conserving sender identity for social communication. Unraveling evolutionary drivers for echolocation call design in bats has so far been hampered by the lack of adequate model organisms sharing a phylogenetic origin and facing similar sensorial challenges. We thus believe that knowledge of the echolocation call diversity of New World molossid bats may prove to be landmark to understand the evolution and functionality of species-specific signal design in bats.     

Differences in songflight calls and social calls between two phonic types of the vespertilionid bat Pipistrellus pipistrellus

The pipistrelle ( Pipistrellus pipistrellus ) occurs as two phonic types in Britain, its echolocation calls falling into two distinct frequency bands, with mean frequencies of maximum energy at 55 kHz and 46 kHz. These are termed the 55 kHz and 45 kHz phonic types here for simplicity. Songflight calls produced by males in the mating season, probably to attract females, differed between the two phonic types in the number of components in the calls and the call parameters measured. Songflight calls of the 55 kHz phonic type, which generally consisted of three components, were of higher frequencies than those of the 45 kHz phonic type, usually of four components. There were also significant differences in call parameters among individuals. A discriminant analysis of songflight calls classified 100% of individuals to the correct phonic type. The relationships between echolocation call frequency and songflight call frequency differed significantly between phonic types. Social calls produced during flight also differed between phonic types, in the number of components and call parameters measured. Social calls were compared to songflight calls of each phonic type. Social calls of the 55 kHz phonic type did not differ significantly from songflight calls; there were small but significant differences between the two types of calls of the 45 kHz phonic type. The study provides support for the hypothesis that the phonic types should be treated as sibling species. If songflight calls are used for mate choice, the differences may allow reproductive isolation between the two phonic types. The functions of songflight calls and social calls need to be investigated through experimental studies to explain the implications of the differences between phonic types.     

Echolocation calls produced by Kuhl's pipistrelles in different flight situations

Flexibility in the echolocation call structure of bats can improve their performances, because, in some situations, some signal designs are better than others. Hence, at least some bats should adjust their echolocation calls according to the setting in which they are operating but also to the specific task at hand, that is their behavioral intention. We studied variation in the echolocation calls of Pipistrellus kuhlii emitted during four flight situations that were similar in setting but differed in behavioral context: emergence from a roost, commuting to and from foraging sites, foraging and returning to a roost. Echolocation calls produced by P. kuhlii differed significantly according to the flight situation. Call types differed most distinctly between foraging and commuting. We also found a high variance in the emergence calls we recorded, perhaps reflecting pre- and post-takeoff calls. Discriminant function analysis on calls emitted while foraging, commuting or returning to the roost classified the calls to the correct group 73.3% of the time. The differences between bats' echolocation calls in different flight situations might indicate an intrinsic change in the bat's behavior. Recognizing these differences could be crucial when using call variables to identify bat species.     

Prey-Correlated Spectral Changes in Echolocation Sounds of the Indian False Vampire Megaderma lyra

False Vampires ( Megaderma lyra ) are gleaning bats which emit brief (1 ms) and faint echolocation signals consisting of four harmonics of a shallow frequency downward modulated fundamental (27–19 kHz). The complete signal spans a frequency range from 100 to 19 kHz. In sound recordings from three experimental animals we show that Megaderma lyra shifts the dominant frequency in the echolocation signals in relation to the type of prey offered and to flight style. During roaming flights the mean peak frequency was 63.2 ± 9 kHz (third harmonic). In prey catching flights, peak frequencies were shifted into the fourth harmonic. In flights towards a dish of crawling mealworms, mean peak frequency was raised to 91.2 ± 3.3 kHz. When the bats flew towards living mice the dominant frequency was further increased to 99.8 ± 5.2 kHz, and the second and third harmonic were at least 10 dB fainter or no longer recordable. The additional frequency shift when flying towards mice was not only a consequence of the dominance of the fourth harmonic but also of an additional rise of the fundamental harmonic by nearly 2 kHz. These prey-correlated frequency shifts in echolocation calls showed little variation between the three experimental animals and were reproducible over time. They occurred at or even before take-off of the bats. This is the first report of target-correlated transient adaptations in echolocation calls of any bat species.