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We recorded bat activity simultaneously at ground level and 30 m height using Anabat II bat detectors and Anabat ZCAIM recording units mounted on masts at seven sites of varying habitat type in eastern England. At the lower detectors 6194 passes were recorded; 90.02% of calls were from Pipistrellus; 6.91 % were from Nyctalus/Eptesicus; 1.70% were Myotis/Plecotus; 0.13% were Barbastella and 1.24% could not be determined to species. At the higher detectors 484 passes were recorded from Pipistrellus (65.08%) and Nyctalus/Eptesicus (34.09%) and 0.83% were not determined to species. Total bat passes and the number of Pipistrellus passes were significantly higher on the lower detectors compared with the upper detectors. The difference between numbers of passes of Nyctalus/Eptesicus at the upper and lower detectors was not significant, despite proportionally more passes of these species being recorded at height. No extra species groups were recorded at height than were recorded at ground level except at one site where closed canopy broadleaved woodland surrounded the mast. Here, Nyctalus/Eptesicus species were recorded only at height. Overall, 28% of Nyctalus/Eptesicus passes and 5% of Pipistrellus passes were recorded exclusively at height, and not at ground level. Assessing bat activity levels from ground level detectors only can therefore be misleading, particularly when surveying high-flying species that are most likely to be at risk from wind turbine developments.
The acoustic structure of echolocation pulses emitted by Japanese pipistrellePipistrellus abramus (Temminck, 1840) bats during different phases of aerial hawking is described here for the first time. Behavioural observations of the foraging flight in conjunction with acoustical analysis of echolocation pulses indicated a flight path consisting of four distinct phases following the reconnaissance or search phase. Short (∼4.68 ms) and relatively broadband frequencymodulated (FM) pulses (∼23.55 kHz bandwidth) were emitted at a repetition rate of 15 Hz during presumed target approach. Presumed insect capture consisted of an early and a late buzz phase. Both buzz types were emitted at high repetition rates (111 Hz in early to 222 Hz in late) and consisted of very short, broadband FM pulses (1.26 ms in early to 0.3 ms in late). There was also a characteristically sharp drop in both the peak and terminal frequencies of each echolocation pulse during the transition from early to late buzz. No pulses were recorded during the final phase of foraging referred to as a “post-buzz pause”. Thus the foraging behaviour of this species consisted of five sequential phases involving four broad types of echolocation pulses.
Researchers often use harp traps and mist nets to capture bats, and need to be aware of factors that affect trap capture efficiency. Ultrasound reflects from small targets in a frequency-dependent manner, so we predicted that higher frequency sound pulses would return stronger echoes from the fine wires and net of bat traps than would lower frequency signals. We also predicted that mist nets would return stronger echoes than harp traps because mist nets are made of a higher density (and often diameter) of material. Ensonification experiments with pulses of 20–110 kHz showed that both harp traps and mist nets reflected higher frequency pulses more strongly. Pocketed areas of mist nets returned stronger echoes than harp traps although at some frequencies differences between trap types were small. Our results provide one verified reason why harp traps are more effective than mist nets at capturing bats, and also predict that bats using high frequency echolocation calls may be more difficult to trap than species emitting low frequency signals. Interspecies differences in how traps are detected are therefore likely sources of bias in field surveys. Observations of bats encountering harp traps in the field showed less than 4% of encounters resulted in capture, and only 8.8% of encounters could be interpreted as a failure to detect the trap. A comparison between two species that differ in echolocation call and flight characteristics (Rhinolophus hipposideros and Myotis nattereri) showed no difference in trap detection or avoidance. However, differences in behaviour during trap encounters were apparent.
Differences in skull morphology between two cryptic species of bat, Pipistrellus pipistrellus (n = 14) and P. pygmaeus (n = 15), originating from Great Britain, were investigated. Four different data sets were analysed: (1) 23 landmarks and (2) 26 landmarks on the dorsal and ventral sides of the cranium, respectively, (3) 49 landmarks on the upper jaw, and (4) 34 landmarks on the labial side of the mandible. For almost all data sets, when compared within sex groups, P. pipistrellus were significantly larger than P. pygmaeus; the biggest difference being observed in the mandible size. Interspecific differences in shape, analysed by Principal Component Analysis and Discriminant Function Analysis (DFA) of the Procrustes superimposed landmarks, were also mostly visible in the mandible, and were related to dietary differences between the species. For example, the longer and more upright canines of P. pipistrellus allow them to pierce harder prey, the bigger molars ease its processing, and the shortened body of the mandible and the more developed coronoid process presumably generate a stronger bite. The shape and size of the mandible proved to be a good characteristic for distinguishing both cryptic taxa. A procedure for estimating missing landmarks for 3D geometric morphometric purposes was created. Our procedure of finding the missing landmarks had no effect on the within-group loss of variation. DFA of data sets with reconstructed versus orginal (but reduced) landmarks yielded similar results (three versus two misclassified specimens in leave-one-out cross-validation).
The meridional serotine Eptesicus isabellinus (Temminck, 1840) has recently been identified as a cryptic species that occurs in the south of the Iberian Peninsula. Little is known about its ecology. We used morphological analysis of droppings from an urban semiarid Mediterranean landscape to determine seasonal differences in diet. We identified 15 insect prey types and found significant seasonal differences in the consumption of seven prey types. The diet of E. isabellinus was dominated by Scarabaeidae (Coleoptera) and Diptera, and there were seasonal changes in the consumption of Carabidae (Coleoptera), Lepidoptera and Cercopidae (Hemiptera). Shannon-Wiener and Levins' indices showed that the diet was more diverse during the post-hibernation and pregnancy periods. Pianka's index showed that there was relatively low dietary overlap between periods. The seasonal changes recorded between different physiological periods could be related to changes in the energetic needs of the individual or to variation in preyavailability. Although E. isabellinus mainly eats Scarabaeidae and Diptera, it can show flexibility for example by hunting insects that fly around blossoms such as chafers Melolontha spp.
We monitored a barbastelle (Barbastella barbastellus) maternity roost for four months using a portable CCTV system, time synchronised with ultrasound recorders. We discovered three patterns of vocal activity not previously described. When barbastelles investigated the roost entrance, calls resembling the approach phase of echolocation were produced consisting of a group of 10 or more broadband pulses of low amplitude, not detected more than a few metres from the roost. The other two distinct vocal patterns were produced during swarming around the roost tree. The first consisted of broadband pulses which were similar to those in the approach pattern, but which tended to be produced in a dynamic group rather than a fixed pattern. They were of higher amplitude than the approach echolocation pulses and were considered to be swarming echolocation pulses. This pattern frequently morphed into sequences containing social calls with a quasi-constant frequency (QCF) tail of very high amplitude. These social calls were likely produced to warn approaching bats on a potential collision course, similar to calls previously described in other species as ‘honking’. In addition to these three vocalisations, standard echolocation pulses were emitted, as well as hooked calls presumably having a social context. These latter two vocalisations were less commonly recorded in our study. Pulse duration was shortest for approach echolocation and longest for the hooked social calls. Amplitude was lowest for approach echolocation and highest for swarming honking calls. The QCF tail in swarming honking was steeper than for swarming social calls. CCTV video evidence (by motion detection) proved invaluable for relating behaviour to call type.
We review the distribution and conservation status of Old World fruit bats (Pteropodidae) in tropical and subtropical China. Our results are based on a series of surveys conducted between 1999 and 2008 by Chinese researchers and international collaborators, spanning almost the entire range of pteropodids within China over nine provinces. Additionally we provide new information on morphometrics and notes on ecology. We also review earlier Chinese literature on fruit bats because much of this has previously been inaccessible to western scientists, and we evaluate the reliability of some of these older records. Thirteen species of fruit bats have been reputed to occur in China, including one species restricted to Taiwan. We classified Chinese fruit bats according to distribution and status as follows: Resident (six species, including one species formerly found in Taiwan and now restricted to its neighbouring islands), Marginal (three species), Questionable (one species) and Alien (three species). Consequently, only five species are encountered with any regularity in mainland China and Hainan Island: Cynopterus sphinx, Eonycteris spelaea, Macroglossus sobrinus, Rousettus leschenaultii, and Sphaerias blanfordi. Three species in the genus Pteropus are referred to in old records, but these bats are clearly not native to mainland China.
The low- and high-frequency components of a rustling sound, created when prey (freshly killed frog) was jerkily pulled on dry and wet sandy floors and asbestos, were recorded and played back to individual Indian false vampire bats (Megaderma lyra). Megaderma lyra responded with flight toward the speakers and captured dead frogs, that were kept as reward. The spectral peaks were at 8.6, 7.1 and 6.8 kHz for the low-frequency components of the sounds created at the dry, asbestos and wet floors, respectively. The spectral peaks for the high-frequency sounds created on the respective floors were at 36.8, 27.2 and 23.3 kHz. The sound from the dry floor was more intense than that of from the other two substrata. Prey movements that generated sonic or ultrasonic sounds were both sufficient and necessary for the bats to detect and capture prey. The number of successful prey captures was significantly greater for the dry floor sound, especially to its high-frequency components. Bat-responses were low to the wet floor and moderate to the asbestos floor sounds. The bats did not respond to the sound of unrecorded parts of the tape. Even though the bats flew toward the speakers when the prey generated sounds were played back and captured the dead frogs we cannot rule out the possibility of M. lyra using echolocation to localize prey. However, the study indicates that prey that move on dry sandy floor are more vulnerable to predation by M. lyra.
The development of vocalizations during postnatal growth in the flat-headed bats, Tylonycteris pachypus and T. robustula in South China is described. Females of both species gave birth to twins at the end of May, and the infants flew in the last ten days of June. Vocalizations served as precursors to echolocation calls and as isolation calls (i-calls) used to attract mothers. As the infants grew, the frequency of i-calls and precursor calls increased. The duration of i-calls increased little before 6-day old and then decreased. At the same time, the duration of echolocation precursor calls decreased. The directive calls that the mother or the infant emitted when searching for each other are also described. Female directive calls are lower in frequency and longer in duration than their echolocation calls, and the duration of infant directive calls is longer than those of the i-calls and precursor calls.
We describe the echolocation calls, flight morphology and diet of the endemic Chinese bat Myotis pequinius Thomas, 1908. Orientation calls are broadband, and reach low terminal frequencies. Diet comprised 80% beetles by volume. Wing shape and call design suggest that the bats fly in cluttered habitats, and the possession of moderately long ears and the dietary composition imply they forage at least sometimes by gleaning. Myotis pequinius resembles a larger Oriental version of the western Palaearctic species M. nattereri. Phylogenetic analysis based on sequences of the cytochrome b gene of mitochondrial DNA (1,140 base pairs) from a range of Palaearctic Myotis species confirmed that M. pequinius is close to the nattereri group, and is a sister-species to the eastern Palaearctic M. bombinus. One bat sequenced from China could not be identified from available species descriptions. It was smaller than M. pequinius, and also differed from it in sequence divergence by 6.7%, suggesting the existence of additional, cryptic taxonomic diversity in this group. Our phylogenetic analysis also supports the recognition of M. schaubi as a species distinct from M. nattereri in Transcaucasia and south-western Asia. Myotis nattereri tschuliensis is more closely related to M. schaubi than to M. nattereri, and is best considered either as a subspecies of M. schaubi, or possibly as a distinct species.
Variation in the acoustic structure of bat echolocation calls can often provide sufficient information for reliable and efficient species identification. The aim of this study was to investigate the use of echolocation call structure to identify a number of bats in the families Vespertilionidae, Emballonuridae, Nycteridae and Megadermatidae from Thailand. These species typically emit echolocation calls with a frequency-modulated (FM) sweep dominating part or all of their calls. A total of 510 echolocation calls from free-flying individuals were recorded throughout Thailand. According to the frequency-time spectra, these calls were categorized into four types: broadband FM (eight species), narrowband FM (seven species), long multiharmonic (four species) and short multiharmonic (three species). Discriminant function analysis was used to classify calls from individual bats to species. Correct classification levels were 85.9% for individuals emitting broadband FM calls (six species with adequate sample sizes), 70.4% for narrowband FM bats (seven species), 84.4% for species emitting long multiharmonic calls (four species) and 96.7% for species emitting short multiharmonic calls (two species with adequate sample sizes). However, classification rates were often low at the species level. Acoustic identification of bats emitting FM calls should be approached with caution in species-rich communities, in contrast with the identification of rhinolophoid bats where many species use distinctive constant frequencies that can facilitate identification, and provides a basis for rapid acoustic surveys of large areas in Thailand, and potentially other parts of Southeast Asia.
We conducted surveys of bats in China between 1999 and 2007, resulting in the identification of at least 62 species. In this paper we present data on 19 species, comprising 12 species from the family Rhinolophidae and seven from the Hipposideridae. Rhinolophids captured were Rhinolophus affinis, R. ferrumequinum, R. lepidus, R. luctus, R. macrotis, R. siamensis, R. marshalli, R. rex, R. pearsonii, R. pusillus, R. sinicus and R. stheno. Because of extensive morphological similarities we question the species distinctiveness of R. osgoodi (may be conspecific with R. lepidus), R. paradoxolophus (which may best be treated as a subspecies of R. rex), R. huananus (probably synonymous with R. siamensis), and we are skeptical as to whether R. sinicus is distinct from R. thomasi. Hipposiderids captured were Hipposideros armiger, H. cineraceus, H. larvatus, H. pomona, H. pratti, Aselliscus stoliczkanus and Coelops frithii. Of these species, two rhinolophids (Rhinolophus marshalli and R. stheno) and one hipposiderid (Hipposideros cineraceus) represent new species records for China. We present data on species’ ranges, morphology and echolocation call frequencies, as well as some notes on ecology and conservation status. China hosts a considerable diversity of rhinolophid and hipposiderid bats, yet threats to their habitats and populations are substantial.
We studied the chiropteran diversity of Andaman Islands between July 2012 and January 2016 from 38 different localities spread throughout the islands. Our surveys revealed the presence of 17 species of bats. One species — lesser bamboo bat Tylonycteris pachypus — reported earlier could not be found during the present study. Our study adds four new species records (Andersen's roundleaf bat Hipposideros pomona, diadem roundleaf bat Hipposideros diadema, Indian pipistrelle Pipistrellus coromandra, and greater bamboo bat Tylonycteris robustula) for the Andaman Islands. A brief review of each species including details on its distribution (earlier and present), and external and craniodental measurements is provided. We also provide a database of echolocation calls for the Islands' echolocating bat fauna, and confirm the reliability of acoustic monitoring as a method to noninvasively document chiropteran diversity there. This database allows comparisons with taxa from the mainland that are currently considered conspecific, but which show acoustic divergence, and deserve further taxonomic study.
Bats are an important component of biodiversity in Southeast Asia, and are key indicators of habitat quality. Acoustic analysis of echolocation calls not only allows the identification of bat species that are difficult to capture, but also allows for rapid and standardised ways to survey and monitor bats over large areas. However keys based on call parameters must also account for geographic variation in call parameters, and consider any effects of morphology and sex on call frequency. Presence-only modelling can predict likely geographic locations of specific taxa, and can used to refine decision making so that species unlikely to occur in a specific region can be omitted from more localised acoustic libraries. Here we develop an acoustic library for the echolocation calls of rhinolophoid bats in Thailand, and use presence-only modelling (Maxent) to explore how species with similar calls in a library developed at the national level can sometimes be largely allopatric, and hence identifiable, once geographic range is predicted. Presence-only modelling can also be used to explore whether species with similar calls adjust call frequency in likely areas of sympatry. We analysed calls from fourteen species of rhinolophid and twelve hipposiderid species from Thailand. Calls from a further three rhinolophid and one hipposiderid species are also described but not analysed statistically because of small sample sizes. Even without considering geographic variation, 69% of rhinolophid (14 species with a minimum of five individuals/ species) and 91% of hipposiderid calls (12 species) could be classified successfully to species using two call parameters (frequency of most energy (FMAXE) and duration) in a discriminant function analysis. Most of the discrimination was achieved because species often utilised different frequency bands. Morphology can also affect call frequency both across and within species. In both rhinolophids and hipposiderid families there was a negative relationship between FMAXE and forearm length. Within species, FMAXE was negatively related to forearm length in Rhinolophus microglobosus, R. pusillus and R. thomasi, and positively related to forearm length in R. affinis and R. pearsonii. Male R. pusillus called at higher frequencies than females, although there was no sexual size dimorphism in forearm length. Call frequency was negatively related to relative humidity in R. pusillus, suggesting that bats called at lower frequencies in humid environments to counter increases in atmospheric attenuation. Presence-only modelling was used to show that some species with similar call frequencies (e.g., R. lepidus and R. microglobosus; R. yunanensis and R. trifoliatus are predicted to occur largely in allopatry, and so could be identified reliably in particular parts of the country. Presenceonly modeling can assist in predicting areas of overlap between species with similar call frequencies. Other species (e.g., R. malayanus, R. coelophyllus) overlap in frequency when data from all of Thailand are combined, but seem to avoid call overlap when syntopic. Hence acoustic identification can be improved by taking into account local distribution patterns and patterns of species coexistence. The creation of call libraries at a local scale would have extensive potential as a resource to monitor changes in species distributions through time.
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