Anim. Syst. Evol. Divers. Vol. 31, No. 3: 160-175, July 2015 http://dx.doi.org/10.5635/ASED.2015.31.3.160
Echolocation Call Structure of Fourteen Bat Species in Korea
Dai Fukui1,2,*, David A. Hill3,4, Sun-Sook Kim2, Sang-Hoon Han2
1The University of Tokyo Hokkaido Forest, Graduate School of Agricultural and Life Sciences, The University of Tokyo, Hokkaido 079-1563, Japan 2Division of Animal Resources, National Institute of Biological Resources, Incheon 404-708, Korea 3Wildlife Research Center, Kyoto University, Kyoto 606-8203, Japan 4Primate Research Institute, Kyoto University, Aichi 484-8506, Japan
ABSTRACT
The echolocation calls of bats can provide useful information about species that are generally difficult to observe in the field. In many cases characteristics of call structure can be used to identify species and also to obtain information about aspects of the bat’s ecology. We describe and compare the echolocation call structure of 14 of the 21 bat spe cies found in Korea, for most of which the ecology and behavior are poorly understood. In total, 1,129 pulses were analyzed from 93 echolocation call sequences of 14 species. Analyzed pulses could be classified into three types according to the pulse shape: FM/CF/FM type, FM type and FM/QCF type. Pulse structures of all species were con sistent with previous studies, although geographic variation may be indicated in some species. Overall classification rate provided by the canonical discriminant analysis was relatively low. Especially in the genera Myotis and Murina, there are large overlaps in spectral and temporal parameters between species. On the other hand, classification rates for the FM/QCF type species were relatively high. The results show that acoustic monitoring could be a powerful tool for assessing bat activity and distribution in Korea, at least for FM/QCF and FM/CF/FM species. Keywords: bats, Chiroptera, Korean peninsula, echolocation call, interspecific variation
INTRODUCTION ing echolocation call structure can provide insights into the ecology of species that are otherwise difficult to study. Bats are the most unusual and specialized of all terrestrial Recently the use of ultrasonic bat detectors to make acous mammals. They are the only extant vertebrates apart from tic surveys of bat activity has been rapidly increasing (e.g., birds that are capable of powered flight and, with the excep Racey et al., 1998; Loeb and O’Keefe, 2006; Morris et al., tion of Pteropodidae species, have mastered the night skies 2010; Minderman et al., 2012; Stahlschmidt et al., 2012). largely by using echolocation (biosonar) to perceive their Acoustic survey can be especially useful in habitats where surroundings (Griffin, 1958; Thomas et al., 2004). Echoloca capture is generally difficult, such as large open fields and tion calls show a great diversity in form, duration and ampli high in the forest canopy (Kunz et al., 2009). Also, acoustic tude, and key characteristics are correlated with the environ surveys can be carried out at multiple sites simultaneously by ment in which the bat species generally flies (Jones and Teel using automated recording systems. To conduct such acous ing, 2006). Typically, bat species which forage in an open tic surveys, however, it is necessary to have a reference col habitat will emit a low-frequency ultrasonic call that will lection of calls of known bat species and then reliable meth not rapidly attenuate, with a long duration and a narrow- ods should be developed to identify the resident species. frequency bandwidth. By contrast, bat species foraging in The structure of echolocation calls varies, not only bet dense vegetation (i.e., a cluttered habitat) will emit short ween species, but also within species. A variety of factors broadband calls, which are suited to localizing prey in clut have been shown to influence call structure, including geo ter (Schnitzler et al., 2003; Jones, 2005). Therefore, describ graphical variation, foraging habitat and foraging mode (Al
This is an Open Access article distributed under the terms of the Creative *To whom correspondence should be addressed Commons Attribution Non-Commercial License (http://creativecommons.org/ Tel: 81-167-42-2111, Fax: 81-167-42-2689 licenses/by-nc/3.0/) which permits unrestricted non-commercial use, distribution, E-mail: [email protected] and reproduction in any medium, provided the original work is properly cited. pISSN 2234-6953 eISSN 2234-8190 Copyright The Korean Society of Systematic Zoology Echolocation Calls of Korean Bats dridge and Rautenbach, 1987; Norberg and Rayner, 1987; Thomas et al., 1987; Fenton, 1990; Barclay and Brigham, 1991; Obrist, 1995; Barclay et al., 1999). This means that reference calls recorded in a particular region, or in a parti cular habitat type, may not be applicable to other regions, or to other habitat types. Therefore, as far as possible, echo location call structure should be described and species iden tification methods developed for the particular region and habitat type where they are to be used. Numerous previous studies have analyzed and described echolocation call struc ture of a variety of bat species, although the majority has been done in Europe and North America (e.g., Russo and Jones, 2002; Rydell et al., 2002; Redgwell et al., 2009; Brit zke et al., 2011). The Korean Peninsula is an important area to consider in biogeographical studies of East Asian mammals. At least 21 bat species are known to be distributed in Korea (Yoon, 2010). However, the ecology and behavior of Korean bats are almost unknown, except for some cave- and house-dwel ling species (Chung et al., 2009, 2010b, 2011; Kim et al., 2009, 2012). Also, although Chung et al. (2010a, 2010c) have described the echolocation call structure of three Kor ean bat species, the echolocation call structure of most Kor ean bat species has not been analyzed or described yet. The aim of this study is to describe the echolocation call structure of 14 of the 21 Korean bat species. In this paper, we com pare characteristics of echolocation call structure between Fig. 1. Map of the locations where bats were captured or re- species. This is the first study to present an analysis of inter corded for this study. specific differences in echolocation calls of Korean bats.
had been included in My. daubentonii in Asia, we follow MATERIALS AND METHODS Matveev et al. (2005), who proposed it as a valid species for the “eastern” group of My. daubentonii. Yoon (2010) has From 2011 to 2012, fourteen species of bats (Eptesicus sero described My. nattereri from Korea, but the east Asian popu tinus, Hypsugo alaschanicus, Miniopterus fuliginosus, Mur lation is now considered to be My. bombinus (Simmons, ina leucogaster, Mu. ussuriensis, Myotis aurascens, My. for 2005). Although H. (Pipistrellus) alaschanicus had been mosus, My. ikonnikovi, My. macrodactylus, My. bombinus, treated as a subspecies of H. (Pipistrellus) savii, we follow My. petax, Pipistrellus abramus, Plecotus ognevi, and Rhi Horácek et al. (2000), who argued that H. alaschanicus from nolophus ferrumequinum) were caught from a range of loca Mongolia, China, the Russian Far East, Korea and Japan, tions throughout the country (Fig. 1), using mist nets and should be recognized as a valid species of the genus Hyp harp traps in forests, or using a hand net at day or night roosts sugo (Pipistrellus) and different from H. savii. Although (caves and houses). After capturing bats, we checked their Yoon (2010) treated the bent-winged bat from Korea as Mi. sex, maturity and reproductive condition, and measured schreibersii, we follow Tian et al. (2004), who proposed the their forearm length using calipers and body weight using a Asian population as a separate species, Mi. fuliginosus. In balance. In this study scientific names follow Yoon (2010), addition, although P. auritus is described in Yoon (2010), except for My. aurascens, My. bombinus, My. petax, H. ala recent genetic and morphological analyses indicate that the schanicus, Mi. fuliginosus, and P. ognevi. The taxonomic Far Eastern population is a distinct species P. ognevi (Spit status of Korean small-sized Myotis bats is not well studied. zenberger et al., 2006). Although My. aurascens is not described in Yoon (2010), we follow Tsytsulina et al. (2012), who have suggested the Call recording occurrence of My. aurascens in Korea. Although My. petax Calls were recorded using a Pettersson D240x bat detector
Anim. Syst. Evol. Divers. 31(3), 160-175 161 Dai Fukui, David A. Hill, Sun-Sook Kim, Sang-Hoon Han
(Pettersson Elektronik AB, Uppsala, Sweden) linked to a measured the following parameters from each pulse: dura solid state recorder (R-05; Roland, Shizuoka, Japan). The tion (D), maximum frequency (FMAX, the highest frequency D240x was set to time-expansion mode, in which it records of the pulse), minimum frequency (FMIN, the lowest fre 3.4 s of ultrasound, slows the signal down ten times to bring quency of the pulse), peak frequency (PF, frequency of max it into the audible range, and plays it back. This technique imum energy of the pulse), and interpulse interval (IPI, time preserves all characteristics of the original sound and allows from the beginning of measured pulse to the beginning of accurate measurement of acoustic parameters. The record next pulse). D, FMAX, FMIN, and IPI were measured from ings were made in the following four situations. (1) Hand- spectrograms, and PF from power spectra. release: Captured bats except all individuals of Rhinolophus ferrumequinum and some individuals of other species (see Statistical procedures below) were released by hand on a forest road (width >5 m) If calls of the species were recorded in more than two situa near the point of capture, and the calls produced from 1-2 tions, the differences in call parameters between recording seconds after release were recorded. Recordings were made situations (hand-release, flight room, free flight and perch) with the detector held at the same height as, and about 2 m were analyzed using linear mixed models (LMMs). In the away from the released bat. (2) Free-flight: If we were able analyses, the recording situation was the fixed effect, and the to make positive species identification of flying bats (e.g., call sequence was treated as a random effect to account for checking individuals at a night roost, or individuals that went potential differences among individuals. A value of p<0.05 into mist nets after observed flight), we recorded echoloca was considered statistically significant. Next, we compared tion calls during their flight. (3) Flight room: Some bats were the pulse structures between each species by canonical dis brought back to the laboratory to be prepared as specimens. criminant analysis (CDA) using all five parameters. In the Before making specimens, these individuals were released in CDA, the last pulse in each call sequence was excluded a flight room (4.7 m length, 3.0 m width, and 2.7 m height) because it did not have a following call, and therefore there in National Institute of Biological Resources (NIBR), and was no IPI (as defined above). All analyses were performed echolocation calls were recorded during their flight inside in the R environment for statistical computing (R Develop the room. (4) Perch: For R. ferrumequinum, perch calls were ment Core Team, 2012) and its extended nlme and MASS recorded when the bats hung from the roof of a mesh bag (40 packages (Pinheiro et al., 2012). cm square by 40 cm height).
Call analysis RESULTS The structure of recorded calls was analyzed using Bat Sound 3.1 software (Pettersson Elektronik AB) with a sampling rate In total, 93 echolocation call sequences were recorded from of 44.1 kHz and a Hanning window. Fast Fourier Transform 93 individuals of 14 species (Table 1). From these 93 call size was 512 for sonograms and power spectra. All pulses in sequences, 1,129 pulses were analyzed. Descriptive statis each 3.4 s recorded call (call sequence) were analyzed. We tics of calls from all species are shown in Appendix 1. Analy
Table 1. Species recorded and number of individuals in each recording situation Recording situations Free flight Hand-released Flight room Perch Rhinolophus ferrumequinum 12 0 0 25 Myotis aurascens 0 2 2 0 Myotis formosus 0 3 0 0 Myotis ikonnikovi 0 7 5 0 Myotis macrodactylus 0 3 0 0 Myotis bombinus 0 3 0 0 Myotis petax 0 2 0 0 Plecotus ognevi 0 0 3 0 Murina ussuriensis 0 2 3 0 Murina leucogaster 0 6 0 0 Eptesicus serotinus 0 3 0 0 Hypsugo alaschanicus 1 3 0 0 Miniopterus fuliginosus 1 4 0 0 Pipistrellus abramus 0 3 0 0
162 Anim. Syst. Evol. Divers. 31(3), 160-175 Echolocation Calls of Korean Bats
Fig. 2. Sonograms of 3 species of bats in Korea. For Myotis aurascens, two types of calls recorded in different situations (flight room and hand-released) are shown. zed pulses could be classified into the following three types tant-frequency (CF) component preceded and followed by according to the pulse shape. a brief, frequency-modulated (FM) sweep (Fig. 2). Mean bandwidth was 4.7-19.5 kHz. Calls of the individuals from FM/CF/FM type species Jeju island had significantly higher FMAX and PF than those Rhinolophus ferrumequinum produced typical FM/CF/FM from the peninsula (LMMs, FMAX: t= -9.29, p<0.05; echolocation pulses, i.e., pulses with a long, strictly cons PF: t= -12.22, p<0.05) (Appendix 1). The only significant
Anim. Syst. Evol. Divers. 31(3), 160-175 163 Dai Fukui, David A. Hill, Sun-Sook Kim, Sang-Hoon Han
Fig. 3. Sonograms of 4 species of bats in Korea. difference between the free-flight and perch calls was in D 60.0 kHz (Figs. 2-4, Appendix 1). Pulses emitted by the (LMM, t= -2.51, p=0.02). genus Myotis had broad bandwidth (>50 kHz). Pulses emit ted by My. bombinus had particularly broad bandwith (71.4- FM type species 113.1 kHz) (Fig. 3, Appendix 1) similar to those of genus Mu Pulses of nine species of the genera Murina, Myotis, and rina. My. formosus and My. ikonnikovi also emitted pulses Plecotus were classified as this call type. All six Myotis spe with relatively broader bandwidth (71.6-87.7 kHz and 56.8- cies captured in this study had similar mean PF (about 40.0- 92.9 kHz, respectively) (Figs. 2, 3, Appendix 1). D for these
164 Anim. Syst. Evol. Divers. 31(3), 160-175 Echolocation Calls of Korean Bats
Fig. 4. Sonograms of 4 species of bats in Korea. species was short (1.7-5.9 ms) but slightly longer than for type species (38.5-39.2 kHz) (Appendix 1). Pulses emitted the genus Murina or Plecotus. For pulses of My. aurascens by this species had shorter bandwidth (22.4-27.0 kHz) and and My. ikonnikovi, which were recorded in two situations a distinct harmonic (Fig. 4). The mean PF of Murina ussuri (flight room and hand release), there were no significant ensis was the highest among those of the nine FM type spe differences between situations in any parameter except for cies (70.0-82.9 kHz) (Appendix 1). Although it was variable, bandwidth of My. ikonnikovi (LMM, t=2.27, p=0.047). the mean PF of Murina leucogaster was also relatively high Plecotus ognevi had the lowest mean PF of the nine FM er (45.7-67.9 kHz) than other FM type species (Appendix
Anim. Syst. Evol. Divers. 31(3), 160-175 165 Dai Fukui, David A. Hill, Sun-Sook Kim, Sang-Hoon Han
Fig. 5. Sonograms of 4 species of bats in Korea. For Miniopterus fuliginosus, two types of calls recorded in different situations (free flight and hand-released) are shown.
1). D for both species was relatively short (1.6-4.7 ms in FM/QCF type species Mu. ussuriensis and 1.9-3.5 ms in Mu. leucogaster), and Calls of Eptesicus serotinus, Hypsugo alaschanicus, Miniop bandwidth was broad (67.9-85.4 kHz and 67.3-102.4 kHz, terus fuliginosus, and Pipistrellus abramus were all defined respectively) (Fig. 3, Appendix 1). There were no significant as FM/QCF type (Figs. 4, 5). These pulses had two compo differences in call parameters of Mu. ussuriensis between nents: they began with steep FM, and moved to shallow FM different recording situations. (quasi-constant frequency, QCF) in the latter part of the
166 Anim. Syst. Evol. Divers. 31(3), 160-175 Echolocation Calls of Korean Bats
Table 3. Coefficients of first 3 canonical discriminants of classi- fication of 14 species by linear discriminant analysis 0 0 0 0 0 0 1 0 0 0 0 0 17 19 37 19 51.4 mus
Pi. abra CD1 CD2 CD3 FMAX - 0.0210 0.0293 - 0.0052 FMIN 0.1459 0.0492 - 0.1648 0 0 0 0 0 0 0 0 0 0 0 0 67 3 70 67 95.7 nosus PF 0.0366 0.0597 0.0291 Mi. fuligi D 0.0647 0.0182 0.1115 IPI - 0.0079 0.0014 - 0.0035 Bandwidth - 0.0425 0.0281 0.0143
0 0 0 0 0 0 0 4 0 0 1 23 0 0 28 23 82.1 Proportion of trace (%) 79.1 13.2 5.2 nicus H. alascha Model relied on 5 parameters (FMAX, FMIN, PF, D, and IPI). FMAX, maximum frequency; FMIN, minimum frequency; PF, frequency of maximum energy; D, duration; IPI, inter-pulse interval. tinus 0 0 0 0 0 0 0 0 0 0 36 0 0 0 36 36 100.0 E. sero
pulse. Pulses of these species had relatively longer D and
0 0 5 24 6 7 0 1 3 36 0 0 0 0 82 36 45.1 shorter bandwidth than the FM type species. Among these gaster Mu. leu co species, pulses of E. serotinus had the lowest mean PFs (29.7 -31.6 kHz). D and bandwidth of E. serotinus were 8.0-8.9 ms and 32.7-42.3 kHz, respectively (Appendix 1). H. ala 0 0 0 19 0 0 0 0 48 0 0 0 6 0 73 48 65.8
riensis
Mu. ussu Mu. schanicus also had lower mean PFs (34.0-36.2 kHz) than all other species except for E. serotinus. D and bandwidth of H. alaschanicus were 2.4-7.8 ms and 15.6-34.2 kHz, respecti
0 0 0 0 0 0 0 49 0 0 0 0 0 0 49 49 100.0 vely (Appendix 1). Hand-released calls of H. alaschanicus Pl. ognevi had significantly higher PF than free-flight calls (LMM, t= 4.40, p=0.048), whereas there were no differences in other True species True 0 1 0 11 10 0 7 0 0 0 0 1 1 0 31 7 22.6 parameters. The mean PFs, Ds, and bandwidths of the pulses My. petax My. from Mi. fuliginosus were 50.3-54.2 kHz, 2.9-7.2 ms, and 9.4-37.6 kHz, respectively (Appendix 1). Free-flight calls
0 1 0 0 0 46 0 0 0 1 3 0 0 0 51 46 90.2 of M. fuliginosus had significantly longer D and IPI than binus My. bom hand-released calls (LMMs, D: t= -3.53, p=0.038; IPI: t= -6.74, p=0.007). The mean PFs for Pi. abramus were slightly lower than those for Mi. fuliginosus (47.0-48.1 kHz),
0 3 0 2 40 0 4 0 0 0 0 0 1 1 51 40 76.5
dactylus whereas there were no distinct differences in Ds (5.2-6.0 ms) My. macro My. or bandwidths (19.9-30.0 kHz) (Appendix 1).
CDA nikovi 0 20 0 154 1 0 4 0 1 7 5 0 0 0 192 154 79.7 My. ikon My. CDA using 5 parameters resulted in 76.4% of 1,036 records of echolocation pulses being correctly classified to one of
0 0 23 26 0 0 0 0 0 11 0 0 0 0 60 23 38.3 mosus the 16 species (Table 2). The first 3 discriminant functions My. for My. explained 97.5% of total variation (Table 3). Pulses from three species (Eptesicus serotinus, Plecotus ognevi, and Rhi 0 15 1 9 11 0 1 0 0 5 4 0 0 0 46 15 32.6
scens nolophus ferrumequinum) were all classified correctly, and My. aura My. pulses from Miniopterus fuliginosus and Myotis bombinus were classified with more than 90% accuracy. On the other hand, pulses from My. petax were the least accurately classi
230 0 0 0 0 0 0 0 0 0 0 0 0 0 230 230 100.0 quinum fied (22.6%), and were mainly misclassified as being from R. ferrume R. My. ikonnikovi and My. macrodactylus. Pulses from Murina leucogaster, My. aurascens and My. formosus were also poorly classified (45.1%, 32.6%, and 38.3%, respectively). In the canonical score plot, the cluster of R. ferrumequinum was distinct from clusters of other species (Fig. 6). There R. ferrumequinum R. aurascens My. formosus My. ikonnikovi My. macrodactylus My. bombinus My. petax My. Pl. ognevi ussuriensis Mu. leucogaster Mu. serotinus E. H. alaschanicus Mi. fuliginosus Pi. abramus n n correct correct % Classified as
Summary of classification of 16 species by canonical discriminant analysis Table 2. Summary of classification 16 species 76.4%. was rate correct classification Overall IPI). and D, PF, FMIN, (FMAX, parameters 5 on Model relied interval. inter-pulse IPI, duration; D, energy; maximum of frequency PF, frequency; minimum FMIN, frequency; FMAX, maximum was considerable overlap among the clusters of the Myotis
Anim. Syst. Evol. Divers. 31(3), 160-175 167 Dai Fukui, David A. Hill, Sun-Sook Kim, Sang-Hoon Han
Eptesicus serotinus 6 Hypsugo alaschanicus Miniopterus fuliginosus Murina leucogaster Murina ussuriensis Myotis aurascens Myotis formosus 4 Myotis ikonnikovi Myotis macrodactylus Myotis bombinus Myotis petax Pipistellus abramus Plecotus ognevi 2 Rhinolophus ferrumequinum CD2
0
-2
-4
-5 0 5 10 CD1
Fig. 6. Individual scores on canonical discriminant functions 1 and 2. species, although that of My. bombinus was relatively dis versen, 1989). In East Asia, the PF of this species also decre tinct from other species. The clusters of FM/QCF type spe ases from west to east (Beijing-China: 75.1 kHz, Jilin Pro cies had relatively little overlap with each other. vince-China: 69.6 kHz, Kyushu-Japan: 67-69 kHz, western Honshu-Japan: 67-68 kHz, eastern Honshu-Japan: 65-67 kHz, Hokkaido-Japan: 63-66 kHz) (Huihua et al., 2003; DISCUSSION Fukui et al., 2004; Matsumura, 2005; Sun et al., 2008), so shows the same trend at a smaller spatial scale. The PF of Of the 14 bat species treated in this study, call structures of this species from the Korean peninsula in this study (67.8- 11 species were described from Korea for the first time. The 69.5 kHz) (Appendix 1) is consistent with this trend. How pulse parameters measured for each species were generally ever, we found that the Jeju population of R. ferrumequinum consistent with those of other studies in surrounding regions, has distinctively higher PF (72 kHz) (Appendix 1) than pop such as China and Japan, whereas some species seem to ulations in surrounding regions such as Kyushu and the Kor show geographical variation in pulse parameters. ean Peninsula. This phenomenon may be due to the isola Rhinolophus ferrumequinum was the only species in this tion effect. In fact, the PF of R. ferrumequinum (69-70 kHz) study with FM/CF/FM type calls. The pulse structures of this (Matsumura, 2005) from Tsushima island, which is also iso species have been described previously for other regions lated from the continent and Japanese main islands, is higher (Vaughan et al., 1997; Parsons and Jones, 2000; Russo and than surrounding populations. Yoshino et al. (2008) revealed Jones, 2002; Huihua et al., 2003; Fukui et al., 2004; Sun that restricted dispersal of females causes echolocation call et al., 2008), and our results agree with those of a previous frequency variation within rhinolophid bats between regions study of this species in Korea (Chung et al., 2010c). It is because the CF component is determined by mother-off known that PF of R. ferrumequinum shows a geographic spring transmission. Further phylogenetic analysis in east cline along which frequency decreases continuously from Asia is needed to confirm the isolation effect. west to east in its distributional range (Heller and von Hel Pulse characteristics of all six Myotis species were typical
168 Anim. Syst. Evol. Divers. 31(3), 160-175 Echolocation Calls of Korean Bats
FM calls that have been described for Myotis species by pre other two species seem to show variation between studies. vious studies (e.g., Russo and Jones, 2002). Our results sug For example, PF of Eptesicus serotinus varies with regions gest that all Myotis species in this study seem to be “narrow (U.K.: 33.6 kHz, Switzerland: 26.8 kHz, Italy: 29.9 kHz, space gleaning foragers” or “edge space aerial/trawling fora Greece: 31.7 kHz) (Parsons and Jones, 2000; Russo and gers”, following the call categorisation described by Schnitz Jones, 2002; Obrist et al., 2004; Papadatou et al., 2008). ler et al. (2003). In fact, all individuals of Myotis were caught Also, the PF of Miniopterus fuliginosus was different from at the forest edge or interior, and we had frequently observed that of Japanese Mi. fuliginosus (47.2-49.8 kHz) (Funakoshi, “Myotis-like” bats flying above water surfaces (D. Fukui, 2010). It is possible that there are geographic variations in personal observation). Of the six species, pulse structures of the pulse frequency. We cannot determine the extent to five (My. formosus, My. ikonnikovi, My. macrodactylus, My. which these differences represent geographic variation, how bombinus, and My. petax) have been described in previous ever, because recording conditions may have varied between studies (e.g., Fukui et al., 2004, 2007; Sun et al., 2008), and studies. It is known that echolocation pulse parameters cha our results are all consistent with these studies. However, nge with flying situation (Schnitzler and Kalko, 2001), and we cannot assess geographical variation for Myotis species our results showed differences in some pulse parameters because measured parameters of these species showed large of H. alaschanicus and Mi. fuliginosus between recording intra-specific variation (Appendix 1), as has been noted by situations. To reveal geographic variations in the calls of other studies (Fukui et al., 2004; Obrist et al., 2004). FM/QCF and FM type species, we would need to compare Pulse characteristics of Plecotus ognevi were similar to calls of each population that were recorded under the same those of Pl. auritus from Europe, and of other Plecotus spe conditions. cies. The PFs for Pl. ognevi in this study were 38.5-39.6 In this study, the overall classification rate provided by the kHz (Appendix 1), whereas those of Pl. auritus from British, CDA was similar to or lower than previous studies where Swiss and Italian populations are 29.9, 37.7, 33.1 kHz, res discriminant analysis was applied (e.g., Parsons and Jones, pectively (Parsons and Jones, 2000; Russo and Jones, 2002; 2000; Russo and Jones, 2002; Fukui et al., 2004; Papadatou Obrist et al., 2004). In Japan, the mean PF of Pl. sacrimon et al., 2008). This result may have been caused by low clas tis, which was treated as Pl. auritus before Spitzenberger et sification rates of FM type species, apart from Myotis bom al. (2006), is 36.7 kHz (Fukui et al., 2007). Recent genetic binus, whose calls are identifiable because of their broad and morphological analyses indicate that the east Asian pop bandwidth. Generally, echolocation calls from most Myotis ulations should be divided into several species, and the Kor and Murina species show similar pulse structures and large ean population is considered to be Pl. ognevi (Spitzenberger overlap in spectral and temporal parameters (Vaughan et al., et al., 2006). Thus, this variation in mean PF may represent 1997; Russo and Jones, 2002; Hughes et al., 2011). Eight inter-specific variation, although recording situation may of the 14 species in this study were Myotis or Murina spe also have influenced the results. cies. On the other hand, the CDA showed relatively high Pulse characteristics of both Murina species in this study classification rates for the FM/QCF type species, except for were similar to those described for three Murina species in Pipistrellus abramus. Previous studies have also shown high Malaysia (Kingston et al., 1999) and two in Japan (Fukui classification rates on the FM/QCF type species (Parsons et al., 2004). Short, broad-band and low intensity FM calls and Jones, 2000; Russo and Jones, 2002; Papadatou et al., are thought to be used for the detection of arthropod prey in 2008; Hughes et al., 2011). This may be related to the QCF clutter (Simmons et al., 1979; Neuweiler, 1989; Schnitzler component in the pulse. In general, the QCF component et al., 2003). Echolocation calls of Murina in this study area seems to be less variable than the FM component in spectral would be suitable for highly cluttered forest understory. Al parameters. Coefficients of first canonical discriminants in most all individuals in this study were caught inside forests dicate that the most important variable in the model was (D. Fukui, personal observation). As with the genus Myotis, FMIN, which is measured from the QCF component of the it is difficult to compare between populations because of FM/QCF type pulse. Our results indicate that acoustic moni large intra-specific variation in the measured parameters. toring in Korea may be a powerful tool to assess bat activity, Pulse structures of four FM/QCF type species in this including identification to species, at least for the four FM/ study were also consistent with previous studies, which have QCF and one FM/CF/FM species. described pulse structures of these species in other regions. Although pulse frequencies of Hypsugo alaschanicus and Conclusion Pipistrellus abramus were similar to those of Japanese This study reveals echolocation call structures of the 14 Kor populations (PF of H. alaschanicus: 33.9-36.4 kHz, mean ean bat species and those of 11 species are described from PF of Pi. abramus: 46.9 kHz) (Fukui et al., 2002, 2013), the Korea for the first time. Patterns of habitat use and foraging
Anim. Syst. Evol. Divers. 31(3), 160-175 169 Dai Fukui, David A. Hill, Sun-Sook Kim, Sang-Hoon Han behavior of bats have been interpreted in terms of the struc pipistrelle bat (Pipistrellus abramus) in non-reproductive ture of echolocation calls. In theory, it should be possible to season by using radio-tracking. Korean Journal of Envi predict the kinds of habitat that each species in a bat com ronment and Ecology, 24:487-492 (in Korean with English munity is likely to be associated with on the basis of these abstract). characteristics. Such predictions could then be used to devel Chung CU, Han SH, Lim CW, Kim SC, Lee HJ, Kwon YH, Kim CY, Lee CI, 2010c. General patterns in echolocation op plans for conservation management that would enhance call of greater horseshoe bat Rhinolophus ferrumequi habitats for bats. 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172 Anim. Syst. Evol. Divers. 31(3), 160-175 Echolocation Calls of Korean Bats Locality Mitan-myeon, Gangwon-do Mitan-myeon, Mitan-myeon, Gangwon-do Mitan-myeon, Hallim-eup, Jeju-do Hallim-eup, Jeju-do Hallim-eup, Jeju-do Hallim-eup, Jeju-do Hallim-eup, Jeju-do Hallim-eup, Jeollanam-do Jido-eup, Jeollanam-do Jido-eup, Jeollanam-do Haeje-myeon, Jeollanam-do Haeje-myeon, Jeollanam-do Sonbul-myeon, Chungcheongbuk-do Jincheon-eup, Chungcheongbuk-do Jincheon-eup, Jeju-do Jochon-eup, Gangwon-do Yeongweol, Gangwon-do Yeongweol, Jeollabuk-do Seolcheon-myeon, Jeollabuk-do Seolcheon-myeon, Yeongweol, Gangwon-do Yeongweol, Gyeongsangbuk-do Buseok-myeon, Mitan-myeon, Gangwon-do Mitan-myeon, Yeongweol, Gangwon-do Yeongweol, Mitan-myeon, Gangwon-do Mitan-myeon, Mitan-myeon, Gangwon-do Mitan-myeon, Gangwon-do Mitan-myeon, Jeju-do Daejeong-eup, Jeju-do Daejeong-eup, Jeju-do Daejeong-eup, Jeju-do Daejeong-eup, Jeju-do Daejeong-eup, Jeju-do Daejeong-eup, Jeju-do Pyoseon-myeon, Jeju-do Pyoseon-myeon, Jeju-do Pyoseon-myeon, Jeju-do Pyoseon-myeon, Jeju-do Pyoseon-myeon, Situation Perch Free flight Perch Perch Perch Perch Perch Perch Perch Perch Perch Free flight Free flight Free flight Perch Perch Perch Free flight Free flight Free flight Perch Free flight Free flight Free flight Free flight Free flight Perch Perch Perch Perch Perch Perch Perch Perch Perch Perch Perch (kHz)
0.76 2.09 0.90 2.08 2.43 1.11 1.17 0.91 0.90 0.86 1.42 0.99 0.91 1.07 0.71 0.56 0.99 1.21 2.62 3.44 2.83 1.37 1.35 1.46 1.40 0.37 1.00 3.06 1.92 1.80 1.19 1.98 1.35 0.92 1.58 2.01 2.84 ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± 17.3 18.4 18.5 12.8 10.0 8.4 12.6 4.7 13.4 13.5 9.3 9.5 13.9 10.7 11.7 17.3 14.1 17.5 13.3 14.6 13.6 12.9 12.7 9.0 13.2 6.1 10.6 13.7 14.4 10.4 12.3 11.3 19.5 14.0 15.2 10.8 11.2 Band width Band 26.78 26.11 100.78 54.31 3.68 81.17 9.06 26.15 95.56 34.16 27.59 36.92 10.57 21.20 24.09 7.70 16.63 11.36 21.41 11.40 28.81 31.00 40.80 23.61 11.95 109.77 65.64 20.11 29.11 56.18 8.04 11.79 13.17 17.19 4.20 7.57 20.89 (ms)
± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± IPI 46.3 31.0 56.3 101.9 168.8 139.8 97.9 74.7 84.7 131.6 100.3 103.5 56.6 140.4 160.3 169.9 60.9 80.0 105.9 77.5 76.4 120.2 94.0 115.3 96.4 100.3 189.5 87.8 198.7 40.8 44.3 101.3 115.9 38.5 134.6 94.9 68.2 1.67 1.61 3.89 13.79 7.57 9.06 9.18 10.14 8.54 3.63 5.59 6.54 4.95 2.95 4.76 7.32 4.48 5.47 14.81 18.98 5.98 10.81 14.09 13.51 4.87 11.11 12.52 13.05 4.36 6.93 3.42 6.28 2.38 4.05 8.29 13.98 7.32 (ms) ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ±
D 22.6 17.5 31.1 52.2 53.3 54.6 42.3 42.7 44.1 30.5 28.0 52.2 33.0 61.5 35.0 51.8 29.6 46.3 65.7 50.3 25.5 63.6 60.2 63.9 64.5 62.2 55.7 45.7 57.1 24.7 23.8 50.6 52.2 29.7 40.0 47.8 35.4 0.00 0.44 0.00 0.00 0.00 0.49 0.00 0.15 0.24 0.30 0.50 0.12 0.00 0.00 0.00 0.37 0.15 0.00 0.00 0.00 0.11 0.26 0.26 0.19 0.00 0.20 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 0.00 ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± (kHz)
PF 68.9 69.4 72.0 71.0 71.0 71.4 70.7 69.1 68.5 68.0 69.5 69.1 66.8 67.7 72.0 68.9 69.1 68.3 68.3 68.3 68.3 69.3 66.0 69.3 68.3 67.8 72.0 71.0 71.0 72.0 72.0 71.0 72.0 72.0 72.0 72.0 71.9 0.75 1.98 0.90 2.08 2.43 2.54 0.86 1.28 0.99 0.85 3.35 1.09 0.81 0.91 2.84 1.37 0.98 0.71 0.90 0.44 0.89 1.38 1.47 1.81 1.63 1.22 3.07 1.18 1.27 2.01 1.39 0.37 1.00 0.92 3.06 1.58 2.01 (kHz)
± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± 52.3 52.0 53.8 58.6 60.3 63.7 60.8 57.8 52.1 55.7 65.3 56.4 53.3 55.5 63.5 60.5 52.5 55.4 55.4 56.3 59.4 54.2 54.4 61.3 56.9 55.3 66.2 60.9 61.7 53.0 58.4 58.6 57.9 57.2 61.8 61.8 61.5 FMIN 0.14 0.28 0.00 0.00 0.00 0.20 0.00 0.17 0.00 0.40 0.15 0.17 0.16 0.00 0.15 0.00 0.23 0.00 0.00 0.14 1.94 0.31 0.23 0.20 0.28 0.38 0.30 0.24 0.14 0.12 0.14 0.00 0.00 0.00 0.00 0.00 0.00 (kHz)
± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± 69.5 70.4 71.4 70.3 72.2 71.4 69.5 69.4 68.3 70.0 69.7 67.3 69.0 72.8 70.0 70.0 68.7 69.3 69.8 72.2 69.0 66.8 70.2 68.6 72.3 72.2 72.3 72.4 72.4 72.3 72.2 72.4 72.2 72.6 72.7 72.3 69.1 FMAX Descriptive statistics for each echolocation call sequence of species Descriptive 7 10 9 8 5 8 4 5 4 5 9 5 7 5 7 8 7 6 6 11 8 11 10 8 9 8 7 5 9 6 6 3 8 8 7 8 10 No. 1 2 3 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 37 4 36 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Rhinolophus ferrumequinum Call ID Appendix 1.
Anim. Syst. Evol. Divers. 31(3), 160-175 173 Dai Fukui, David A. Hill, Sun-Sook Kim, Sang-Hoon Han Locality Cheongju, Chungcheongbuk-do Hongcheon, Gangwon-do Hampyeong, Jeollanam-do Hampyeong, Jeollanam-do Haeje-myeon, Hongcheon, Gangwon-do Jeju-do Sanghyo-dong, Haeje-myeon, Jeollanam-do Haeje-myeon, Jincheon, Chungcheongbuk-do Sanghyo-dong, Jeju-do Sanghyo-dong, Jeju-do Hawon-dong, Jeollabuk-do Namweon, Jincheon, Chungcheongbuk-do Sanghyo-dong, Jeju-do Sanghyo-dong, Jincheon, Chungcheongbuk-do Gangwon-do Mitan-myeon, Yeongju, Gyeongsangbuk-do Yeongju, Gangwon-do Yeongweol, Gangwon-do Daehwa-myeon, Mitan-myeon, Gangwon-do Mitan-myeon, Yeongweol, Gangwon-do Yeongweol, Daehwa-myeon, Gangwon-do Daehwa-myeon, Yeongju, Gyeongsangbuk-do Yeongju, Hwacheon, Gangwon-do Hwacheon, Yeongju, Gyeongsangbuk-do Yeongju, Yeongju, Gyeongsangbuk-do Yeongju, Jecheon, Chungcheongbuk-do Hongcheon, Gangwon-do Hongcheon, Gangwon-do Hongcheon, Gangwon-do Hongcheon, Gangwon-do Situation Flight room Release call Release Flight room call Release Release call Release call Release Release call Release call Release Release call Release call Release call Release Release call Release Release call Release Release call Release Flight room Release call Release call Release Flight room Flight room Release call Release Flight room Flight room Flight room Flight room Flight room Release call Release Release call Release Release call Release Release call Release Release call Release (kHz)
4.28 9.17 14.09 12.75 5.98 10.87 15.34 14.39 9.96 7.20 8.96 9.81 6.96 4.19 1.96 5.17 7.07 1.56 8.01 10.88 3.02 6.29 4.57 6.81 7.30 4.31 11.74 4.45 4.13 4.16 ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± 54.0 90.6 75.8 64.7 85.9 52.2 53.8 74.9 56.4 50.4 87.7 71.6 87.5 67.3 22.7 67.7 53.0 22.4 82.8 27.0 79.6 56.8 86.6 92.9 83.3 86.3 89.6 113.1 109.4 71.4 Band width Band 20.64 14.90 35.19 5.40 2.46 2.69 18.42 9.59 19.57 9.50 20.79 14.07 13.87 19.44 12.86 47.49 8.64 15.42 41.81 18.33 35.55 23.78 15.75 46.00 6.47 12.44 3.00 7.19 9.35 15.24 (ms)
± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± IPI 65.8 58.7 71.9 91.5 72.7 41.1 147.2 78.9 46.7 68.2 57.4 64.0 52.2 121.3 63.3 90.5 54.7 67.7 64.9 61.3 46.6 100.9 58.7 58.8 121.3 58.8 66.3 53.0 66.0 62.7 0.24 0.34 0.41 0.36 0.37 0.54 0.36 0.29 0.45 0.23 0.37 0.59 0.21 0.41 0.26 0.60 0.17 0.21 0.36 0.35 0.23 0.42 0.33 0.27 0.89 0.32 0.28 0.24 0.27 0.39 (ms)
± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± D 1.7 2.6 2.2 2.8 3.9 2.6 5.9 3.7 2.6 3.5 3.3 3.7 2.4 5.1 2.7 2.9 3.2 1.4 2.6 2.8 1.2 3.0 1.8 2.6 3.6 4.0 2.6 3.6 3.7 2.9 4.13 1.70 2.95 5.17 2.10 4.86 2.11 1.45 5.54 4.30 8.46 2.99 6.47 1.72 2.16 9.04 1.92 0.86 1.29 1.25 1.70 3.62 1.64 4.61 6.43 1.33 1.27 1.72 2.49 2.79 ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± (kHz)
PF 49.7 59.7 55.4 54.3 54.4 51.9 42.3 48.2 49.9 50.5 39.0 50.4 56.9 46.9 57.4 40.6 55.9 38.5 57.3 46.5 39.2 60.6 39.6 62.6 46.7 51.2 56.7 58.1 54.0 60.9 2.17 1.83 2.74 1.48 1.42 1.76 0.84 1.65 1.84 2.57 0.81 2.04 1.66 1.24 1.91 1.83 0.68 0.97 1.92 3.13 1.90 1.53 2.07 1.55 4.07 1.84 2.56 1.09 0.94 2.79 (kHz)
± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± 31.4 41.6 34.8 33.1 33.8 40.5 27.3 34.0 34.5 39.9 21.7 33.3 25.3 32.9 36.5 17.2 26.7 42.5 39.9 33.9 26.7 43.7 25.4 44.2 26.6 35.5 38.9 39.9 39.2 38.8 FMIN 5.35 11.71 8.79 13.73 7.91 14.29 4.48 4.12 6.85 8.73 5.47 2.08 6.82 6.47 14.02 5.18 8.44 7.12 3.40 9.74 9.36 4.18 4.15 3.10 (kHz)
8.89 8.71 6.78 1.41 2.28 3.06 ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± FMAX 85.4 132.2 97.8 110.6 86.0 126.5 109.0 81.1 91.0 90.3 134.8 121.0 134.6 124.0 104.4 88.5 49.4 109.8 107.7 87.0 49.1 126.5 52.4 123.8 83.4 122.1 131.8 123.2 125.4 128.4 Continued 15 17 6 12 20 13 15 17 26 8 22 30 22 11 18 10 8 19 15 14 31 12 13 22 27 16 11 23 18 19 No. 1 2 1 12 1 3 4 2 1 2 3 2 1 3 3 1 2 1 2 2 3 3 4 5 6 7 8 9 10 11 Myotis aurascens Call ID Myotis macrodactylus Myotis formosus Myotis bombinus Myotis ikonnikovi Myotis petax Plecotus ognevi Appendix 1.
174 Anim. Syst. Evol. Divers. 31(3), 160-175 Echolocation Calls of Korean Bats Locality Jochon-eup, Jeju-do Jochon-eup, Yeongweol, Gangwon-do Yeongweol, Gangwon-do Yeongweol, Toji-myeon, Jeollanam-do Toji-myeon, Hawon-dong, Jeju-do Hawon-dong, Toji-myeon, Jeollanam-do Toji-myeon, Hawon-dong, Jeju-do Hawon-dong, Gyeongju, Gyeongsangbuk-do Gyeongju, Jecheon, Chungcheongbuk-do Hawon-dong, Jeju-do Hawon-dong, Gyeongsangbuk-do Gyeongju, Jecheon, Chungcheongbuk-do Cheongju, Chungcheongbuk-do Gyeongju, Gyeongsangbuk-do Gyeongju, Jecheon, Chungcheongbuk-do Cheongju, Chungcheongbuk-do Cheongju, Chungcheongbuk-do Samcheog, Gangwon-do Jecheon, Chungcheongbuk-do Sanghyo-dong, Jeju-do Sanghyo-dong, Jecheon, Chungcheongbuk-do Jecheon, Chungcheongbuk-do Mitan-myeon, Gangwon-do Mitan-myeon, Cheongju, Chungcheongbuk-do Jecheon, Chungcheongbuk-do Gangwon-do Yeongweol, Situation Free flight Release call Release call Release Flight room Release call Release Flight room Release call Release Release call Release Flight room Release call Release call Release Release call Release call Release Release call Release Release call Release call Release Release call Release Release call Release Release call Release Release call Release call Release Release call Release Free flight call Release Release call Release call Release (kHz)
9.60 3.69 6.33 5.38 5.21 19.24 11.87 3.70 18.31 5.00 7.56 3.05 2.24 8.42 2.84 18.19 2.37 6.69 3.62 11.06 4.37 4.92 2.16 4.91 8.84 3.15 ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± 70.9 85.4 67.9 73.6 82.2 9.4 15.9 30.8 12.9 37.6 19.9 33.1 25.8 30.0 42.3 32.7 15.6 34.2 39.7 22.4 90.3 96.3 84.2 96.7 102.4 67.3 Band width Band 61.85 5.02 2.61 14.73 9.43 56.12 21.01 18.21 12.71 10.41 18.47 8.22 30.20 25.75 19.72 8.60 20.26 17.52 50.86 8.66 29.37 27.95 29.03 14.81 14.82 28.56 (ms)
± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± IPI 208.1 102.2 38.2 48.9 65.9 51.6 68.4 136.3 36.5 88.1 109.4 59.5 79.6 89.4 47.8 45.5 75.5 40.3 38.9 67.8 138.0 145.1 175.9 70.7 132.5 114.4 1.12 0.51 0.38 0.88 0.42 1.30 0.84 0.23 0.30 0.51 0.58 0.19 0.49 0.57 0.46 0.13 0.93 0.66 0.32 1.29 0.66 0.63 1.16 0.36 0.73 0.92 (ms)
± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± D 7.2 7.0 3.3 4.2 2.9 4.7 3.3 6.0 1.6 5.6 5.4 1.9 4.0 5.2 3.2 2.1 3.1 3.1 2.7 3.5 8.9 8.3 7.8 2.4 8.0 7.3 0.67 0.40 0.89 10.70 0.97 17.28 1.12 0.41 21.83 0.76 0.66 16.04 15.47 0.69 13.71 4.33 2.58 11.79 10.10 4.94 1.10 0.58 0.73 1.53 0.89 0.34 ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± (kHz)
PF 50.3 35.5 51.9 74.4 52.9 82.9 54.2 48.1 79.8 50.9 47.0 67.9 70.0 47.3 77.1 54.9 59.5 53.4 60.6 45.7 31.1 29.7 34.0 36.2 31.6 36.0 0.41 0.49 1.62 2.23 0.37 3.80 0.75 0.64 2.14 0.58 0.45 0.60 4.71 0.78 5.46 1.41 2.30 3.33 2.77 3.68 0.44 0.37 0.66 0.95 0.50 0.36 (kHz)
± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± 47.4 32.1 46.8 49.6 49.7 48.0 48.8 44.7 51.0 47.2 43.4 38.4 41.2 44.1 44.4 32.9 42.7 30.2 33.0 25.2 32.8 25.0 30.7 29.3 24.9 32.9 FMIN SD. 11.56 5.54 18.61 9.09 4.64 5.83 3.14 4.95 3.34 6.04 19.96 ± (kHz)
2.21 3.10 8.21 2.74 18.12 2.16 6.63 3.51 11.49 7.70 4.46 2.17 4.21 8.47 3.11 ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± FMAX 56.8 48.0 77.6 120.6 62.6 133.4 86.4 64.6 118.9 80.3 69.2 128.7 114.8 74.1 126.6 129.2 126.8 126.9 135.4 67.6 100.1 57.7 46.3 63.5 64.6 55.4 Continued 5 10 6 19 10 16 28 8 12 22 10 5 12 22 19 10 12 19 20 10 22 10 6 9 19 11 No. 1 2 1 3 4 2 4 3 5 1 2 4 3 1 5 2 3 4 5 1 6 2 3 1 2 3 Miniopterus fuliginosus Murina ussuriensis Call ID means as presented are Values No., number of measured pulses; FMAX, maximum frequency; FMIN, minimum frequency; PF, frequency of maximum energy; D, duration; IPI, inter-pulse interval. inter-pulse IPI, duration; D, energy; maximum of frequency PF, frequency; minimum FMIN, frequency; maximum pulses; FMAX, measured of number No., Pipistrellus abramus Murina leucogaster Eptesicus serotinus Hypsugo alaschanicus Appendix 1.
Anim. Syst. Evol. Divers. 31(3), 160-175 175