Bat Echolocation Researc h - Bat Conservation International

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Echolocation calls of Euderma maculatum (Chiroptera: Vespertilionidae): use in orientation and communication. Journal of Mammalogy 65:122-126. LIMPENS, H. J. G. A., W. BONGERS, and W. HELMER. 2001. Territorium en strategie van de rosse vleermuis in de paartijd. Zoogdier 12:26-27. MANKSE, U., and U. SCHMIDT. 1976. Visual acuity of the vampire bat, Desmodus rotundus, and its dependence upon light intensity. Zeitschrift fur Tierpsychologie 42:215-221. MARIMUTHU, G., J. HABERSETZER, and D. LEIPPERT. 1995. Active acoustic gleaning from the water surface by the Indian false vampire bat, Megaderma lyra. Ethology 99:61-74. MILLER, L. A., and A. SURLYKKE. 2001. How some insects detect and avoid being eaten by bats: tactics and countertactics of prey and predator. Bioscience 51:570-581. OBRIST, M. K. 1995. Flexible bat echolocation: the influence of individual, habitat, and conspecifics on sonar signal design. Behavioral Ecology and Sociobiology 36:207-219. POPPER, A. N., and R. R. 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y the African bat Cardioderma cor (Megadermatidae). Journal of Comparative Physiology A161:59-66. SCHNITZLER, H.-U., and E. K. V. KALKO. 2001. Echolocation by insect-eating bats. Bioscience 51:557-569. SCHUMM, A., D. KRULL, and G. NEUWEILER. 1991. Echolocation in the notch-eared bat, Myotis emarginatus. Behavioral Ecology and Sociobiology 28:255-261. SIEMERS, B. M., P. STILZ, and H.-U. SCHNITZLER. 2001. The acoustic advantage of hunting at low heights above water: behavioral experiments on the European ‘trawling’ bats Myotis capaccini, M. dasycneme and M. daubentoni. The Journal of Experimental Biology 204:3843-3854. SIMMONS, J. A., and R. A. STEIN. 1980. Acoustic imaging in bat sonar: echolocation signals and the evolution of echolocation. Journal of Comparative Physiology A135:61-84. SUTHERS, R. A. 1970. Vision, olfaction and taste. Pp. 265- 309 in Biology of bats. Volume II (W. A. Wimsatt, ed.). Academic Press, New York. THIES, W., E. K. V. KALKO, and H.-U. SCHNITZLER. 1998. The roles of echolocation and olfaction in two neotropical fruit-eating bats, Carollia perspicillata and C. castanea, feeding on Piper. Behavioral Ecology and Sociobiology 42:397-409. VERBOOM, B., A. M. BOONMAN, M. BOONMAN, and H. J. G. A. LIMPENS. 1999. Acoustic perception of landscape elements by the pond bat (Myotis dasycneme). Journal of Zoology (London) 248:59-66. VON HELVERSEN, D., and O. VON HELVERSEN. 1999. Acoustic guide in bat-pollinated flower. Nature 398:759- 760. VON HELVERSEN, D., M. W. HOLDERIED, and O. VON HELVERESEN. 2003. Echoes of bat-pollinated, bellshaped flowers: conspicuous for nectar-feeding bats? Journal of Experimental Biology 206:1025-1034. WILLIAMS, T. C., J. M. WILLIAMS, and D. R. GRIFFIN. 1966. The homing ability of the neotropical bat Phyllostomus hastatus. Science 155:1435-1436. THE PAST AND FUTURE HISTORY OF BAT DETECTORS DONALD R. GRIFFIN Deceased Work done at Concord Field Station, Harvard University, Old Causeway Road, Bedford MA 01730, United States What we can learn about the echolocation abilities of bats has been limited by the sensitivity of microphones; and some of their high-frequency sounds cannot be detected under many conditions, even with the best instruments currently available. For example, sensitive heterodyne bat detectors sometimes fail to detect feeding buzzes from Myotis lucifugus on early summer evenings when they are obviously catching insects. By mounting the bat detector on a long pole and using infrared video showing the bat and the insect it captured, we found that buzzes were detectable only if the microphone was within one or two meters of the bat. Yet at other times buzzes could be detected from the same bats in the same location at distances of ten meters. Many neotropical bats emit much fainter sounds than M. lucifugus, and there is some evidence that shrews and laboratory rats use echolocation, although it has been difficult to detect whatever sounds they may be emitting. There is an “apparatus threshold” which is ordinarily well above the auditory thresholds of bats and shrews, so that there is a sizeable intensity range where we cannot yet determine whether sounds are being used for echolocation or communication. Key words: bat detectors, echolocation, equiptment, insect capture by bats, microphone sensitivity, Myotis lucifugus 6 PAST HISTORY To study the echolocation abilities of bats it is first necessary to detect their orientation sounds. The fact that bats make ultrasonic sounds was discovered by means of G. W. Pierce’s “sonic amplifier,” which was a heterodyne detector with an audio output constructed by modifying an AM radio receiver (Pierce and Griffin 1938). This apparatus had been developed to study the sounds of insects; its circuit is described by Noyes and Pierce (1937) and the microphones by Pierce (1948). The essential first element in any apparatus for detecting bat sounds is the microphone. Pierce used magnetostriction transducers and Rochelle salt crystals as microphones; the latter were somewhat more sensitive, but neither was calibrated nor nearly as sensitive as microphones developed later. Pierce’s apparatus displayed the heterodyne output graphically, but its temporal resolution was not adequate to measure the duration of the ultrasonic orientation sounds used by bats of the family Vespertilionidae. It could record pulse repetition rates and revealed that the interpulse interval dropped by about half when Myotis lucifugus avoided small wires (Galambos and Griffin 1942). Furthermore it enabled us to demonstrate that emission and reception of orientation sounds were necessary for obstacle avoidance (Griffin and Galambos 1941; Griffin 1958). The next step was to use the Western Electric 640 Bat Echolocation Research: tools, techniques & analysis
Page 1 and 2: Bat Echolocation R esearc h tools,
Page 3 and 4: TABLE OF CONTENTS CO N T R I B U T
Page 5 and 6: CONTRIBUTING AUTHORS Ingemar Ahlén
Page 7 and 8: I N T R O D U C T I O N Bat Echoloc
Page 9 and 10: BAT NATURAL HISTORY AND ECHOLOCATIO
Page 11: species (Thies et al. 1998). Before
Page 15 and 16: participants in this symposium. Sin
Page 17 and 18: INTRODUCTION Bat detectors convert
Page 19 and 20: less sensitive than a heterodyne de
Page 21 and 22: 14 advanced computer software made
Page 23 and 24: COMMUTING AND FORAGING BEHAVIOR Fig
Page 25 and 26: NIGHTLY ACTIVITY BASED ON CAPTURE W
Page 27 and 28: 20 Recordings of echolocation calls
Page 29 and 30: 22 and analyzing echolocation calls
Page 31 and 32: 24 Resource partitioning in rhinolo
Page 33 and 34: Section 2 ACOUSTIC INVENTORIES ULTR
Page 35 and 36: f tuned = f bat . As an example, wi
Page 37 and 38: The chance of detection also depend
Page 39 and 40: ent species. But in the field, it i
Page 41 and 42: Section 2: Acoustic Inventories Fig
Page 43 and 44: Suchflug und Richtungskarakteristik
Page 45 and 46: Journal N Percentage of total (%) A
Page 47 and 48: ential power of data. Hayes (1997)
Page 49 and 50: For instance, we recorded a single
Page 51 and 52: KALCOUNIS, M. C., K. A. HOBSON, R.
Page 53 and 54: and McCracken (this volume) discuss
Page 55 and 56: using other methods (Fig. 4). Ident
Page 57 and 58: Figure 6: Schematic diagram of diff
Page 59 and 60: AMPLITUDE-RELATED PARAMETERS Loudne
Page 61 and 62: tereri. Hand netting on the followi
Page 63 and 64:
flight in bats. Pp. 93-108 in Bat b
Page 65 and 66:
difficult to understand why the aut
Page 67 and 68:
Figure 5: Call sequence showing the
Page 69 and 70:
function analysis and artificial ne
Page 71 and 72:
species identifications. Echolocati
Page 73 and 74:
tially from calls emitted in the op
Page 75 and 76:
LITERATURE CITED BARCLAY, R. M. R.,
Page 77 and 78:
HETERODYNE AND TIME-EXPANSION METHO
Page 79 and 80:
74 QUANTIFYING SOUND FEATURES To qu
Page 81 and 82:
Figure 14: Pulse rhythm diagrams fo
Page 83 and 84:
A FINAL WORD Figure 21: Heterodyne
Page 85 and 86:
80 INTRODUCTION Ultrasonic detector
Page 87 and 88:
82 munity was uncovered when ultras
Page 89 and 90:
84 INTRODUCTION Being nocturnal, ba
Page 91 and 92:
A Figure 2: Surveys of bat activity
Page 93 and 94:
LITERATURE CITED Figure 5A: Pipistr
Page 95 and 96:
90 In order to set a foundation for
Page 97 and 98:
92 diagram, e.g., a spectrogram, fo
Page 99 and 100:
ecording time of the cassette. When
Page 101 and 102:
96 detail is not helpful for specie
Page 103 and 104:
tification, providing the immediate
Page 105 and 106:
spectrum, which is a graph of the e
Page 107 and 108:
since there is no evidence that the
Page 109 and 110:
104 Figure 5: Different modes in se
Page 111 and 112:
currently the case. Even at the sim
Page 113 and 114:
108 INTRODUCTION Much of our knowle
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Figure 1: On-axis search call recor
Page 117 and 118:
112 SUMMARY To summarize, flight-ca
Page 119 and 120:
114 SIGNAL PROCESSING TECHNIQUES FO
Page 121 and 122:
al a situation as possible. Ideally
Page 123 and 124:
For all networks, correct identific
Page 125 and 126:
ety of London 69:111-128. JONES, G.
Page 127 and 128:
eveal species-specific vocal signat
Page 129 and 130:
Figure 6: Comparison of Myotis cili
Page 131 and 132:
ity of the two Myotis spp. calls (F
Page 133 and 134:
Section 4 RESOURCES, RESEARCH, AND
Page 135 and 136:
A The data extracted from the diffe
Page 137 and 138:
used. This theoretically makes it p
Page 139 and 140:
ple, will not show individual calls
Page 141 and 142:
Figure 10: Fast Fourier transform p
Page 143 and 144:
ACKNOWLEDGEMENTS I thank the organi
Page 145 and 146:
munity (now called the European Uni
Page 147 and 148:
staff to collect all of the require
Page 149 and 150:
and van Parijs 1993). Early studies
Page 151 and 152:
Figure 5: Variation in the echoloca
Page 153 and 154:
47:60-69. JONES, G., and R. D. RANS
Page 155 and 156:
RECORDING TECHNIQUES Section 4: Res
Page 157 and 158:
monly observed with such broadband
Page 159 and 160:
ple sonograms and power spectra can
Page 161 and 162:
ences between Myotis californicus a
Page 163 and 164:
intensity). Our ability to detect m
Page 165 and 166:
• The detector is kept stationary
Page 167 and 168:
the number of bat passes in real ti
Page 169 and 170:
SHERWIN, R. E., W. L. GANNON, and S
Page 171:
tion trends, while they also raise
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