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The bat fauna of Papua New Guinea (PNG) is more diverse than that of Australia, yet knowledge of the distribution and ecology of PNG's 57 microchiropteran species is particularly poor, almost a third being known from five or less localities. The lack of knowledge is partly due to problems with use of both mist nets and harp traps in PNG. Bat detectors may help overcome some of these problems, however, they have not been widely used in PNG primarily due to the lack of a body of reference calls to aid species identification. Using the Anabat system, we recorded 744 reference echolocation call sequences from eight microchiropteran species captured in PNG. Calls were analysed using Analook software and described. The characteristic frequency of the predominant harmonics were: Aselliscus tricuspidatus112-113 kHz; Hipposideros cervinus136.5-138 kHz; H. diadema54-59 kHz; H. maggietaylorae121-123 kHz; Mormopterus cf. beccarii44-50 kHz; Mosia nigrescens45-60 kHz; Rhinolophus arcuatus70-72 kHz; and Rhinolophus euryotis52-56 kHz. Comparison with published calls of some of these species from Australia, south-east Asia and elsewhere in PNG suggest regional variations occur within PNG and abroad and/or that there are taxonomic issues such as cryptic species. This emphasises the need for far greater collection of reference echolocation calls, the development of regional PNG call libraries with vouchered specimens and cautions against using reference libraries developed in Australia or elsewhere.

Aldridge, H.D.J., Obrist, N.M., Merriam, H.G. and Fenton, M.B. 1990. Roosting, vocalizations, and foraging by the African bat, Nycteris thebaica. Journal of Mammalogy 71 (2): 242-246.
Aldridge, H. D. J., Rautenbach, I.L. 1987. Morphology, echolocation and resource partitioning in insectivorous bats. The Journal of Animal Ecology 56 (3): 763-778.
Armstrong, K.N. and Coles, R.B. 2007. Echolocation differences between geographic isolates of Rhinonicteris aurantia(Chiroptera: Hipposideridae): implications of nasal chamber size. Journal of Mammalogy 88: 94-104.
Barclay, R.M.R., Fullard, J.H., and Jacobs, D.S. 1999. Variation in the calls of the hoary bat ( Lasiurus cinereus): influence of body size, habitat structure and geographic location. Canadian Journal of Zoology 77: 530-534.
Barratt, E.M., Deaville, R., Burland, T.M. and Bruford, M.W. 1997. DNA answers the call of the pipisterelle bat species. Nature 387: 138-139.
Bonaccorso, F. 1998 Bats of Papua New Guinea. Conservation International Tropical Field Guide Series, Washington, DC.
Broders, H.G., Findlay, S.F., and Zhang, L. 2004. Effects of clutter on echolocation call structure of Myotis septentrionalis and M. lucifugus. Journal of Mammalogy 85 (2): 273-281.
Brown, P. E. and Berry, R. D. 1983. Echolocation behaviour of a “Flycatcher” bat Hipposideros diadema. The Journal of the Acoustical Society of America74(S1), S32.
Bullen, R.D. and McKenzie, N. L. 2002. Scaling bat wingbeat frequency and amplitude. The Journal of Experimental Biology 205: 2615-2626.
Chen, M., Feng, J., Liu, Y., Zhang, X., Li, Z., Zhou, J. and Zhang, S. 2004. Relationship between echolocation calls and age in the bat species of Rhinolophus pearsoni. Zoological Research 25 (1): 15-19.
Churchill, S. 1998 Australian Bats. Reed New Holland Publishing, Sydney.
Corben, C. and Fellers, G.M. 2001. Choosing the ‘correct’ bat detector - a reply. Acta Chiropterologica 3 (2): 245-256.
Corben, C. and O'Farrell, M.J. 1999. Techniques for the effective use of Anabat in identifying free-flying bat species. Anabat system manual.
Crome, F.H.J. and Richards, G.C. 1988. Bats and gaps: Microchiropteran community structure in a Queensland rainforest. Ecology 69 (6): 1960-1969.
Fenton, M.B. 1982. Echolocation calls and patterns of hunting and habitat use of bats (Microchiroptera) from Chillagoe, north Queensland. Australian Journal of Zoology 30: 417-425.
Fenton, M.B., Audet, D., Obrist, M.K., and Rydell, J. 1995. Signal strength, timing, and self-deafening: The evolution of echolocation in bats. Paleobiology 21 (2): 229-242.
Filewood, L.W. 1983. The possible occurrence in New Guinea of the ghost bat ( Macroderma gigas: Chiroptera: Megadermatidae). Australian Mammalogy 6: 35-36.
Fukui, D., Agtsuma, N., and Hill, D.A. 2004. Acoustic identification of eight species of bat (Mammalia: Chiroptera) inhabiting forests of southern Hokkaido, Japan: Potential for conservation monitoring. Zoological Science 21: 947-955.
Gebia, O. and Balun, L. 2004. Species diversity and composition in the Darai limestone karst and Libano alluvial forest in Gulf and Southern Highlands Provinces. KICDP Scientific Report: SR-04-01. Unpublished report to WWF.
Griffiths, T.A., Koopman, K.F. and Starrett, A. 1991. The systematic relationship of Emballonura nigrescens to other species of Emballonura and to Coleura(Chiroptera: Emballonuridae). American Museum Novitates 2996: 1-16.
Grinnell, A.D. and Hagiwara, S. 1972. Adaptations of the auditory nervous system for echolocation. Studies of New Guinea bats. Zeitscrift fur vergleichende Physiologie 76: 41-81.
Guillen, A., Juste, B., and Ibanez, C. 2000. Variation in the frequency of the echolocation calls of Hipposideros ruber in the Gulf of Guinea: an exploration of the adaptive meaning of the constant frequency value in rhinolophid CF bats. Journal of Evolutionary Biology 13: 70-80.
Hall, L.S. and Richards, G. 1995. Diadem Leafnosed-bat Hipposideros diadema. Pp. 459-460 in The Mammals of Australia, Second Edition, edited by R. Strahan. Reed Books, Chatswood, NSW.
Helgen, K. 2005. Systematics of the Pacific monkey-faced bats (Chiroptera: Pteropidae) with a new species of Pteralopex and a new Fijian genus. Systematics and Biodiversity 3 (4): 433-453.
Helgen, K. 2006. A taxonomic and geographic overview of mammals of Papua. Pp. 689-749 in The Ecology of Papua edited by A.J. Marshall and B. M. Beehler. Periplus editions, Singapore.
Heller, K.G. and Von Helversen, O. 1989. Resources partitioning of sonar frequency bands in rhinolophoid bats. Oecologia 80: 178-186.
Holderied, M.W. and von Helversen, O. 2003. Echolocation range and wingbeat period match in aerial-hawking bats. Proceedings of the Royal Society Biological Sciences 270(1530): 2293-2299.
Holderied, M.W., Korine, C., Fenton, M.B., Parsons, S., Robson, S. and Jones, G. 2005. Echolocation call intensity in the aerial hawking bat Eptesicus bottae(Vespertilionidae) studied using stereo videogrammetry. Journal of Experimental Biology 208: 1321-1327.
Jacobs, D.S., Eick, G.N., Schoeman, M.C. and Matthee, C.A. 2006. Cryptic species in an insectivorous bat Scotophilus dinganii. Journal of Mammalogy 87: 161-170.
Jenkins, P.D. and Hill, J.E. 1981. The status of Hipposideros galeritus(Cantor, 1846) and Hipposideros cervinus(Gould 1854) (Chiroptera: Hipposideridae). Bulletin of the British Museum of Natural History, Zoology 41: 279-294.
Jones, G. 1994. Scaling of wingbeat and echolocation pulse emission rates in bats: Why are aerial insectivorous bats so small? Functional Ecology 8(4): 450-457.
Jones, G. and Barlow, K.E. 2004. Cryptic species of echolocating bats. Pp. 345-349 in Echolocation in bats and dolphins, edited by J.A. Thomas, C.F. Moss, and M. Vater. University of Chicago Press, Chicago, Illinois.
Jones, G. and Holderied, M.W. 2007. Bat echolocation calls: adaptation and convergent evolution. Proceedings of the Royal Society Biological Sciences 274(1612): 905-912.
Kazial, K.A., Burnett, S.C. and Masters, W.M. 2001. Individual and group variation in echolocation calls of big brown bats, Eptesicus fuscus(Chiroptera: Vespertilionidae). Journal of Mammalogy 82(2): 339-351.
Kingston, T. and Rossiter, S.J. 2004. Harmonic hopping in Wallacea's bat. Nature 429: 654-657.
Kingston, T., Jones, G., Zubaid, A. and Kunz, T.A. 2000. Resource partitioning in rhinolophoid bats revisited. Oecologia 124: 332-342.
Kingston, T., Lara, M.C., Jones, G., Zubaid, A., Kunz, T.H. and Schneider, C.J. 2001. Acoustic divergence in two cryptic Hipposideros species: a role for social selection? Proceedings of the Royal Society of London, Series B Biological Sciences 268: 1381-1386.
Law, B.S., Reinhold, L. and Pennay, M. 2002. Geographic variation in the echolocation calls of Vespadelus spp. (Vespertilionidae) from New South Wales and Queensland, Australia. Acta Chiroptologica 4 (2): 201-215.
Leary, T. and Mamu, T. 2004. Conserving Papua New Guinea's forest fauna through community planning. Pp. 186-207 in Conservation of Australia's Forest Fauna(Second Edition), edited by D. Lunney. Royal Zoological Society of New South Wales, Mosman, NSW.
Li, G., Liang, B., Wang, Y., Zhao, H., Helgen, K.M., Liangkong, L., Jones, G. and Zhang, S. 2007. Echlocation calls, diet and phylogenetic relationships of Stoliczka's trident bat Aselliscus stoliczkanus(Hipposideridae). Journal of Mammalogy 88(3):736-744.
Macias, S. and Mora, E.C. 2003. Variation of echolocation calls of Pteronotus quadridens(Chiroptera: Mormoopidae) in Cuba. Journal of Mammalogy 84(4):1428-1436.
Macias, S., Mora, E.C., Koch, C. and Von Helversen, O. 2005. Echolocation behaviour of Phyllops falcatus(Chiroptera: Phyllostomidae): unusual frequency range of the first harmonic. Acta Chiropterologica 7(2): 275-283.
Matsumura, S. 1979. Mother-infant communication in a horseshoe bat ( Rhinolophus ferrumequinum nippon): development of vocalization. Journal of Mammalogy 60(1): 76-84.
McKenzie, N.L. 1995. Beccari's Freetail-Bat Mormopterus beccarii. Pp. 480-481 in The Mammals of Australia(Second Edition) edited by R. Strahan. Reed Books, Chatswood, NSW.
McKenzie, N.L. and Bullen, R.D. 2008. Beccari's Free-tailed Bat Mormopterus beccarii. Pp. 486-487 in The Mammals of Australia(Third Edition) edited by S. Van Dyck and R. Strahan. Reed New Holland Books, Chatswood, NSW.
McKenzie, N.L., Start, A.N. and Bullen, R.D. 2002. Foraging ecology and organisation of a desert bat fauna. Australian Journal of Zoology 50 (5): 529-545.
Milne, D.J. 2002 Key to the Bat Calls of the Top End of the Northern Territory. Parks and Wildlife Commission of the Northern Territory. Technical Report 71.
Novick, A. 1958. Orientation in palaeotropical bats. I. Microchiroptera. Journal of Experimental Biology 138: 81-153.
Novick, A. 1977. Acoustic orientation. Pp. 74-289 in Biology of Bats-Volume III edited by W.A. Wimsatt. Academic Press, New York.
O'Farrell, M.J. and Miller, B.W. 1999. Use of vocal signatures for the inventory of free-flying Neotropical bats. Biotropica. 31(3): 507-516.
O'Farrell, M., Miller, B.W. and Gannon, W.L. 1999. Qualitative identification of free-flying bats using the Anabat detector. Journal of Mammalogy 80(1): 11-23.
Obrist, M., Aldridge, H.D.J.N. and Fenton, M.B. 1989. Roosting and echolocation behavior of the African bat, Chalinolobus variegatus. Journal of Mammalogy 70(4): 828-833.
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.
Paijmans, K. 1976 New Guinea Vegetation. Australian National University Press, Canberra.
Papadatou, E., Butlin, R.K. and Altringham, J.D. 2008. Identification of bat species in Greece from their echolocation calls. Acta Chiropterologica 10(1): 127-143.
Parsons, S. 1998. The effect of recording situation on the echolocation calls of the New Zealand lesser short-tailed bat ( Mystacina tuberculata Gray). New Zealand Journal of Zoology 25: 147-156.
Parsons, S., Boonman, A.J. and Obrist, M.K. 2000. Advantages and disadvantages of techniques for transforming and analyzing chiropteran echolocation calls. Journal of Mammalogy 81(4): 927-938.
Pavey, C.R. and Burwell, C.J. 1995. Fawn Leafnosed Bat Hipposideros cervinus. Pp. 457-458 in The Mammals of Australia(Second Edition) edited by R. Strahan. Reed Books, Chatswood, NSW.
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Surlykke, A., Futtrup, V. and Tougaard, J. 2003. Prey-capture success revealed by echolocation signals in pipistrelle bats ( Pipistrellus pygmaeus). The Journal of Experimental Biology 206: 93-104.
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Contents

Data & Figures

References

Aldridge, H.D.J., Obrist, N.M., Merriam, H.G. and Fenton, M.B. 1990. Roosting, vocalizations, and foraging by the African bat, Nycteris thebaica. Journal of Mammalogy 71 (2): 242-246.
Aldridge, H. D. J., Rautenbach, I.L. 1987. Morphology, echolocation and resource partitioning in insectivorous bats. The Journal of Animal Ecology 56 (3): 763-778.
Armstrong, K.N. and Coles, R.B. 2007. Echolocation differences between geographic isolates of Rhinonicteris aurantia(Chiroptera: Hipposideridae): implications of nasal chamber size. Journal of Mammalogy 88: 94-104.
Barclay, R.M.R., Fullard, J.H., and Jacobs, D.S. 1999. Variation in the calls of the hoary bat ( Lasiurus cinereus): influence of body size, habitat structure and geographic location. Canadian Journal of Zoology 77: 530-534.
Barratt, E.M., Deaville, R., Burland, T.M. and Bruford, M.W. 1997. DNA answers the call of the pipisterelle bat species. Nature 387: 138-139.
Bonaccorso, F. 1998 Bats of Papua New Guinea. Conservation International Tropical Field Guide Series, Washington, DC.
Broders, H.G., Findlay, S.F., and Zhang, L. 2004. Effects of clutter on echolocation call structure of Myotis septentrionalis and M. lucifugus. Journal of Mammalogy 85 (2): 273-281.
Brown, P. E. and Berry, R. D. 1983. Echolocation behaviour of a “Flycatcher” bat Hipposideros diadema. The Journal of the Acoustical Society of America74(S1), S32.
Bullen, R.D. and McKenzie, N. L. 2002. Scaling bat wingbeat frequency and amplitude. The Journal of Experimental Biology 205: 2615-2626.
Chen, M., Feng, J., Liu, Y., Zhang, X., Li, Z., Zhou, J. and Zhang, S. 2004. Relationship between echolocation calls and age in the bat species of Rhinolophus pearsoni. Zoological Research 25 (1): 15-19.
Churchill, S. 1998 Australian Bats. Reed New Holland Publishing, Sydney.
Corben, C. and Fellers, G.M. 2001. Choosing the ‘correct’ bat detector - a reply. Acta Chiropterologica 3 (2): 245-256.
Corben, C. and O'Farrell, M.J. 1999. Techniques for the effective use of Anabat in identifying free-flying bat species. Anabat system manual.
Crome, F.H.J. and Richards, G.C. 1988. Bats and gaps: Microchiropteran community structure in a Queensland rainforest. Ecology 69 (6): 1960-1969.
Fenton, M.B. 1982. Echolocation calls and patterns of hunting and habitat use of bats (Microchiroptera) from Chillagoe, north Queensland. Australian Journal of Zoology 30: 417-425.
Fenton, M.B., Audet, D., Obrist, M.K., and Rydell, J. 1995. Signal strength, timing, and self-deafening: The evolution of echolocation in bats. Paleobiology 21 (2): 229-242.
Filewood, L.W. 1983. The possible occurrence in New Guinea of the ghost bat ( Macroderma gigas: Chiroptera: Megadermatidae). Australian Mammalogy 6: 35-36.
Fukui, D., Agtsuma, N., and Hill, D.A. 2004. Acoustic identification of eight species of bat (Mammalia: Chiroptera) inhabiting forests of southern Hokkaido, Japan: Potential for conservation monitoring. Zoological Science 21: 947-955.
Gebia, O. and Balun, L. 2004. Species diversity and composition in the Darai limestone karst and Libano alluvial forest in Gulf and Southern Highlands Provinces. KICDP Scientific Report: SR-04-01. Unpublished report to WWF.
Griffiths, T.A., Koopman, K.F. and Starrett, A. 1991. The systematic relationship of Emballonura nigrescens to other species of Emballonura and to Coleura(Chiroptera: Emballonuridae). American Museum Novitates 2996: 1-16.
Grinnell, A.D. and Hagiwara, S. 1972. Adaptations of the auditory nervous system for echolocation. Studies of New Guinea bats. Zeitscrift fur vergleichende Physiologie 76: 41-81.
Guillen, A., Juste, B., and Ibanez, C. 2000. Variation in the frequency of the echolocation calls of Hipposideros ruber in the Gulf of Guinea: an exploration of the adaptive meaning of the constant frequency value in rhinolophid CF bats. Journal of Evolutionary Biology 13: 70-80.
Hall, L.S. and Richards, G. 1995. Diadem Leafnosed-bat Hipposideros diadema. Pp. 459-460 in The Mammals of Australia, Second Edition, edited by R. Strahan. Reed Books, Chatswood, NSW.
Helgen, K. 2005. Systematics of the Pacific monkey-faced bats (Chiroptera: Pteropidae) with a new species of Pteralopex and a new Fijian genus. Systematics and Biodiversity 3 (4): 433-453.
Helgen, K. 2006. A taxonomic and geographic overview of mammals of Papua. Pp. 689-749 in The Ecology of Papua edited by A.J. Marshall and B. M. Beehler. Periplus editions, Singapore.
Heller, K.G. and Von Helversen, O. 1989. Resources partitioning of sonar frequency bands in rhinolophoid bats. Oecologia 80: 178-186.
Holderied, M.W. and von Helversen, O. 2003. Echolocation range and wingbeat period match in aerial-hawking bats. Proceedings of the Royal Society Biological Sciences 270(1530): 2293-2299.
Holderied, M.W., Korine, C., Fenton, M.B., Parsons, S., Robson, S. and Jones, G. 2005. Echolocation call intensity in the aerial hawking bat Eptesicus bottae(Vespertilionidae) studied using stereo videogrammetry. Journal of Experimental Biology 208: 1321-1327.
Jacobs, D.S., Eick, G.N., Schoeman, M.C. and Matthee, C.A. 2006. Cryptic species in an insectivorous bat Scotophilus dinganii. Journal of Mammalogy 87: 161-170.
Jenkins, P.D. and Hill, J.E. 1981. The status of Hipposideros galeritus(Cantor, 1846) and Hipposideros cervinus(Gould 1854) (Chiroptera: Hipposideridae). Bulletin of the British Museum of Natural History, Zoology 41: 279-294.
Jones, G. 1994. Scaling of wingbeat and echolocation pulse emission rates in bats: Why are aerial insectivorous bats so small? Functional Ecology 8(4): 450-457.
Jones, G. and Barlow, K.E. 2004. Cryptic species of echolocating bats. Pp. 345-349 in Echolocation in bats and dolphins, edited by J.A. Thomas, C.F. Moss, and M. Vater. University of Chicago Press, Chicago, Illinois.
Jones, G. and Holderied, M.W. 2007. Bat echolocation calls: adaptation and convergent evolution. Proceedings of the Royal Society Biological Sciences 274(1612): 905-912.
Kazial, K.A., Burnett, S.C. and Masters, W.M. 2001. Individual and group variation in echolocation calls of big brown bats, Eptesicus fuscus(Chiroptera: Vespertilionidae). Journal of Mammalogy 82(2): 339-351.
Kingston, T. and Rossiter, S.J. 2004. Harmonic hopping in Wallacea's bat. Nature 429: 654-657.
Kingston, T., Jones, G., Zubaid, A. and Kunz, T.A. 2000. Resource partitioning in rhinolophoid bats revisited. Oecologia 124: 332-342.
Kingston, T., Lara, M.C., Jones, G., Zubaid, A., Kunz, T.H. and Schneider, C.J. 2001. Acoustic divergence in two cryptic Hipposideros species: a role for social selection? Proceedings of the Royal Society of London, Series B Biological Sciences 268: 1381-1386.
Law, B.S., Reinhold, L. and Pennay, M. 2002. Geographic variation in the echolocation calls of Vespadelus spp. (Vespertilionidae) from New South Wales and Queensland, Australia. Acta Chiroptologica 4 (2): 201-215.
Leary, T. and Mamu, T. 2004. Conserving Papua New Guinea's forest fauna through community planning. Pp. 186-207 in Conservation of Australia's Forest Fauna(Second Edition), edited by D. Lunney. Royal Zoological Society of New South Wales, Mosman, NSW.
Li, G., Liang, B., Wang, Y., Zhao, H., Helgen, K.M., Liangkong, L., Jones, G. and Zhang, S. 2007. Echlocation calls, diet and phylogenetic relationships of Stoliczka's trident bat Aselliscus stoliczkanus(Hipposideridae). Journal of Mammalogy 88(3):736-744.
Macias, S. and Mora, E.C. 2003. Variation of echolocation calls of Pteronotus quadridens(Chiroptera: Mormoopidae) in Cuba. Journal of Mammalogy 84(4):1428-1436.
Macias, S., Mora, E.C., Koch, C. and Von Helversen, O. 2005. Echolocation behaviour of Phyllops falcatus(Chiroptera: Phyllostomidae): unusual frequency range of the first harmonic. Acta Chiropterologica 7(2): 275-283.
Matsumura, S. 1979. Mother-infant communication in a horseshoe bat ( Rhinolophus ferrumequinum nippon): development of vocalization. Journal of Mammalogy 60(1): 76-84.
McKenzie, N.L. 1995. Beccari's Freetail-Bat Mormopterus beccarii. Pp. 480-481 in The Mammals of Australia(Second Edition) edited by R. Strahan. Reed Books, Chatswood, NSW.
McKenzie, N.L. and Bullen, R.D. 2008. Beccari's Free-tailed Bat Mormopterus beccarii. Pp. 486-487 in The Mammals of Australia(Third Edition) edited by S. Van Dyck and R. Strahan. Reed New Holland Books, Chatswood, NSW.
McKenzie, N.L., Start, A.N. and Bullen, R.D. 2002. Foraging ecology and organisation of a desert bat fauna. Australian Journal of Zoology 50 (5): 529-545.
Milne, D.J. 2002 Key to the Bat Calls of the Top End of the Northern Territory. Parks and Wildlife Commission of the Northern Territory. Technical Report 71.
Novick, A. 1958. Orientation in palaeotropical bats. I. Microchiroptera. Journal of Experimental Biology 138: 81-153.
Novick, A. 1977. Acoustic orientation. Pp. 74-289 in Biology of Bats-Volume III edited by W.A. Wimsatt. Academic Press, New York.
O'Farrell, M.J. and Miller, B.W. 1999. Use of vocal signatures for the inventory of free-flying Neotropical bats. Biotropica. 31(3): 507-516.
O'Farrell, M., Miller, B.W. and Gannon, W.L. 1999. Qualitative identification of free-flying bats using the Anabat detector. Journal of Mammalogy 80(1): 11-23.
Obrist, M., Aldridge, H.D.J.N. and Fenton, M.B. 1989. Roosting and echolocation behavior of the African bat, Chalinolobus variegatus. Journal of Mammalogy 70(4): 828-833.
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.
Paijmans, K. 1976 New Guinea Vegetation. Australian National University Press, Canberra.
Papadatou, E., Butlin, R.K. and Altringham, J.D. 2008. Identification of bat species in Greece from their echolocation calls. Acta Chiropterologica 10(1): 127-143.
Parsons, S. 1998. The effect of recording situation on the echolocation calls of the New Zealand lesser short-tailed bat ( Mystacina tuberculata Gray). New Zealand Journal of Zoology 25: 147-156.
Parsons, S., Boonman, A.J. and Obrist, M.K. 2000. Advantages and disadvantages of techniques for transforming and analyzing chiropteran echolocation calls. Journal of Mammalogy 81(4): 927-938.
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