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Passive Acoustic Monitoring of Tothed Whales, with Implications for Mitigation, Monitoring and Biology

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Passive Acoustic Monitoring of Tothed Whales, with Implications for Mitigation, Monitoring and Biology PASSIVE ACOUSTIC MONITORING OF TOOTHED WHALES, WITH IMPLICATIONS FOR MITIGATION, MANAGEMENT AND BIOLOGY Ph.d. thesis 2010 Line Anker Kyhn NATIONAL ENVIRONMENTAL RESEARCH INSTITUTE AU AARHUS UNIVERSITY PASSIVE ACOUSTIC MONITORING OF TOOTHED WHALES, WITH IMPLICATIONS FOR MITIGATION, MANAGEMENT AND BIOLOGY Ph.d. thesis 2010 Line Anker Kyhn NATIONAL ENVIRONMENTAL RESEARCH INSTITUTE AU AARHUS UNIVERSITY Data sheet Title: Passive acoustic monitoring of toothed whales, with implications for mitigation, management and biology Subtitle: Ph.d. thesis Author: Line Anker Kuhn Affi liation: Dep. of Arctic Environment, National Environmental Research Institute, Aarhus University Publisher: National Environmental Research Institute (NERI)© Aarhus University – Denmark URL: http://www.neri.dk Date of publication: November 2010 Please cite as: Kyhn, L.A. 2010: Passive acoustic monitoring of toothed whales, with implications for mitigation, management and biology. PhD thesis. Dep. of Arctic Environment, NERI. National Environmental Research Institute, Aar- hus University, Denmark. 166 pp. Reproduction permitted provided the source is explicitly acknowledged. Abstract: Small cetaceans like harbour porpoises are notoriously diffi cult to study due to various technical, physical and economical constraints in observing them in their marine environment. However, toothed whales are vocal animals and their social life as well as successful orientation and feeding depends on emission and reception of sound. When humans, by means of a hydrophone, are the receivers of these sounds, analysis may provide important information to researchers about the behavioural ecology, social interactions and sensory physiology of toothed whales, as well as it may be the means for acoustic monitoring. This ph.d. thesis presents a novel approach for counting and estimating density of harbour porpoises using acoustic dataloggers. This method is especially valuable for low density areas where traditional visual surveys become very expensive per observa- tion. The thesis further presents thorough descriptions of sounds from six small species that all use the same echolocation signal type, a narrow band high frequency (NBHF) click. Such thorough sound descriptions are a prerequisite of acoustic monitoring. The NBHF click is further discussed in light of possible adaptive values, since the click has evolved by convergent evolution four times. In agreement with previously stated hypothesis it is concluded through modelling that the NBHF click type most likely evolved as acoustic camoufl age against predation from killer whales, since all energy in the NBHF click is at frequencies above the upper hearing limit of killer whales. Keywords: Acoustic monitoring, abundance, density, detection function, harbour porpoise, echolocation, click source parameters, habitat specialization, acoustic species differentiation. Supervisors: Dr. Hans Malte, Biological Sciences, Aarhus University Dr. Jakob Tougaard, National Environmental Research Institute, Aarhus University Layout & photos: Line Anker Kyhn ISBN (DMU): 978-87-7073-210-9 Printed by: Rosendahls – Schultz Grafi sk a/s Number of pages: 166 Circulation: 50 Internet version: The report is available in electronic format (pdf) at NERI’s website http://www.dmu.dk/Pub/PHD_LAK.pdf Content Preface 5 Dansk resumé 6 Summary 8 Acknowledgements 10 A. Review Chapter I 11 Kyhn, L.A. Passive acoustic monitoring of Narrow Band High Frequency species. A review. (Unpublished manuscript). Appendix Source parameter definitions 77 B. Passive Acoustic Monitoring Chapter II 79 Kyhn, L.A., Tougaard, J., Thomas, L., Duve, L.R., Steinback, J., Amundin, M., Desportes, G. and Teilmann, J. From echolocation clicks to animal density – acoustic sampling of harbour porpoises with static dataloggers. (Drafted for Applied Ecology). C. Narrow Band High Frequency Sounds Chapter III 103 Kyhn, L.A., Tougaard, J., Jensen, F., Wahlberg, M., Stone, G., Yoshinaga, A., Beedholm, K. and Madsen, P.T. (2009). Feeding at a high pitch: Source parameters of narrow band, high frequency clicks from echolocating off-shore hourglass dolphins and coastal Hector’s dolphins. J. Acoust. Soc. Am. 125(3): 1783–1791. Chapter IV 115 Kyhn, L.A., Jensen, F. H., Beedholm, K., Tougaard, J. Hansen, M. and Madsen, P.T. (2010). Echolocation in sympatric Peale’s dolphins (Lagenorhynchus australis) and Commerson’s dolphins (Cephalorhynchus commersonii) producing narrow-band high-frequency clicks. The Journal of Experimental Biology 213: 1940-1949. Chapter V 127 Tougaard, J. and Kyhn, L.A. (2010). Echolocation sounds of hourglass dolphins (Lagenorhynchus cruciger) are similar to the narrow band high-frequency echolocation sounds of the dolphin genus Cephalorhynchus. Marine Mammal Science 26(1): 239-245. Chapter VI 137 Kyhn, L.A., Tougaard, J., Beedholm, K., Ash, E., Williams, R. Jensen, F.H. and Madsen, P.T. (Manuscript). Clicking in killer country: Is evolution of echolocation click parameters driven by predation or habitat? (Drafted for Behavioural Ecology and Sociobiology). Ph.d. thesis for the partial fulfilment of the requirements for the degree of Doctor of Philosophy (ph.d. thesis). Submitted to the Faculty of Science, Aarhus University, 29th October 2010. Supervisors: Dr. Hans Malte, Biological Sciences, Aarhus University Dr. Jakob Tougaard, National Environmental Research Institute, Aarhus University. _____________________________________________________________________ Defence date: December 8th 2010. Opponents: Dr. Paul Nachtigall, Hawai'i Institute of Marine Biology, Coconut Island, Marine Mammal Research Program Dr. Jonathan Gordon, Sea Mammal Research Unit, University of St. Andrews, Scotland. Assoc. Prof. Angela Fago (Chairman) Zoophysiology, Dept. of Biological Sciences, Aarhus University _____________________________________________________________________ All photos taken by the author Preface This thesis represents the partial fulfilment of the requirements for the degree of Doctor of Philosophy (ph.d.) at the Faculty of Science, Aarhus University. The thesis is written in accordance with regulations outlined by the Faculty of Science, in May 2002, and within the three year frame. When I was a child I once stumbled upon a dead harbour porpoise on the beach. I was fascinated and astonished – could that really be a whale? The picture has stuck to my mind and the memory of the amazement is clear. But what astonishes me to day is, that there still is so much to be learned about cetaceans: Imagine that in 2007 Jakob Tougaard and I were the first to record sounds from hourglass dolphins! And there are still species that have never been recorded. That more knowledge is ever important is underlined by the fact that one cetacean species have died out since I was a child. As a scientist it is crucial to obtain knowledge by posing and testing hypothesis scientifically in order to objectively assess the problems that many cetacean species encounter to day from for example human encroachment of their habitats. My hope for this thesis work is that I may contribute with knowledge on a cheap methodology to study and count small cetaceans by acoustic monitoring in all parts of the world, and I hope that this may contribute some of the required knowledge to ensure that we shall experience no more extinctions of cetaceans. This thesis deals with the sounds of six narrow band high frequency species that I have recorded in different parts of the world during the last three years. In this thesis I also present a method to combine acoustics and monitoring with the aim of being able to assess population densities acoustically. The thesis consists of six manuscripts of which three are published in peer reviewed journals (III, IV & V), two are ready for submission (II & VI) and awaits fruitful comments from my opponents, and the last manuscript, chapter I, represents, in accordance with the regulations outlined by the Faculty of Science, a larger review where I put my main results of chapter II to VI in to the context of other studies. Copenhagen, October 27th 2010 Line Anker Kyhn 5 Dansk resumé Dette er et resumé af ph.d. afhandlingen “Passive acoustic monitoring of small toothed whales, with implications for mitigation, management and biology” udarbejdet af Line Anker Kyhn på Afdelingen for Arktisk Miljø, Danmarks Miljøundersøgelser, & Zoofysiologi, Biologisk Institut, begge Århus Universitet. Jeg har arbejdet med en særlig gruppe af små tandhvaler der alle producerer samme ekkolokaliseringssignal. Signalet – klikket - er særligt bemærkelsesværdigt, fordi al energien er indeholdt i et smalt frekvensbånd, med energi udelukkende over hundrede kilohertz (kHz). Klikket kaldes et NBHF klik (Narrow Band High Frequency). Klikkets egenskaber gør det let, at designe et filter, der kan sortere signalet fra den omgivende støj. Der er derfor udviklet dataloggere der automatisk kan opsamle kliks fra eksempelvis marsvin (Phocoena phocoena, L.). Metoden anvendes i Danmark til at beskrive relative ændringer i forekomsten af marsvin over tid. Det er dog langt at foretrække, at kunne vurdere ændringer på populationsniveau. Jeg har i denne afhandling arbejdet med, at udvikle en metode til at tælle marsvin ved brug af akustiske dataloggere. Den metode præsenterer jeg i kapitel II. For at omregne akustiske registreringer per tidsenhed til antal dyr per arealenhed kræver det, at man kan beskrive en dataloggers detektionsfunktion. En detektionsfunktion beskriver hvad sandsynligheden er for, at en datalogger registrerer et klik
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