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Globicephala Melas) Pulsed Calls and Complex Whistles

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Globicephala Melas) Pulsed Calls and Complex Whistles STRUCTURAL VARIATION AND COMMUNICATIVE FUNCTIONS OF LONG-FINNED PILOT WHALE (GLOBICEPHALA MELAS) PULSED CALLS AND COMPLEX WHISTLES by Leah Nemiroff Submitted in partial fulfillment of the requirements for the degree of Master of Science at Dalhousie University Halifax, Nova Scotia January 2009 © Copyright by Leah Nemiroff, 2009 DALHOUSIE UNIVERSITY DEPARTMENT OF BIOLOGY The undersigned hereby certify that they have read and recommend to the Faculty of Graduate Studies for acceptance a thesis entitled “STRUCTURAL VARIATION AND COMMUNICATIVE FUNCTIONS OF LONG-FINNED PILOT WHALE (GLOBICEPHALA MELAS) PULSED CALLS AND COMPLEX WHISTLES” by Leah Nemiroff in partial fulfillment of the requirements for the degree of Master of Science. Dated: 28 January, 2009 Supervisor: _________________________________ Readers: _________________________________ _________________________________ _________________________________ ii DALHOUSIE UNIVERSITY DATE: January 28, 2009 AUTHOR: Leah Nemiroff TITLE: STRUCTURAL VARIATION AND COMMUNICATIVE FUNCTIONS OF LONG-FINNED PILOT WHALE (GLOBICEPHALA MELAS) PULSED CALLS AND COMPLEX WHISTLES DEPARTMENT: Department of Biology DEGREE: MSc CONVOCATION: May YEAR: 2009 Permission is herewith granted to Dalhousie University to circulate and to have copied for non-commercial purposes, at its discretion, the above title upon the request of individuals or institutions. _______________________________ Signature of Author The author reserves other publication rights, and neither the thesis nor extensive extracts from it may be printed or otherwise reproduced without the author’s written permission. The author attests that permission has been obtained for the use of any copyrighted material appearing in the thesis (other than the brief excerpts requiring only proper acknowledgement in scholarly writing), and that all such use is clearly acknowledged. iii To Sach, for singing to me about lemons. And to my family, who always believed that I could be a marine biologist, even though I’m scared of fish. iv Table of Contents Table of Contents.................................................................................................................v List of Tables....................................................................................................................viii List of Figures......................................................................................................................x Abstract..............................................................................................................................xv List of Abbreviations Used...............................................................................................xvi Acknowledgements........................................................................................................xviii CHAPTER 1 – INTRODUCTION.................................................................................. 1 1.1 General Introduction ................................................................................................. 1 1.2 The Long-Finned Pilot Whale .................................................................................. 3 1.2.1 Distribution and Abundance ................................................................................ 3 1.2.2 Movement Patterns .............................................................................................. 4 1.2.3 Individual Identification....................................................................................... 4 1.2.4 Social Structure.................................................................................................... 4 1.3 Cetacean Vocalization .............................................................................................. 5 1.3.1 Cetacean Sound Production and Hearing ............................................................ 5 1.3.2 Odontocete Vocalizations .................................................................................... 7 1.3.3 Long-Finned Pilot Whale Vocalizations.............................................................. 8 1.4 Thesis Overview ..................................................................................................... 10 CHAPTER 2 – PILOT WHALE PULSED CALLS .................................................... 11 2.1 Introduction............................................................................................................. 11 2.2 Methods................................................................................................................... 13 2.2.1 Field Sites........................................................................................................... 13 2.2.2 Recording of Vocalizations................................................................................ 14 2.2.3 Acoustic Analysis .............................................................................................. 14 2.2.4 Statistical Analysis............................................................................................. 16 2.3 Results..................................................................................................................... 18 2.3.1 Properties of Pilot Whale Pulsed Calls .............................................................. 18 2.3.2 Principal Components Analysis......................................................................... 23 2.3.3 K-means Cluster Analysis.................................................................................. 27 2.3.3 Hierarchical Cluster Analysis ............................................................................ 27 2.4 Discussion............................................................................................................... 31 2.4.1 The Structural Complexity of Pulsed Calls ....................................................... 31 2.4.2 The Non-Discrete Nature of Pulsed Calls.......................................................... 34 2.4.3 Pilot and Killer Whale Pulsed Calls, an Example of Form Following Function?............................................................................................................ 36 2.5 Summary................................................................................................................. 39 CHAPTER 3 – RATING THE SIMILARITY OF PILOT WHALE CALLS USING STATISTICAL AND PERCEPTUAL MEASURES.......... 40 3.1 Introduction............................................................................................................. 40 v 3.2 Methods................................................................................................................... 42 3.2.1 Field Sites........................................................................................................... 42 3.2.2 Recording of Vocalizations................................................................................ 42 3.2.3 Call Extraction ................................................................................................... 42 3.2.4 Statistical Measures ........................................................................................... 43 3.2.4.1 Acoustic Analysis...............................................................................................43 3.2.4.2 Statistical Analysis .............................................................................................44 3.2.5 Perceptual Measures .......................................................................................... 44 3.2.5.1 Online Comparison of Calls...............................................................................44 3.2.5.2 Statistical Analysis .............................................................................................48 3.2.6 Statistical Measures vs. Perceptual Measures.................................................... 50 3.3 Results..................................................................................................................... 51 3.3.1 Similarities from Quantitative Statistical Measures .......................................... 51 3.3.2 Similarities from Qualitative Perceptual Measures ........................................... 51 3.3.3 Statistical Measures vs. Perceptual Measures.................................................... 56 3.4 Discussion............................................................................................................... 57 3.4.1 The Perceptual Method...................................................................................... 57 3.4.2 Human Perception of Pilot Whale Calls ............................................................ 59 3.4.3 The Statistical Method ....................................................................................... 60 3.4.4 Comparing Perceptual Measures with Statistical Measures .............................. 60 3.5 Summary................................................................................................................. 61 CHAPTER 4 – SOURCES OF VARIATION AND COMMUNICATIVE FUNCTIONS OF PILOT WHALE PULSED CALLS AND COMPLEX WHISTLES..................................................................... 63 4.1 Introduction............................................................................................................. 63 4.2 Methods................................................................................................................... 65 4.2.1 Field Methods ...................................................................................................
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