Acoustic Ecology of Humpback Whales (Megaptera Novaeangliae) in the Hawaiian Archipelago
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ACOUSTIC ECOLOGY OF HUMPBACK WHALES (MEGAPTERA NOVAEANGLIAE) IN THE HAWAIIAN ARCHIPELAGO A DISSERTATION SUBMITTED TO THE OFFICE OF GRADUATE EDUCATION OF THE UNIVERSITY OF HAWAIʻI AT MĀNOA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY (MARINE BIOLOGY) MAY 2017 By Jessica Chen Dissertation Committee: Whitlow W.L. Au, Chairperson Marc O. Lammers Paul E. Nachtigall Adam A. Pack Eva-Marie Nosal © Copyright 2017 Jessica Chen All rights reserved. i ACKNOWLEDGEMENTS I would like to thank my advisor, Whitlow Au, for all his support and supervision since my entrance into the graduate program and throughout my doctorate research. It takes a lot of patience to guide a biologist through the field of acoustics, and his assistance was priceless. I would like to thank my dissertation committee, Paul Nachtigall, Adam Pack, Marc Lammers, and Eva-Marie Nosal for their valuable guidance and support. I have learned so much from each of you and could not have made it this far without your advice and cooperation. My research involved many parts and people, so I wish to thank all who contributed to the research. Many thanks to my colleagues, past and present, in the Marine Mammal Research Program (MMRP) of the Hawaiʻi Institute of Marine Biology (HIMB) for their guidance and support: Alison Stimpert, Adrienne Copeland, Lee Shannon, Giacomo Giorli, Aude Pacini, Michael Richlen, Alexis Rudd, Adam Smith, Ali Bayliss, Aliza Milette-Winfree, and Helen Ou. Additional thanks to “honorary labmates” Brendan Rideout, Andy Dewald, and Anne Rosinski. I would also like to thank the many volunteers who came out to assist with fieldwork and data collection: Andrea Bendlin, Chelsie Counsell, Elia Herman, Merra Howe, Zachary Jambor, Anke Kuegler, Sheldon Plentovich, Mark Royer, Sabena Siddiqui, Hilary Street, Chelsea Szydlowski, Allison Taranto, Grant Thompson, Dana Wilkes, and Eden Zhang. Thanks to Adrienne Copeland and Melanie Abecassis for help with statistical analysis and R scripts. If I have missed anyone, know that you have my gratitude for your help throughout the years. I could not have carried out my research without the partnership of the Hawaiian Islands Humpback Whale National Marine Sanctuary and Ed Lyman. Their logistical and vessel support was essential for the success of my research. The exemplary captaining skills of Ed Lyman, Joe Carrier, and Carmen DeFazio were priceless. Thanks also to Oceanwide Science Institute and Marc Lammers for vessel support. Bill Burgess always did his best to provide troubleshooting and data analysis support for the Acousonde and Bioacoustic probe tags. I also want to thank my family and friends for their continued support throughout the years. Their love and encouragement allowed me to persevere through the many academic and research challenges, and complete my PhD. ii This research was supported by the University of Hawaiʻi (UH) Sea Grant College Program, the UH School of Ocean and Earth Science and Technology, the NOAA Coral Reef Conservation Program, the NOAA/HIMB Northwestern Hawaiian Islands Partnership Program, NOAA’s Pacific Islands Fisheries Science Center - Coral Reef Ecosystem Division, the Hawaiian Islands Humpback Whale National Marine Sanctuary, and the Office of Naval Research. Additional funding was provided by the University of Hawaiʻi E. A. Kay Award. Research was conducted under NOAA National Marine Fisheries Service scientific research permit numbers 14682, 14682-01, and 14585, State of Hawaiʻi Special Activity Permit numbers 2014-02, 2014-10, PMAL-2015-204, and PMAL-2017-206, and University of Hawaiʻi Institutional Animal Care and Use Committee protocol numbers 13-1769, 13-1769-2, 13-1769-3. iii ABSTRACT Humpback whale songs are some of the most studied of cetacean vocalizations, however some of their other non-song vocalizations are less researched. During the winter breeding season, humpback whales are a consistent source of sound in the waters surrounding Hawaii. This dissertation research focused on quantifying and describing both song and non-song vocalizations at various spatial and temporal scales. Song units from the Main and Northwestern Hawaiian Islands were identified and quantified. The pattern in song unit occurrence suggested that the songs varied geographically but there is no clear divide between the humpback songs from Main and Northwestern Hawaiian Islands, showing instead a gradient of change along the island chain. Sound levels were examined across time of day and over the course of a year for patterns of sound contribution from humpback whales around the island of Kauai. Sound levels were approximately 6 dB higher during the winter season compared to summer, when possible sound sources include humpback whales, wind, and waves. Sound levels were approximately 1 dB higher during the day compared to night during the humpback season. A calf off the coast of Maui was tagged with an Acousonde acoustic and movement recording tag. The tag recorded song from a singing escort and revealed that a calf may be exposed to sound levels from 126 to 158 dB re 1 µPa. Additional mother-calf groups were tagged in the Maui area to study the vocalizations and associated behaviors. Twelve call types were identified from tags deployed on mothers and calves, and correlated with travel and surface activity. These studies are important to understanding the way humpback whales communicate and the vocalizations are used in order to understand their use of the environment. As the levels of anthropogenic noise in the ocean increase, this knowledge is important for preserving the acoustic environment of the humpback whales. iv TABLE OF CONTENTS Acknowledgements ................................................................................................................. ii Abstract ................................................................................................................................... iv List of tables .......................................................................................................................... vii List of figures ........................................................................................................................ viii List of abbreviations and symbols ......................................................................................... x Chapter 1. Introduction ......................................................................................................... 1 1.1 Acoustics as a tool to study wildlife .................................................................................... 1 1.2 Humpback whales ............................................................................................................... 3 1.3 Humpback whale acoustics ................................................................................................. 6 1.4 Research objectives ............................................................................................................. 8 1.5 Research significance ......................................................................................................... 8 Chapter 2. Variation in the song units utilized by humpback whales wintering in the Northwestern and Main Hawaiian Islands ......................................................................... 10 2.1 Abstract ............................................................................................................................. 10 2.2 Introduction....................................................................................................................... 11 2.3 Methods ............................................................................................................................. 14 2.4 Results ............................................................................................................................... 17 2.5 Discussion ......................................................................................................................... 21 Chapter 3. Spatial and temporal changes in soundscape around the island of Kauai, Hawaii ................................................................................................................................................ 24 3.1 Abstract ............................................................................................................................. 24 3.2 Introduction....................................................................................................................... 25 3.3 Methods ............................................................................................................................. 27 3.4 Results ............................................................................................................................... 30 3.5 Discussion ......................................................................................................................... 44 Chapter 4. First measurements of the received levels of humpback whale song produced by a singing escort in close proximity to a calf in the Hawaiian breeding grounds ............. 48 4.1 Abstract ............................................................................................................................. 48 4.2 Introduction....................................................................................................................... 49 4.3 Methods ............................................................................................................................